Handbook of Muscle Variations and Anomalies in Humans - A - Eve K. Boyle, Vondel S. E. Mahon, Rui Diogo - 2022 - CRC Press - 9780367538620 - Anna's Archive

Handbook of Muscle Variations and
Anomalies in Humans
Handbook of Muscle Variations and
Anomalies in Humans
A Compendium for Medical Education, Physicians,
Surgeons, Anthropologists, Anatomists, and Biologists
Eve K. Boyle
Department of Anatomy
Howard University College of Medicine
Washington, District of Columbia
Vondel S. E. Mahon
R Adams Cowley Shock Trauma Center
University of Maryland Medical Center
Baltimore, Maryland
Rui Diogo
Department of Anatomy
and
Center for the Advanced Study of Human Paleobiology
The George Washington University
First edition published 2022
by CRC Press
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ISBN: 978-0-367-53862-0 (hbk)

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DOI: 10.1201/9781003083535
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Contents
Preface..................................................................................................................................................................................vii
Acknowledgments.................................................................................................................................................................ix
About the Authors.................................................................................................................................................................xi
Contributors ....................................................................................................................................................................... xiii
Chapter 1 Introduction ...................................................................................................................................................... 1

Eve K. Boyle, Vondel S. E. Mahon, and Rui Diogo
Chapter 2 Head and Neck Muscles ...................................................................................................................................9

Eve K. Boyle, Vondel S. E. Mahon, Rui Diogo, Warrenkevin Henderson, Hannah Jacobson,
Noelle Purcell, and Kylar Wiltz
Chapter 3 Upper Limb Muscles .................................................................................................................................... 111

Eve K. Boyle, Vondel S. E. Mahon, and Rui Diogo
Chapter 4 Trunk Muscles..............................................................................................................................................207

Eve K. Boyle, Vondel S. E. Mahon, Rui Diogo, and Rowan Sherwood
Chapter 5 Lower Limb Muscles.................................................................................................................................... 253

Eve K. Boyle, Vondel S. E. Mahon, Rui Diogo, and Malynda Williams
References ......................................................................................................................................................................... 341

Index .................................................................................................................................................................................. 375
v
Preface
Most textbooks and atlases of human anatomy describe in many species of animals—and a comprehensive review
only a few cases of variations in the “normal” population of the literature. This book combines three important goals:
and/or of anomalies of the muscles. These variations and (1) provide a comparative context, based on evolutionary
anomalies are also often excluded from the curricula in col- thinking and comparative zoological anatomy, for the exis-
leges of medicine and dentistry in the U.S.A. and in many tence of variations and defects in humans, particularly for
other countries. However, it is known that each person medical students, professors, and students of human gross
within the “normal” population has at least a few muscle anatomy; (2) summarize the major types of variations and
variations, and there are millions of individuals born glob- anomalies found in the skeletal muscles of humans; and (3)
ally every year with muscle anomalies. It is highly likely include schematic fgures for an easy visualization of these
that during their careers, almost all medical profession- variations and anomalies. This approach dovetails with
als working with any aspect related to muscles—includ- classical and contemporary approaches to medical instruc-
ing muscle physiology, gross anatomy, physiotherapy, and tion and clinical practice.
surgery—will be faced with at least a few cases of muscle This book may be of interest to students of medical and
variations and/or anomalies. There are therefore crucial other health sciences—such as dentistry, physiotherapy, and
gaps of knowledge between what teachers of medical and orthopedics—instructors, teaching assistants, professors,
dentistry schools teach, what students learn, what textbooks and other medical researchers and pathologists; students
and atlases show, and the reality that medical professionals and teachers at nonmedical colleges, universities, and high
will face in practice during their careers. schools where human gross anatomy is also taught; and sci-
This is the frst textbook and anatomical atlas to be exclu- entists and research students in biological felds including
sively focused on the variations and anomalies of human biological anthropology, comparative anatomy, functional
muscles. It is based on the authors’ own work—our lab has morphology, zoology, and evolutionary and developmental
been particularly focused on human variations and defects biology.
vii
Acknowledgments
We would all like to thank Boyani Moikobu, Latonya Vondel S. E. Mahon would like to thank God, Lovon
Aaron, Tah-jai Sharpe, Rowan Sherwood, and Brandon Mahon, Nancy Welsh, Mary Klunk, Izel Glover, Ann Brown,
Wallace for their assistance in formatting references. We all Janet Carlile, Dan Thompson, the University of Maryland
also want to thank Chuck Crumly for always thinking a step Medical Center R. Adams Cowley Shock Trauma Center, the
ahead and inviting us to do this book for Taylor & Francis, Johns Hopkins University School of Medicine Department
and for helping us defne key aspects about the organization of Art as Applied to Medicine, the CUNY Baccalaureate for
of the book. Many thanks also to Ana Lucia Eberhart. Unique and Interdisciplinary Studies, the Thomas W. Smith
Eve K. Boyle would like to thank the many mentors and Academic Fellowship, The Art Students League of New York,
academic advisors who have facilitated her understand- the Medgar Evers College of the City University of New York,
ing of musculoskeletal anatomy and provided opportuni- the AMI, and his beloved family, mentors, and friends.
ties to teach and conduct research. Special thanks to Drs. Rui Diogo would like to thank the numerous colleagues
Kirsten Brown, Jeremy DeSilva, Sergio Almécija, Bernard that have worked with him on the subjects covered in this
Wood, Briana Pobiner, and Shiloh Jones. Thank you also to book, as well as to the institutions that have funded those
the students, staff, and faculty of the George Washington works, and to Howard University, where he teaches anat-
University School of Medicine and Health Sciences and the omy and undertakes research about a plethora of anatomi-
Georgetown University School of Medicine. cal, evolutionary, and societal topics.
ix
About the Authors
Eve K. Boyle is an evolutionary anthropologist with exper- healthcare and medical education. He is also a proud mem-
tise in informal science education, public engagement, proj- ber of the Association of Medical Illustrators (AMI).
ect management, and broadening participation initiatives.
She received her Ph.D. from the Center for the Advanced Rui Diogo is a multi-awarded researcher, speaker, and
Study of Human Paleobiology of George Washington writer who is renowned worldwide for addressing broader
University in 2019. As a postdoctoral fellow at Howard scientifc questions and societal issues using state-of-the-
University from 2019 to 2020, she was the frst project art empirical data from many different felds of science.
manager for the Visible Ape Project. She served as an He obtained his Bachelor’s degree in Biology from the
AAAS Science & Technology Policy Fellow at the National University of Aveiro, Portugal, and later did a Ph.D. at
Science Foundation from 2020 to 2022. the University of Liege, Belgium, a postdoc at the King's
College of London, and then a master’s and a Ph.D. at the
Vondel S. E. Mahon is a Medical Illustrator at Mahon Department of Biology of George Washington University,
Illustrations LLC and at a private enterprise. He received U.S.A. A wonderer and a wanderer, he has done research,
his M.A. in Medical and Biological Illustration from the gave speeches, or traveled in more than 120 countries. He
Department of Art as Applied to Medicine at the Johns is the author of more than 150 papers in top journals and
Hopkins University School of Medicine. For his master’s 20 books, including Learning and Understanding Human
thesis, he developed a novel tool for orthopedic trauma edu- Anatomy and Pathology—used at several medical schools
cation with the Johns Hopkins University Computer Aided worldwide; Evolution Driven by Organismal Behavior—
Medical Procedures Laboratory and the Johns Hopkins often listed among the ten best evolutionary books in 2017;
Medicine Technology Innovation Center. He holds an and the already highly-acclaimed Meaning of Life, Human
interest in the use of Extended Reality (XR) technology in Nature, and Delusions.
xi
Contributors
Warrenkevin Henderson Rowan Sherwood
Boston University Department of Anthropology
Boston, Massachusetts University of Michigan
Ann Arbor, Michigan
Hannah Jacobson
George Washington University Kylar Wiltz
Washington, District of Columbia Howard University College of Medicine
Noelle Purcell
George Washington University Malynda Williams
Washington, District of Columbia Howard University College of Medicine
xiii
1 Introduction
Eve K. Boyle
Vondel S. E. Mahon
Rui Diogo
CONTENTS
Why Do Humans Have Anatomical Variations and Anomalies? ........................................................................................... 1
Toward a More Holistic Approach to Human Anatomy and Evolution ................................................................................. 4
Brief Notes about Our Methodology .....................................................................................................................................6
Portions of this introduction are reproduced by permission other species. For instance, one of the more severe malfor-
from John Wiley and Sons: Diogo, R., Smith, C.M. and mations in humans is cyclopia, the presence of a single eye.
Ziermann, J.M. 2015. Evolutionary developmental pathol- We never had an ancestor that was truly a cyclops, but many
ogy and anthropology: A new feld linking development, other mammals also have cyclopic malformations.
comparative anatomy, human evolution, morphological This phenomenon is now known as Alberch’s (1989)
variations and defects, and medicine. Dev Dyn 244:1357– ill-named “logic of monsters.” According to this theory,
1374. © 2015 Wiley Periodicals, Inc. which has been strongly supported by a plethora of empiri-
cal data, there is a parallel between the variation/anomalies
in normal/abnormal individuals of a certain taxon and the
WHY DO HUMANS HAVE ANATOMICAL
fxed diversity observed in wildtype individuals of other
VARIATIONS AND ANOMALIES? taxa. According to Alberch (1989), this parallel might be
When Charles Darwin wanted to convince a highly skep- achieved through regulation of a conserved developmental
tical scientifc community and general public that we program (i.e., a set of genetic and/or epigenetic interactions)
evolved from other primates in The Descent of Man (1871), such that the structure of these internal interactions con-
he started by discussing human anatomical variations and strains the realm of possible variation upon which selection
anomalies. Why? Because he knew that a strong way to can operate. This program may lead to the breakdown in the
show that we descend from other animals is to highlight evolution of some clades, but within most clades this would
the commonalities we share with them and particularly, the lead to death of the embryos. This leads us to distinguish
commonalities that are only present in some humans, as a between variant features and anomalies. By defnition,
reminder of the nonhuman ancestors we had several mil- anatomical variants are structures present in the common
lions of years ago. Our anatomical variations and anoma- population, that is, the population that is karyotypically
lies, in particular those that are related to the presence of normal and that does not have severe congenital malforma-
evolutionary reversions, are indeed the most direct, strong tions, or so-called “birth defects.” Anatomical anomalies
evidence of our evolutionary past. This is because there are are precisely often the result of genetic mutations or severe
features that are present in adults of other species that are congenital malformations, such as cyclopia, a feature that is
typically absent in humans, but for some reason can still never found in the common population. However, this does
be found in some human adults. The most likely scientifc not mean that anatomical anomalies are always more severe
explanation, Darwin argued, is that such features were than variants. For instance, a certain limb muscle miss-
present in our adult ancestors and then were evolutionary ing in an individual from the common population would
lost but evolutionarily “re-acquired.” Not only that: even for be seen as a variant, while the very same feature seen in,
some anatomical features that appear in humans that have let’s say, a person with Down syndrome, would be seen as
nothing to do with traits typically present in other animals, an anatomical anomaly. According to the “logic of mon-
these usually also show that we share evolutionary and sters,” this parallel between the variation and anomalies in
developmental commonalities with them because such vari- so-called “normal” vs “abnormal” individuals of a certain
ations and anomalies are also present in some cases within taxon is exactly to be often expected.
DOI: 10.1201/9781003083535-1 1
2 Handbook of Muscle Variations and Anomalies in Humans
As noted by Diogo et al. (2015), such a parallel between result from similar developmental mechanisms (Hodin
the more common phenotypic variations seen in the normal 2000). The “logic of monsters” is thus framed in an “inter-
human population and malformations seen in birth defects nalist” view of evolution and development that contrasts
is also to be expected according to Shapiro et al.’s (1983) with the more “externalist” view of adaptationists, who
“lack of homeostasis” model. This model was in large part maintain that selection by the external environment is the
formulated based on observations of human trisomic indi- main evolutionary force. For example, frogs and salaman-
viduals and states that the presence of a whole extra func- ders tend to exhibit loss of digit one and reduction of digit
tioning chromosome or of a large chromosome segment fve due to developmental constraints. This pattern is found
causes a disruption of evolved genetic balance. Because of in frogs living in different environments that are exposed to
the obligatory integration of the entire genotype, this dis- different external factors (Alberch and Gale 1985). That is,
ruption affects the products of the trisomic chromosome and the recurrent loss/reduction of such digits in frogs seems to
other chromosomes. This results in decreased physiological have much more to do with internal mechanisms than with
and developmental buffering against genetic and environ- specifc external conditions, although of course the latter
mental forces, which leads to decreased developmental and might play some role in the specifc frog groups and/or geo-
physiological homeostasis where the pathways and pro- logical time in which those loses/reductions happen.
cesses that will be the most often and seriously affected are The works of Alberch, as well as of his colleague Stephen
those that are more unstable. This instability thus leads to Jay Gould and various other authors in the 1970s and 1980s,
variations in the normal population. An illustrative exam- have led to the rise of evolutionary developmental biology
ple predicted by both the “logic of monsters” and “lack of (Evo-Devo), which has revived interest in comparative anat-
homeostasis” hypotheses is the presentation of palmaris omy, particularly of soft tissues, and of “teratology,” now
longus. The absence of palmaris longus (see Fig 3.14) is a designated as the study of congenital malformations. These
common human polymorphism. Polymorphisms are differ- are felds that were somewhat dormant for many years after
ent from variations, because although they are also present having a prominent role in anatomical and biological stud-
in the “normal” population as variants are, they are present ies in the 18th, 19th, and early 20th centuries, with only a
in more than 2% of the “normal” population, while varia- few notable works in the latter half of the 20th century (e.g.,
tions are present in less than 2%. So, within the “normal” Willis 1958). Following this trend, one of us (RD) has cre-
population there are variants—present in less than 2% of ated, with other colleagues a new subfeld within Evo-Devo:
the population; polymorphisms—present between 2% and Evolutionary Developmental Pathology (Evo-Devo-Path),
50% of the population; and the common phenotype—pres- which is exactly focused on understanding the evolutionary
ent in more than 50% of the population. In the case of the and developmental basis of human variations and anomalies
palmaris longus, this absence of muscle is a polymorphism (Diogo et al. 2015; Diogo and Wood 2016). This new sub-
because the muscle is absent in about ~15%–20% of the feld—which was specifcally applied to human anatomy,
“normal” human population. Interestingly, the absence of by Diogo et al. (2016)—is in a way a comeback to the study
the muscle is often amplifed in humans with severe con- of the links between normal and abnormal development
genital malformations. For example, palmaris longus was and pathologies, which began to be intensively undertaken
reported to be absent in 74% (105) of 141 defective upper centuries ago by authors such as St. Geoffroy St. Hilaire,
limbs reviewed by Smith et al. (2015), further reinforcing but that were then somewhat neglected for decades dur-
that variants and polymorphisms are often amplifed in ing the 20th century. Fortunately, more and more authors
people with congenital malformations. are realizing the importance of investigating these links,
However, with exception to this similarity between the although with Evo-Devo they have been mainly studied
“logic of monsters” and the “lack of homeostasis” mod- by focusing principally on osteological or superfcial fea-
els, they differ fundamentally in their assumptions and tures (e.g., absence of a certain bone, shape of head), with
predictions. The “lack of homeostasis” model argues that fewer studies being done about the muscular system of both
defects are mostly random and disorganized due to a gen- humans and nonhuman animals with anatomical variations
eral lack of homeostasis. The “logic of monsters” predicts, or anomalies. However, now that we understand the devel-
instead, that defects are mostly “logical” and “constrained” opmental—including both genetic and epigenetic—and
because developmental constraints are kept intact by inter- evolutionary basis for why we have anomalies and varia-
nal homeostasis, even when things go “wrong” in both evo- tions, it is time to develop Evo-Devo-Path to the detailed
lution and embryonic development. That is why the “logic study of soft tissues such as muscles.
of monsters” model predicts—contrary to the “lack of For instance, we now know that evolutionary reversions
homeostasis” model—that congenital malformations mir- did play a substantial role in primate/human evolution. In
ror, or amplify, variations, and that both of them further a phylogenetic work about muscle evolution in primates,
mirror evolutionary events that occurred in other taxa at Diogo and Wood (2012b) have shown that 28 out of the 220
completely different geological times. This prediction has (12.7%) evolutionary changes unambiguously optimized in
been supported by studies demonstrating that similar pat- the most accepted primate phylogenetic tree are reversions
terns of intra-specifc diversity within a taxon (plasticity) to a plesiomorphic (ancestral) state. Of those 28 reversions,
and inter-specifc diversity among different taxa usually six were directly related to our own evolution because they
Introduction 3
occurred at nodes that lead to the origin of modern humans. phenotypic patterns often seen in different trisomies, for
Nine of these reversions go against Dollo’s law, which states instance regarding the axial skeleton in fetuses with tri-
that once a complex structure is lost, it is unlikely to be somy 18 vs fetuses with trisomy 21 (Keeling et al. 1997),
regained. Our Evo-Devo-Path studies support the idea that further supporting the ill-named “logic of monsters.” One
reacquisition of morphological structures in adults that have hypothesis we aim to test in future works is that the dis-
been absent for long periods of time is possible because the appearance of certain muscles during early developmental
associated developmental pathways were maintained in the stages of karyotypically normal humans is related to apop-
members of that taxon. tosis, and that the persistence of these atavistic muscles
For example, chimpanzees display a reversion of a later in ontogeny in individuals with trisomies 13, 18, and
synapomorphy of the Hominidae (great apes and modern 21 is associated with delayed development specifcally due
humans) that was acquired at least 15.4 million years ago. to decreased muscle apoptosis. Some authors suggest that
Adult chimpanzees have two contrahentes digitorum mus- humans with DS show an increase of apoptosis in struc-
cles (see Figure 3.18) in addition to adductor pollicis (see tures such as neurons, granulocytes, and lymphocytes
Figure 3.19), which is the only contrahens muscle present in (e.g., Elsayed and Elsayed 2009). If our studies support the
the typical phenotype in adults of other apes. Chimpanzees hypothesis that these individuals have decreased muscle
thus have contrahentes muscles going to digit four and the apoptosis that leads to the presence of additional muscles in
other to digit fve. Developmental studies of hand muscles later ontogenetic stages, this will imply a more nuanced pat-
show that karyotypically normal human embryos do exhibit tern of apoptosis (i.e., a mosaic scenario where apoptosis is
contrahentes going to various fngers, but these muscles increased in some tissues and decreased in others). Within
usually are reabsorbed or incorporated within other struc- the numerous Down syndrome cases listed in the tables of
tures during later embryonic development (Cihak 1972). Dunlap et al. (1986), there are 12 supernumerary muscles,
In karyotypically abnormal humans, such as those with as opposed to two absences. Therefore, Down syndrome
trisomies 13, 18, or 21, the contrahentes often persist after individuals seem to present with more accessory muscles
birth as “atavisms” (see Figure 3.18), which are structures in general, supporting our hypothesis these individuals may
that present as developmental anomalies or variations and have decreased apoptosis in skeletal muscles. Bersu (1980)
resemble the common adult character state of the ancestors also suggested that the persistence of some embryonic mus-
of the taxon to which the individual belongs. Cihak (1972) cles in later stages of development of trisomic individuals
showed that intermetacarpales muscles of the hand are also likely has to do with failure of normal cell death or some
present as discrete muscles in early stages of karyotypically other process of involution. If there is indeed a contrast in
normal human embryonic development but later join with apoptosis between the nervous system (e.g. more apoptosis
fexor breves profundi muscles to form the dorsal interossei of nerve cells) and muscular system (i.e. less apoptosis and
(see Figure 3.18). Therefore, the evolutionary reversions that presence of extra muscles), our hypothesis might also shed
result in the presence of contrahentes and intermetacarpales light on the etiology of hypotonia (low muscle tone) that is
muscles in chimpanzees are likely related to heterochronic present in individuals with Down syndrome.
(specifcally paedomorphic) events in the chimpanzee lin- Some authors suggest that cases in which complex
eage (Diogo and Wood 2012b). So, it is very likely that many structures are formed early in “normal” ontogeny but
variants and anomalies that are present in adult humans and later become lost/indistinct during development (so-called
that were present in our adult ancestors are structures that “hidden variation”), may allow organisms to have greater
were always present in the embryos of our ancestors, but ontogenetic potential early in development. If faced with
that then became absent, or fused with other structures, in external perturbations (e.g., climate change, habitat occu-
later stages of development. But, for some reason, either pied by new species), evolution can use that potential (adap-
genetic or epigenetic, they persist until later stages of devel- tive plasticity: e.g., West-Eberhard 2003). However, authors
opment in certain people as variations or anomalies. That such as Gould (1977, 2002) and Alberch (1989) suggested
is precisely what happens to features such as the presence that these cases support instead a “constrained” (internalist)
of various hand or foot contrahentes muscles, or a platysma rather than an “adaptationist” (externalist) view of evolu-
cervicale muscle of the head (see Figure 2.3). tion. This is because it is not likely that the persistence of
We have been studying cadaveric material from fetal, some muscles in later developmental stages of karyotypi-
neonatal, and adult humans with trisomies and from mouse cally abnormal humans is due to natural selection and adap-
models for Down syndrome (e.g., Ts65Dn) to investigate the tive evolution. This corresponds with the idea defended by
developmental mechanisms related to the atypical devel- Galis and Metz (2007: 415–416): “without denying the evo-
opment of muscles. Research on the skeletons of trisomic lutionary importance of phenotypic plasticity and genetic
individuals has supported the idea that for some features assimilation, we think that for the generation of macro-
there is in fact a developmental delay. For example, some evolutionary novelties the evidence for the impact of hidden
nasal bones develop after the 24th week in cases of trisomy variation is limited” (see also Levinton 2001).
21, while in karyotypically normal humans the develop- Further studies are needed to determine if hidden varia-
mental onset is the 10th week (e.g., Keeling et al. 1997). tion plays a role as important in generating evolutionary
Interestingly, these studies provided examples of different novelties, and they seem to play in the reappearance of
some traits associated with these novelties, as in anatomical embryonic palmaris longus of those humans, and of nonhu-
reversions that violate Dollo’s law. In this sense, it is impor- man primates.
tant to emphasize that “atavistic” muscles of karyotypically Regarding all the muscles of the head and upper and
normal human embryos do not correspond to the muscles of lower limbs as a whole, within the 1540 cases of human
adult primates or of other adult mammals. These muscles muscle congenital malformations that we have compiled
instead correspond to the embryonic muscles of these latter (Smith et al. 2015), 257 (17%) are potential atavisms and
taxa (Diogo and Wood 2012a,b). The developmental path- 352 (23%) can potentially be due to developmental delay.
ways resulting in the presence of these muscles in adults Therefore, if more detailed comparative developmental
were not completely lost in modern humans, even after sev- studies confrm that these numbers, or even just a sub-
eral million years of evolution, likely because these path- stantial portion of them, are truly atavistic and/or due to
ways are recruited in the formation of other structures that developmental delay, these would be signifcant propor-
are present and functional in modern human adults. tions. Interestingly, within those 1540 cases, 650 (42%)
However, it should also be noted there is still some con- concern the presence of abnormal structures (e.g., muscle
fusion about the use of the term “atavism.” Darwin and heads/slips or tendons) and 590 (38%) concern the complete
other 19th century authors considered the presence of a tail- absence, while only 145 (9%) concern fusions and 66 (4%)
like appendage at the height of the coccyx or lumbar spine concern complete duplications of muscles usually present in
in some human newborns to be an “atavism,” a position the normal human population. Despite the substantial pro-
subscribed to by us. Indeed, some authors have described portions of potential atavisms and/or developmental delays
muscles associated with these structures that they suggest mentioned above, those cases are a minority among all
do resemble caudal muscles of other animals (Wiedersheim cases of birth defects/anomalies we have compiled so far.
1895). But it is also clear that many features considered to This supports other research that suggests many other fac-
be “atavisms” in trisomic humans (e.g., Barash et al. 1970; tors are probably involved in leading to anatomical defects
Dunlap et al. 1986) cannot actually be atavisms, because found in these individuals, apart from developmental retar-
those features were never present in our direct ancestors. dation. Such factors can include, for instance, the aberrant
However, some muscle variants and anomalies that we will organization of primordia and misdirected morphogenetic
describe in this book are atavistic by defnition (e.g., dor- movements (Barash et al. 1970).
soepitrochlearis, epitrochleoanconeus, levator claviculae, In a nutshell, it can thus be said that there are likely many
opponens hallucis). What is not so clear is whether their different reasons why humans have anatomical variations
presence is due to a developmental delay, but that is not and anomalies. It is likely that variations are more often
a requirement to consider a feature as an “atavism.” This cases of imprints of our past (Boyle et al. 2020) than are
short discussion highlights the need for more research on anomalies, because, as noted above, variations and anoma-
the ontogeny of these structures in humans and other pri- lies can sometimes refer to the very same feature, but in gen-
mates, which is crucial to clarify these issues. eral the former are not as severe as some of the most severe
According to Hall (1984), the presence of atavistic fea- anomalies. Simply put, people can’t survive and be part of
tures as human variations might represent the maintenance the common population with a cyclopia. Cyclopia is exclu-
of structures that were adult polymorphisms in the past, sively an anomaly, not a thing of our evolutionary past, and
within the normal human population and/or their ancestors. moreover—and likely precisely because of that—it concerns
For instance, as mentioned above, palmaris longus (see only embryos/fetuses in general and never a variation within
Figure 3.14) is present in ~80%–85% of the normal human the common adult human population. It should be noted that
population. Since this muscle is present in about 100% of in this book we are interested in anomalies and variations
the normal population of primates, including most apes, after embryonic development, and therefore we do include
there is apparently a trend toward a decreased frequency of information on fetal musculature, when available. There is
palmaris longus along human evolutionary history (Diogo much we still do not know, both about the specifc develop-
and Wood 2012a,b). If this is indeed the case, it is likely mental mechanisms and about the anatomical patterns, of
that in future generations, palmaris longus may become muscle variations and anomalies. Much more work is done,
present in a very small percentage of normal adult humans and one of the aims of the present book is precisely to pave
and would therefore be considered a rare, “atavistic” vari- the way for such works in the future by providing an atlas of
ation. If in the future palmaris longus will be present in several variants/anomalies found in all body regions of cur-
most human embryos/fetuses but typically absent in adults, rent human individuals of all non-embryonic ages.
those embryonic/fetal stages will indeed somewhat mirror
what happens today in the normal adult human population. TOWARD A MORE HOLISTIC APPROACH
However, although in this case currently the late fetal and
TO HUMAN ANATOMY AND EVOLUTION
adult confgurations of the palmaris longus are essentially
the same in humans (Bardeen 1906), one has to stress again A second and related primary aim of this book is to pave
that the embryonic palmaris longus of those future humans the way for a more holistic approach to human anatomy and
would not directly correspond to the adult palmaris lon- human evolution. Here we contribute to that goal by gather-
gus seen in most humans living today, but instead to the ing and presenting the result of a 20 year-long comparison
Introduction 5
of the muscle variants and anomalies of humans, and Hinchliffe and Johnson (1980: pp. 96–97) described a
between them and those of other animals, in particular our “nearest-neighbor” model for limb muscle to bone relation-
closest relatives, the nonhuman primates. Our Evo-Devo- ships, as well as for muscle to nerve connections. These
Path research includes more traditional Evo-Devo lines of authors argued that such a dynamic, fexible model ensures
research but also pays special attention to data obtained correspondence between these tissues during dramatic
from chordate comparative, developmental, and evolution- evolutionary changes and in congenital malformations.
ary anatomy and from the direct study of normal/abnormal Wolpert’s “positional information” model is similar and
human development (using, e.g., cadaveric collections of suggests that the positional value of a cell/structure (e.g.,
hospitals, museums, and other institutions), with a primary the skeleton) can be recognized by and affect the typical
focus on soft tissues like muscles (e.g., Diogo 2004a,b, patterning/morphogenesis of another cell/structure (e.g.,
2005, 2007; Diogo and Abdala 2007, 2010; Diogo et al. muscles), without changing the overall phenotype/shape of
2008a,b, 2009a,b, 2018, 2019), including primates (e.g., that other cell/structure (Wolpert 1969, 2011). According
Diogo and Wood 2011, 2012b, 2016; Diogo et al. 2010, to this model, muscle cells of limbs do not have positional
2012, 2013a,b, 2017). We hope this work will pave the values, so it would be reasonable to assume that they are
way for a more holistic approach to human anatomy and affected by and follow the positional values of the skeleton,
evolution, as well as help physicians and surgeons, medi- conforming to the expectations of the “nearest neighbor”
cal students, and biologists and anthropologists to better model.
understand where the structures of our body come from, Our soft-tissue studies of humans supported our “nearest
and why they are as they are. As noted above, this applies neighbor” model. As predicted by our model, in a trisomy
to features that (1) have a so-called “normal” confguration 18 fetus with a six-digit hand (partial duplication of thumb)
and are present in more than 50% of the common popula- and a four-digit hand (no thumb), the hand with no thumb
tion; (2) are polymorphisms, and as such, are present in presented all thumb muscles but these muscles went to the
less than 50% and more than 2%; (3) are variations, which most radial digit, which was digit two. As also anticipated
are traits present in up to 2% of the common population; by our model, in the hand with partial thumb duplication
or (4) anomalies, which are found in people with severe there was no duplication of the thumb muscles. This fnd-
malformations or genetic syndromes. The former item ing supports the results of experimental studies on mouse
was discussed Diogo et al. (2016), Learning and under- models demonstrating that limb skeletal and connective/
standing human anatomy and pathology: an evolution- muscle tissue patterning can be uncoupled (e.g., Li et al.
ary and developmental guide for medical students. That 2010). The number of thumb muscles did not change, but
book paved the way not only for a more comprehensive their insertions varied according to the adult topological
understanding of the human body informed by evolution position of the two thumbs. The muscles that typically go
and development, but also for us to compare the common to the radial (e.g., abductor pollicis brevis) and ulnar (e.g.,
phenotype of our internal structures, including muscles, to adductor pollicis) aspects of the thumb now attach onto the
our anatomical polymorphisms, variants, and anomalies, radial side of the most radial thumb and ulnar side of the
which are precisely the subject of the present book. We most ulnar thumb, respectively. Recent comparative studies
provide here an illustrative example to explain how they indicate that the developmental factors for skeletal morpho-
are linked, and how physicians and surgeons, students, logical identity in vertebrates reside less in the embryonic
teachers, and scientists in general can beneft from such a cell lineages, and more in the topological position in the
broader, more holistic approach. embryo, which allows the relative position of bony elements
This example concerns non-pentadactyly, a topic that to be highly conserved in evolution (Hirasawa and Kuratani
has long attracted researchers’ attention because this is 2015).
the most common human limb birth defect. We have stud- Our recent Evo-Devo-Path studies and comparisons
ied alterations in muscle attachments found in congenital have supported, so far, the idea that internal constraints play
non-pentadactyl human limbs and evaluated whether these a central role in both normal and abnormal human devel-
changed patterns are similar to those found in wildtype opment. Our research thus provides support for Alberch’s
nonhuman tetrapods with non-pentadactyl limbs (Smith “logic of monsters,” because there are consistent patterns
et al. 2015). The most signifcant conclusion of those stud- seen in both individuals with different genetic syndromes
ies was that the non-pentadactyl limbs of wildtype taxa and variations of the normal human population. For exam-
(e.g., frogs, salamanders, crocodilians, chickens, humans) ple, Wood (1867a,b, 1868) found that muscle variations are
with birth defects exhibit a consistent “nearest neighbor” much more frequent in the upper limb than in the lower
pattern. In other words, the identity and attachments of the limbs of the normal human population (292 vs 119 cases
distal forelimb and hindlimb muscles seem to be mainly in his sample, i.e., 71% vs 29%, or almost 2.5 times more
related to the physical (topological) position, and not to the upper than lower limb variations). In our sample of 316
number of the anlage or the homeotic identity of the dig- head, upper, and lower limb defects compiled from stud-
its to which the muscles are attached. Topological position ies of severe congenital malformations (see Smith et al.
refers to the adult relationship with other structures, and not 2015), the proportion was exactly the same: 158 upper limb
to the position of the developmental anlagen. defects (50%), 94 head defects (30%), and 64 lower limb
defects (20%), so almost 2.5 times more upper than lower are only present in humans as variations and/or anomalies,
limb defects. Interestingly, most of the defects found in the and therefore have no typical or “normal” presentation
upper limbs (65%) and lower limbs (84%) are also found to be compared with. For most muscles, this section pro-
in the autopods and zeugopods, which are mainly evolu- vides the typical innervation also described by Standring
tionary innovations of tetrapods. This makes sense within (2016). For muscles that are present only as variations and/
an internalist view of evolution, because these phylogeneti- or anomalies, the innervation is listed under “Variations.”
cally more recent innovations are also the last limb regions Innervation may shed light about a muscle’s developmental
to form during development and more prone to develop- origins and is often clinically relevant for variations.
mental changes, variations, and defects. It also makes sense Then, we provide descriptions of the comparative
within an externalist view of evolution, since the more dis- anatomy of each muscle. For most muscles, this section
tal limb regions are in closer contact with prey, substrates, describes the typical presentation and signifcant variations
and objects, and are thus more prone to evolutionary adap- in the apes, the closest living relatives of humans. The apes
tive changes. include gibbons (Hylobates), orangutans (Pongo), gorillas
This connection between development (ontogeny), the (Gorilla), common chimpanzees (Pan troglodytes), and
order in which morphological structures appear in evolu- bonobos (Pan paniscus). When relevant, we included ana-
tion (phylogeny), interactions with the environment (eco- tomical information for other primates or, in some cases,
morphology), and evolvability (adaptation) supports the non-primate animals. “N/A” in this section indicates that,
“logic of monsters” since it predicts a parallel between the to our knowledge, the muscle has not been observed or
variant phenotypes of normal populations and defects in described in the apes.
abnormal individuals. Within the 64 limb muscle defects We then include information on each muscle’s variations
reviewed by Smith et al. (2015) that involve changes of in humans. For most muscles, this section details variations
attachment (origin and insertion), the changes of insertion on the typical presentation that have been described in the
tended to be toward more proximal limb regions, while the literature. For accessory or supernumerary muscles that are
changes of origin tended to be toward more distal regions. present as variations, this section describes its presentation
Overall, most of the attachment changes involved insertion when it is found in humans. Accessory muscles received
changes. This fnding parallels phylogeny, because in pri- their own entries—and were not just listed as variations of
mate and human evolution, more changes involve the inser- other muscles—when enough information was available
tion sites of limb muscles than origin sites. Furthermore, to fll out more than only the “Description” portion of the
this fnding makes sense both within an externalist frame- “Variations” section.
work—as insertions are more distal by defnition and are in For the variations, the prevalence section provides the
closer contact with the outside—and within an internalist prevalence of the variations as listed in the literature. In gen-
framework—as the origin of limb muscles tends to form eral, as noted above, a “typical” confguration is present in
frst in ontogeny, so terminal changes would more likely more than 50% of the common population, polymorphisms
affect their insertion (Diogo and Tanaka 2014; Diogo and are features present in less than 50% and more than 2%, and
Ziermann 2014). Lastly, as noted above, it appears that the variations present in up to 2% of the common population.
upper limb is most prone to phenotypic changes and defects However, as you will see throughout the text, prevalence
among the head and limb regions of the body. Again, more for some presentations that are considered variations often
studies are needed to clarify if these patterns are seen con- exceed 2% and should perhaps be more aptly considered as
cerning other soft tissues such as arteries, veins, nerves, and polymorphisms. “N/A” in this section indicates that, to our
even internal organs, and the information provided in the knowledge, no prevalence data on variations are available.
present book will be hopefully useful to motivate research- We frst included variations described by anatomists in
ers to undertake, and pave the way for, such future works. the 19th century such as Hallett (1848), Macalister (1867a,b;
1875), Knott (1880, 1883a,b), and Wood (1864, 1865, 1866,
1867a,b, 1868, 1870). We then referred to comprehen-
BRIEF NOTES ABOUT OUR METHODOLOGY
sive works such as Mori (1964), Sato (1968a,b,c, 1969,
As noted above, we include information on variations and 1970), Rickenbacher et al. (1985), Bergman et al. (1988),
anomalies present in human muscles after embryonic devel- and Bergman’s Comprehensive Encyclopedia of Human
opment. For each muscle, we provide the Latin name, with Anatomic Variation (2016, edited by Tubbs, Shoja, and
the common English name shown in parentheses, when Loukas) to double-check the information listed in earlier
they are slightly different. We then include the synonyms texts and to better understand the breadth of variations that
of the muscle that we encountered over the course of our exist for each muscle. Lastly, we conducted an exhaustive
research, not meant to be an exhaustive list. In some cases, search for journal articles published through early 2021 to
this section also includes synonyms we believe should be provide additional information on rare variations, variation
attributed to that muscle based on anatomical descriptions. prevalence, and clinical implications of variations.
Next, we describe the typical presentation of each mus- Next, we describe each muscle’s anomalous presenta-
cle. For most muscles, this section provides the typical pre- tions in humans. For most muscles, this section details ana-
sentation as described by Standring (2016). Some muscles tomical anomalies found in individuals with chromosomal
Introduction 7
conditions, developmental defects of the musculoskeletal to our knowledge, no prevalence data on anomalies are
system, or other congenital disorders. “N/A” in this sec- available.
tion indicates that, to our knowledge, no anomalies of that Lastly, in the clinical implications section, we include
muscle have been described. We also include the prevalence information on the clinical relevance of variations and/or
of the anomalies as listed in the literature, mostly provided anomalies. “N/A” in this section indicates that we did not
by Smith et al. (2015). “N/A” in this section indicates that, easily fnd clinical implications of the variations.
2 Head and Neck Muscles
Eve K. Boyle
Vondel S. E. Mahon
Rui Diogo
Warrenkevin Henderson
Boston University
Hannah Jacobson
George Washington University
Noelle Purcell
George Washington University
Kylar Wiltz
CONTENTS
Facial Muscles .....................................................................................................................................................................21
Occipitofrontalis..............................................................................................................................................................21
Synonyms ...................................................................................................................................................................21
Typical Presentation ...................................................................................................................................................21
Comparative Anatomy................................................................................................................................................ 21
Variations.................................................................................................................................................................... 21
Anomalies...................................................................................................................................................................22
Clinical Implications ..................................................................................................................................................22
Orbicularis oculi..............................................................................................................................................................22
Synonyms ...................................................................................................................................................................22
Comparative Anatomy................................................................................................................................................23
Variations....................................................................................................................................................................23
Anomalies...................................................................................................................................................................23
Corrugator supercilii .......................................................................................................................................................23
Synonyms ...................................................................................................................................................................23
Variations....................................................................................................................................................................23
Anomalies...................................................................................................................................................................24
Malaris.............................................................................................................................................................................24
Synonyms ...................................................................................................................................................................24
Variations....................................................................................................................................................................24
DOI: 10.1201/9781003083535-2 9
Anomalies...................................................................................................................................................................24
Orbitozygomaticus ..........................................................................................................................................................24
Synonyms ...................................................................................................................................................................24
Variations....................................................................................................................................................................24
Anomalies...................................................................................................................................................................25
Zygomaticus minor .........................................................................................................................................................25
Synonyms ...................................................................................................................................................................25
Variations....................................................................................................................................................................25
Anomalies...................................................................................................................................................................25
Zygomaticus major..........................................................................................................................................................26
Synonyms ...................................................................................................................................................................26
Variations....................................................................................................................................................................26
Anomalies...................................................................................................................................................................26
Levator labii superioris....................................................................................................................................................27
Synonyms ...................................................................................................................................................................27
Variations.................................................................................................................................................................... 27
Anomalies...................................................................................................................................................................27
Levator labii superioris alaeque nasi ............................................................................................................................... 27
Synonyms ...................................................................................................................................................................27
Variations.................................................................................................................................................................... 28
Anomalies...................................................................................................................................................................28
Clinical Implications .................................................................................................................................................. 28
Levator anguli oris...........................................................................................................................................................28
Synonyms ...................................................................................................................................................................28
Variations....................................................................................................................................................................28
Anomalies...................................................................................................................................................................29
Risorius ...........................................................................................................................................................................29
Synonyms ...................................................................................................................................................................29
Variations....................................................................................................................................................................29
Anomalies...................................................................................................................................................................29
Buccinatorius (buccinator) ..............................................................................................................................................30
Synonyms ...................................................................................................................................................................30
Variations....................................................................................................................................................................30
Anomalies...................................................................................................................................................................30
Head and Neck Muscles 11
Orbicularis oris................................................................................................................................................................30
Synonyms ...................................................................................................................................................................30
Variations....................................................................................................................................................................30
Anomalies...................................................................................................................................................................31
Depressor anguli oris.......................................................................................................................................................31
Synonyms ...................................................................................................................................................................31
Variations.................................................................................................................................................................... 31
Anomalies...................................................................................................................................................................32
Depressor labii inferioris.................................................................................................................................................32
Synonyms ...................................................................................................................................................................32
Variations....................................................................................................................................................................32
Anomalies...................................................................................................................................................................32
Procerus...........................................................................................................................................................................32
Synonyms ...................................................................................................................................................................32
Variations....................................................................................................................................................................33
Anomalies...................................................................................................................................................................33
Nasalis .............................................................................................................................................................................33
Synonyms ...................................................................................................................................................................33
Variations....................................................................................................................................................................33
Anomalies...................................................................................................................................................................33
Depressor septi nasi.........................................................................................................................................................33
Synonyms ...................................................................................................................................................................33
Variations.................................................................................................................................................................... 34
Anomalies...................................................................................................................................................................34
Incisivus labii superioris.................................................................................................................................................. 34
Synonyms ................................................................................................................................................................... 34
Variations....................................................................................................................................................................34
Anomalies...................................................................................................................................................................35
Incisivus labii inferioris...................................................................................................................................................35
Synonyms ...................................................................................................................................................................35
Variations....................................................................................................................................................................35
Anomalies...................................................................................................................................................................35
Mentalis...........................................................................................................................................................................35
Synonyms ...................................................................................................................................................................35
Typical Presentation ................................................................................................................................................... 35

Variations.................................................................................................................................................................... 35
Anomalies................................................................................................................................................................... 36
Platysma myoides (platysma) ............................................................................................................................................ 36
Synonyms ................................................................................................................................................................... 36
Variations.................................................................................................................................................................... 36
Anomalies................................................................................................................................................................... 37
Platysma cervicale........................................................................................................................................................... 37
Synonyms ................................................................................................................................................................... 37
Variations.................................................................................................................................................................... 37
Anomalies................................................................................................................................................................... 38
Muscles of Mastication ........................................................................................................................................................ 38
Masseter .......................................................................................................................................................................... 38
Synonyms ................................................................................................................................................................... 38
Variations.................................................................................................................................................................... 39
Anomalies................................................................................................................................................................... 39
Temporalis....................................................................................................................................................................... 40
Synonyms ................................................................................................................................................................... 40
Variations.................................................................................................................................................................... 40
Anomalies................................................................................................................................................................... 40
Pterygoideus lateralis (Lateral pterygoid)....................................................................................................................... 41
Synonyms ................................................................................................................................................................... 41
Variations.................................................................................................................................................................... 41
Anomalies................................................................................................................................................................... 42
Pterygoideus medialis (Medial pterygoid) ...................................................................................................................... 43
Synonyms ................................................................................................................................................................... 43
Variations.................................................................................................................................................................... 43
Anomalies................................................................................................................................................................... 43
Sphenotemporalis............................................................................................................................................................ 44
Synonyms ................................................................................................................................................................... 44
Variations.................................................................................................................................................................... 44
Anomalies................................................................................................................................................................... 44
Pterygospinous ................................................................................................................................................................ 44
Synonyms ................................................................................................................................................................... 44

Variations....................................................................................................................................................................44
Anomalies...................................................................................................................................................................45
Pterygoideus proprius......................................................................................................................................................45
Synonyms ...................................................................................................................................................................45
Variations....................................................................................................................................................................45
Anomalies...................................................................................................................................................................45
Extrinsic Muscles of the Ear and Middle Ear Muscles........................................................................................................46
Auricularis anterior .........................................................................................................................................................46
Synonyms ...................................................................................................................................................................46
Variations....................................................................................................................................................................46
Anomalies...................................................................................................................................................................46
Auricularis superior.........................................................................................................................................................47
Synonyms ...................................................................................................................................................................47
Variations.................................................................................................................................................................... 47
Anomalies...................................................................................................................................................................47
Auricularis posterior........................................................................................................................................................ 47
Synonyms ...................................................................................................................................................................47
Variations.................................................................................................................................................................... 47
Anomalies...................................................................................................................................................................48
Tensor tympani (Not Illustrated).....................................................................................................................................48
Synonyms ...................................................................................................................................................................48
Variations....................................................................................................................................................................48
Anomalies...................................................................................................................................................................48
Stapedius (Not Illustrated) .............................................................................................................................................. 48
Synonyms ...................................................................................................................................................................48
Variations....................................................................................................................................................................49
Anomalies...................................................................................................................................................................49
Muscles of the Tongue .........................................................................................................................................................49
Intrinsic Muscles of the Tongue (Not Illustrated) ...........................................................................................................49
Synonyms ...................................................................................................................................................................49
Variations....................................................................................................................................................................49
Anomalies...................................................................................................................................................................49
Genioglossus ...................................................................................................................................................................50
Synonyms ................................................................................................................................................................... 50
Variations.................................................................................................................................................................... 51
Anomalies................................................................................................................................................................... 51
Hyoglossus ...................................................................................................................................................................... 51
Synonyms ................................................................................................................................................................... 51
Variations.................................................................................................................................................................... 52
Anomalies................................................................................................................................................................... 52
Styloglossus..................................................................................................................................................................... 52
Synonyms ................................................................................................................................................................... 52
Variations.................................................................................................................................................................... 53
Anomalies................................................................................................................................................................... 53
Myloglossus .................................................................................................................................................................... 53
Synonyms ................................................................................................................................................................... 53
Variations.................................................................................................................................................................... 53
Anomalies................................................................................................................................................................... 54
Styloauricularis ............................................................................................................................................................... 54
Synonyms ................................................................................................................................................................... 54
Variations.................................................................................................................................................................... 54
Anomalies................................................................................................................................................................... 55
Muscles of the Soft Palate.................................................................................................................................................... 55
Tensor veli palatini .......................................................................................................................................................... 55
Synonyms ................................................................................................................................................................... 55
Variations.................................................................................................................................................................... 56
Anomalies................................................................................................................................................................... 56
Levator veli palatini......................................................................................................................................................... 57
Synonyms ................................................................................................................................................................... 57
Variations.................................................................................................................................................................... 57
Anomalies................................................................................................................................................................... 57
Musculus uvulae.............................................................................................................................................................. 57
Synonyms ................................................................................................................................................................... 57
Variations.................................................................................................................................................................... 58
Anomalies................................................................................................................................................................... 58
Palatoglossus ................................................................................................................................................................... 58
Synonyms ...................................................................................................................................................................58
Variations....................................................................................................................................................................58
Anomalies...................................................................................................................................................................58
Palatopharyngeus ............................................................................................................................................................59
Synonyms ...................................................................................................................................................................59
Variations....................................................................................................................................................................59
Anomalies...................................................................................................................................................................60
Salpingopharyngeus (Not Illustrated) .............................................................................................................................60
Synonyms ...................................................................................................................................................................60
Variations....................................................................................................................................................................60
Anomalies...................................................................................................................................................................61
Pharyngeal Muscles .............................................................................................................................................................61
Constrictor pharyngis superior (Superior pharyngeal constrictor)..................................................................................61
Synonyms ...................................................................................................................................................................61
Variations.................................................................................................................................................................... 61
Anomalies...................................................................................................................................................................62
Stylopharyngeus..............................................................................................................................................................62
Synonyms ...................................................................................................................................................................62
Variations.................................................................................................................................................................... 63
Anomalies...................................................................................................................................................................63
Petropharyngeus..............................................................................................................................................................63
Synonyms ...................................................................................................................................................................63
Variations....................................................................................................................................................................63
Anomalies...................................................................................................................................................................64
Constrictor pharyngis medius (Middle pharyngeal constrictor) .....................................................................................64
Synonyms ................................................................................................................................................................... 64
Variations....................................................................................................................................................................64
Anomalies...................................................................................................................................................................64
Constrictor pharyngis inferior (Inferior pharyngeal constrictor) .................................................................................... 65
Synonyms ...................................................................................................................................................................65
Variations....................................................................................................................................................................65
Anomalies...................................................................................................................................................................65
Laryngeal Muscles ...............................................................................................................................................................65
Cricothyroideus (Cricothyroid).......................................................................................................................................65
Synonyms ...................................................................................................................................................................65
Variations....................................................................................................................................................................66
Anomalies...................................................................................................................................................................67
Thyrotrachealis................................................................................................................................................................67
Synonyms ...................................................................................................................................................................67
Variations....................................................................................................................................................................67
Anomalies...................................................................................................................................................................67
Ceratocricoid ...................................................................................................................................................................67
Synonyms ...................................................................................................................................................................67
Variations....................................................................................................................................................................67
Anomalies...................................................................................................................................................................68
Cricoarytenoideus posterior (Posterior cricoarytenoid) ..................................................................................................68
Synonyms ...................................................................................................................................................................68
Variations....................................................................................................................................................................69
Anomalies...................................................................................................................................................................69
Cricoarytenoideus lateralis (Lateral cricoarytenoid)....................................................................................................... 69
Synonyms ...................................................................................................................................................................69
Variations....................................................................................................................................................................69
Anomalies...................................................................................................................................................................70
Arytenoideus transversus (Transverse arytenoid) ...........................................................................................................70
Synonyms ...................................................................................................................................................................70
Variations....................................................................................................................................................................70
Anomalies...................................................................................................................................................................70
Arytenoideus obliquus (Oblique arytenoid)....................................................................................................................70
Synonyms ................................................................................................................................................................... 70
Variations....................................................................................................................................................................71
Anomalies...................................................................................................................................................................71
Thyroarytenoideus (Thyroarytenoid) ..............................................................................................................................71
Synonyms ...................................................................................................................................................................72
Variations....................................................................................................................................................................72
Anomalies...................................................................................................................................................................72
Superior thyroarytenoideus (Superior thyroarytenoid) ...................................................................................................72
Synonyms ...................................................................................................................................................................72
Variations....................................................................................................................................................................72
Anomalies...................................................................................................................................................................73
Infrahyoid Muscles ..............................................................................................................................................................73
Sternohyoideus (Sternohyoid).........................................................................................................................................73
Synonyms ...................................................................................................................................................................73
Variations....................................................................................................................................................................74
Anomalies...................................................................................................................................................................74
Cleidohyoideus (Cleidohyoid) ........................................................................................................................................74
Synonyms ...................................................................................................................................................................75
Variations....................................................................................................................................................................75
Anomalies...................................................................................................................................................................75
Omohyoideus (Omohyoid)..............................................................................................................................................75
Synonyms ...................................................................................................................................................................75
Variations....................................................................................................................................................................75
Anomalies...................................................................................................................................................................76
Sternothyroideus (Sternothyroid)....................................................................................................................................77
Synonyms ...................................................................................................................................................................77
Variations.................................................................................................................................................................... 77
Anomalies...................................................................................................................................................................78
Thyrohyoideus (Thyrohyoid) .......................................................................................................................................... 78
Synonyms ...................................................................................................................................................................78
Variations.................................................................................................................................................................... 78
Anomalies...................................................................................................................................................................78
Levator glandulae thyroideae .......................................................................................................................................... 78
Synonyms ................................................................................................................................................................... 78
Variations....................................................................................................................................................................79
Anomalies...................................................................................................................................................................79
Suprahyoid Muscles.............................................................................................................................................................79
Mylohyoideus (Mylohyoid) ............................................................................................................................................79
Synonyms ...................................................................................................................................................................79
Variations....................................................................................................................................................................80
Anomalies...................................................................................................................................................................80
Geniohyoideus (Geniohyoid) .......................................................................................................................................... 81

Synonyms ................................................................................................................................................................... 81
Variations.................................................................................................................................................................... 81
Anomalies................................................................................................................................................................... 81
Digastricus anterior (Anterior Belly of the Digastric) .................................................................................................... 82
Synonyms ................................................................................................................................................................... 82
Variations.................................................................................................................................................................... 82
Anomalies................................................................................................................................................................... 84
Digastricus posterior (Posterior Belly of the Digastric).................................................................................................. 85
Synonyms ................................................................................................................................................................... 85
Variations.................................................................................................................................................................... 85
Anomalies................................................................................................................................................................... 86
Stylohyoideus (Stylohyoid)............................................................................................................................................. 87
Synonyms ................................................................................................................................................................... 87
Variations.................................................................................................................................................................... 87
Anomalies................................................................................................................................................................... 88
Extraocular Muscles............................................................................................................................................................. 88
Levator palpebrae superioris ........................................................................................................................................... 88
Synonyms ................................................................................................................................................................... 88
Variations.................................................................................................................................................................... 88
Anomalies................................................................................................................................................................... 90
Tensor trochleae .............................................................................................................................................................. 90
Synonyms ................................................................................................................................................................... 90
Variations.................................................................................................................................................................... 90
Anomalies................................................................................................................................................................... 90
Rectus superior (Superior rectus).................................................................................................................................... 90
Synonyms ................................................................................................................................................................... 90
Variations.................................................................................................................................................................... 91
Anomalies................................................................................................................................................................... 91
Rectus inferior (Inferior rectus) ...................................................................................................................................... 91
Synonyms ................................................................................................................................................................... 91
Variations.................................................................................................................................................................... 91
Anomalies................................................................................................................................................................... 92
Rectus medialis (Medial rectus)......................................................................................................................................92

Synonyms ...................................................................................................................................................................92
Variations....................................................................................................................................................................92
Anomalies...................................................................................................................................................................92
Rectus lateralis (Lateral rectus).......................................................................................................................................92
Synonyms ...................................................................................................................................................................92
Variations.................................................................................................................................................................... 93
Anomalies...................................................................................................................................................................93
Obliquus oculi superior (Superior oblique)..................................................................................................................... 93
Synonyms ...................................................................................................................................................................93
Variations.................................................................................................................................................................... 93
Anomalies...................................................................................................................................................................94
Obliquus oculi inferior (Inferior oblique) .......................................................................................................................94
Synonyms ...................................................................................................................................................................94
Variations....................................................................................................................................................................94
Anomalies...................................................................................................................................................................95
Scalene, Prevertebral, and Suboccipital Muscles.................................................................................................................95
Scalenus anterior (Anterior scalene) ...............................................................................................................................95
Synonyms ...................................................................................................................................................................95
Variations....................................................................................................................................................................95
Anomalies...................................................................................................................................................................96
Scalenus medius (Middle scalene) .................................................................................................................................. 96
Synonyms ...................................................................................................................................................................96
Variations....................................................................................................................................................................96
Anomalies...................................................................................................................................................................97
Scalenus posterior (Posterior scalene) ............................................................................................................................ 97
Synonyms ...................................................................................................................................................................97
Variations.................................................................................................................................................................... 97
Anomalies...................................................................................................................................................................98
Scalenus minimus............................................................................................................................................................98
Synonyms ...................................................................................................................................................................98
Variations....................................................................................................................................................................98
Anomalies...................................................................................................................................................................99
Longus colli.....................................................................................................................................................................99
Synonyms ...................................................................................................................................................................99
Variations....................................................................................................................................................................99
Anomalies.................................................................................................................................................................100
Clinical Implications ................................................................................................................................................ 100
Longus capitis ...............................................................................................................................................................100
Synonyms .................................................................................................................................................................100
Typical Presentation .................................................................................................................................................100
Comparative Anatomy.............................................................................................................................................. 100
Variations.................................................................................................................................................................. 100
Anomalies.................................................................................................................................................................101
Rectus capitis anterior ...................................................................................................................................................101
Synonyms .................................................................................................................................................................101
Comparative Anatomy..............................................................................................................................................101
Variations..................................................................................................................................................................101
Anomalies.................................................................................................................................................................101
Clinical Implications ................................................................................................................................................101
Rectus capitis lateralis...................................................................................................................................................101
Synonyms .................................................................................................................................................................101
Variations..................................................................................................................................................................102
Anomalies.................................................................................................................................................................102
Rectus capitis posterior major.......................................................................................................................................102
Synonyms .................................................................................................................................................................102
Variations..................................................................................................................................................................102
Anomalies.................................................................................................................................................................103
Rectus capitis posterior minor.......................................................................................................................................104
Synonyms .................................................................................................................................................................104
Variations.................................................................................................................................................................. 104
Anomalies.................................................................................................................................................................105
Obliquus capitis superior...............................................................................................................................................105
Synonyms ................................................................................................................................................................. 105
Variations..................................................................................................................................................................105
Anomalies.................................................................................................................................................................105
Obliquus capitis inferior................................................................................................................................................105
Synonyms .................................................................................................................................................................105
Variations..................................................................................................................................................................106
Anomalies.................................................................................................................................................................106
Posterolateral Neck Muscles..............................................................................................................................................106

Sternocleidomastoideus (Sternocleidomastoid) ............................................................................................................ 106
Synonyms .................................................................................................................................................................106
Variations.................................................................................................................................................................. 106
Anomalies.................................................................................................................................................................107
Cleido-occipitalis cervicalis .......................................................................................................................................... 107
Synonyms .................................................................................................................................................................107
Variations..................................................................................................................................................................107
Anomalies.................................................................................................................................................................108
Trapezius .......................................................................................................................................................................108
Synonyms .................................................................................................................................................................108
Variations..................................................................................................................................................................108
Anomalies.................................................................................................................................................................109
FACIAL MUSCLES occipitalis is typically differentiated into a main portion

(occipitalis proprius) and a cervico-auriculo-occipitalis
OccipitOfrOntalis (figures 2.1, 2.3) portion (Deniker 1885; Lightoller 1928; Loth 1931; Diogo
Synonyms et al. 2012). A vestigial cervico-auriculo-occipitalis bundle
The occipital and frontal components of occipitofrontalis may be present rarely in common chimpanzees (Seiler
may be referred to as two separate muscles: occipitalis and 1976) and gorillas (Ruge 1887). In orangutans, occipitalis
frontalis. is split into a pars profunda and pars superfcialis (Sullivan
and Osgood 1925; Diogo et al. 2013b).
Typical Presentation
Description Variations
Occipitofrontalis consists of the occipitalis and fron- Description
talis muscles, connected by the epicranial aponeurosis The muscular portions of occipitofrontalis may fuse or
(Standring 2016). Occipitalis extends from the highest decussate across the midline (Macalister 1875; Knott
nuchal line and mastoid region to attach to the epicranial 1883a, Bergman et al. 1988; Watanabe 2016; Raveendran
aponeurosis (Standring 2016). Frontalis extends from the and Anthony 2021). Either the occipital or frontal por-
superfcial fascia of the eyebrows to attach to the epicra- tion may be absent (Macalister 1875; Bergman et al. 1988;
nial aponeurosis, having connections with other forehead Watanabe 2016). The muscle fbers of the frontalis and
muscles (Standring 2016). occipitalis may directly connect (Macalister 1875). Both
the occipitalis and frontalis may be divided into fascicles
Innervation instead of presenting as continuous sheets (Macalister
Occipitofrontalis is innervated by the facial nerve, the 1875; Knott 1883a). When fasciculated, occipitalis may
occipital portion via the posterior auricular branch and the split into superior and inferior parts (Macalister 1875).
frontal portion via the temporal branches (Standring 2016). Occipitalis may join with auricularis posterior or receive
fbers from sternocleidomastoid (Macalister 1875).
Comparative Anatomy Frontalis can vary in its extent and at the level where it
Frontalis has a similar typical presentation in the apes, attaches to the epicranial aponeurosis (Macalister 1875;
extending from the skin of the eyebrow and nose to the Knott 1883a). It may have attachments directly to the
epicranial aponeurosis (Raven 1950; Miller 1952; Gibbs nasal or maxillary bones (Macalister 1875; Bergman
1999; Diogo et al. 2010, 2012, 2013a,b, 2017). Occipitalis et al. 1988). It may send a slip to levator labii superioris
typically presents similarly to humans in common chim- alaeque nasi, or more rarely, the medial palpebral liga-
panzees, bonobos, and gorillas, extending from the occipi- ment (Macalister 1875; Knott 1883a). Temporoparietalis
tal and mastoid regions to the epicranial aponeurosis is occasionally present between frontalis, auricularis ante-
(Gibbs 1999; Diogo et al. 2010, 2013a, 2017). In gibbons, rior, and auricularis superior (Standring 2016).
FIGURE 2.1 Muscles of the face in anterior view. In all fgures, muscles that are present as variations and/or anomalies are bolded.
Prevalence Prevalence
In a sample of 26 cadavers, Raveendran and Anthony (2021) Occipitofrontalis was hypoplastic in all eight cases of
found that fbers of the frontalis muscles decussated only trisomy 18 (100%) examined by Bersu and Ramirez-Castro
near the eyebrows before diverging in seven cases (26.9%), (1977).
decussated up to the middle of the muscles before diverging
in 12 cases (46.2%), and did not decussate at all in seven Clinical Implications
cases (26.9%).
Understanding variations in frontalis decussation is help-
ful for administering botulinum toxin type A therapies
Anomalies (Raveendran and Anthony 2021).
Description
In a fetus with craniorachischisis, the frontalis was situated
Orbicularis Oculi (figure 2.1)
deep to a layer of fatty tissue, and the occipitalis was miss-
ing, bilaterally (Alghamdi et al. 2017). In a female fetus with See also: Malaris, Orbitozygomaticus
trisomy 18, the left and right frontalis muscles were fused
at the midline (Alghamdi et al. 2018). Occipitofrontalis may Synonyms
also be hypoplastic in individuals with trisomy 18 (Bersu This muscle may also be referred to as orbicularis
and Ramirez-Castro 1977). palpebrarum (Macalister 1875).
Typical Presentation Prevalence

Description Sato (1968a) found orbicularis oculi present in all 190 facial
Orbicularis oculi is comprised of orbital, palpebral, and sides (100%) in Kyushu-Japanese males and present in all
lacrimal parts and a ciliary bundle (Standring 2016). The 121 sides (100%) in females.
fbers of the orbital part form complete ellipses around
the orbit and originate from the nasal part of the frontal Anomalies
bone, the frontal process of the maxilla, and the medial Description
palpebral ligament (Standring 2016). These fbers form In a boy with trisomy 13q, the orbicularis oculi muscles
depressor supercilii (Macalister 1875; Standring 2016). covered most of the midface and their lower borders lay
The fbers of the palpebral part originate from the medial below the level of the nostrils (Pettersen 1979). In a fetus
palpebral ligament, course across the eyelids, and end as with trisomy 18 and cyclopia, the orbicularis oculi muscles
the lateral palpebral raphe (Standring 2016). The fbers appeared to converge at the midline and expand laterally
of the lacrimal part originate from the lacrimal bone and (Smith et al. 2015). On the left side of this specimen, orbicu-
insert into the tarsi of the eyelids and the lateral palpebral laris oculi blended with auricularis anterior. In individuals
raphe (Standring 2016). with trisomy 21, Bersu (1980) observed that the superfcial
muscles in the midface region were poorly differentiated
Innervation and replaced by a sheet of muscle that was continuous later-
Orbicularis oculi is innervated by the temporal and zygo- ally with the orbital part of orbicularis oculi.
matic branches of the facial nerve (Standring 2016).
Prevalence
Comparative Anatomy Bersu (1980) found that the midface muscles were replaced
Orbicularis oculi has a similar typical presentation in by a single muscle sheet that was continuous with orbicularis
the apes, as it is divided into orbital, palpebral, and lac- oculi in fve out of fve individuals with trisomy 21 (100%).
rimal parts with a ciliary bundle, depressor supercilii,
bony attachments around the orbit, and an origin from the Clinical Implications
medial palpebral ligament (Deniker 1885; Sullivan and N/A
Osgood 1925; Lightoller 1928; Raven 1950; Miller 1952;
Seiler 1976; Gibbs 1999; Burrows et al. 2006; Diogo et al.
2010, 2012, 2013a,b, 2017). It may send a tendon to the cOrrugatOr supercilii (figure 2.1)
zygomatic bone in orangutans (Sullivan and Osgood 1925;
Synonyms
Diogo et al. 2013b).
N/A
Variations
Description
Description
Orbicularis oculi connects variably with frontalis, corru-
Corrugator supercilii originates from the medial aspect of
gator supercilii, zygomaticus minor, levator labii superi-
the superciliary arch and attaches to the skin over the mid-
oris, and levator labii superioris alaeque nasi (Macalister
dle part of the supraorbital margin (Standring 2016). It is
1875; Standring 2016; Watanabe 2016). It may send
situated deep to, and mingles with, orbicularis oculi and the
a slip to levator palpebrae superioris (Knott 1883a).
frontal portion of occipitofrontalis (Standring 2016).
Zygomaticus minor may originate entirely from orbicu-
laris oculi (Macalister 1875; Knott 1883a). Platysma may Innervation
reach orbicularis oculi (Macalister 1875; Bergman et al.
Corrugator supercilii is innervated by the temporal branches
1988; Watanabe 2016). The orbicularis oculi muscles may
of the facial nerve (Standring 2016).
connect via a transversus glabellae muscle (Burkitt and
Lightoller 1926; Bergman et al. 1988; Watanabe 2016). Comparative Anatomy
There may be a fascicle that surrounds the lacus lacri-
Corrugator supercilii has a similar typical presentation in
malis (Macalister 1875). The orbital part may be absent
the apes, extending from the medial end of the superciliary
(Bergman et al. 1988; Watanabe 2016). The peripheral
arch to the eyebrow region (Deniker 1885; Lightoller 1928;
fbers may be absent (Macalister 1875). The orbital and
Miller 1952; Seiler 1976; Gibbs 1999; Diogo et al. 2010,
palpebral parts may be completely separate (Macalister
2012, 2013a,b, 2017).
1875; Bergman et al. 1988; Watanabe 2016). Orbicularis
oculi is associated with two variable muscles: malaris
Variations
and the orbitozygomatic muscle (see the entries for these
muscles) (Henle 1858, 1871; Lightoller 1925; Hwang et al. Description
2002; Park et al. 2011, 2012; Zufferey 2013; Standring Corrugator supercilii may be completely separate from, or
2016; Watanabe 2016; Kampan et al. 2018). completed fused with, orbicularis oculi (Macalister 1875;
Watanabe 2016). It may be split into several fasciculi or arise angle of the mouth (Henle 1858, 1871; Lightoller 1925; Park
from two or three tendinous slips (Macalister 1875). It may be et al. 2011; Standring 2016; Watanabe 2016). It may blend
bilaminar (Macalister 1875). It may vary in its superior and with zygomaticus major, zygomaticus minor, levator labii
lateral extents (Janis et al. 2007). Corrugator supercilii may be superioris, and/or orbicularis oris (Lightoller 1925; Standring
absent (Macalister 1875; Sato 1968a; Watanabe 2016). 2016). Kampan et al. (2018) found a suspending bundle
between the attachments of malaris (lateral bundle) and the
Prevalence orbitozygomatic muscle (medial bundle) and suggest all three
Sato (1968a) found corrugator supercilii present in 269 out bundles are part of the same malaris muscle complex.
of 328 facial sides (82.01%) in Kyushu-Japanese males and
present in 181 out of 220 sides (82.27%) in females. Innervation
Malaris is innervated by the temporal branches of the facial
Anomalies nerve (Kampan et al. 2018).
Description
Corrugator supercilii was absent in a fetus with trisomy 18 Prevalence
and cyclopia (Smith et al. 2015) and a fetus with craniora- In a sample of 61 hemifaces, Park et al. (2011) found malaris
chischisis (Alghamdi et al. 2017). present in 33 cases (54.1%). It ended at the zygomatic arch
in 17 cases (27.9%), in the cheek region in 11 cases (18%),
Prevalence and at the angle of the mouth in fve cases (8.2%). In a sam-
N/A ple of 22 head halves, Kampan et al. (2018) found lateral
bundles of orbicularis oculi in all cases (100%).
Clinical Implications
Understanding variability in the extent of corrugator super- Anomalies
cilii and its varied relationships with the supraorbital nerve N/A
is important for successfully performing complete resec-
tion of this muscle (Janis et al. 2007, 2008). Clinical Implications
Understanding variations in the presence and presentation
of malaris is helpful for planning and administering perior-
Malaris (figure 2.1) bital botulinum toxin type A (Botox) injections (Park et al.
2011; Zufferey 2013).
See also: Orbicularis oculi
Synonyms OrbitOzygOMaticus (figure 2.1)

This muscle may be referred to as the lateral muscular band See also: Orbicularis oculi
of orbicularis oculi (Park et al. 2011) or the lateral bundle
of the malaris muscle (Henle 1858, 1871; Lightoller 1925). Synonyms
This muscle may be referred to as the medial muscular band
Typical Presentation of orbicularis oculi (Park et al. 2012), the medial bundle
of the malaris muscle (Henle 1858, 1871; Lightoller 1925),
This muscle is only present as a variation.
zygomatico-orbicularis muscle (Diogo et al. 2013a,b, 2017),
Comparative Anatomy or malar levator (Snider et al. 2017).
A “pars orbicularis malaris” of orbicularis oculi was Typical Presentation
described in orangutans as being similar to zygomaticus
minor and blended with zygomaticus major (Sullivan and
Osgood 1925; Diogo et al. 2013b). Pellatt (1979) reported Comparative Anatomy
the presence of malaris in common chimpanzees, and some
Diogo et al. (2013a,b) state that “zygomatico-orbicularis” is
authors designate malaris as a bundle of zygomaticus major
absent in orangutans and common chimpanzees.
in this species (Burrows et al. 2006; Diogo et al. 2013a).
Variations Variations
Description Description
When present, malaris is a thin band of muscle that originates Hwang et al. (2002) used “orbitozygomatic muscle” to refer
from the superfcial temporal fascia lateral to orbicularis oculi to a muscle easily separated from orbicularis oculi that
and is continuous with the inferolateral margin of that muscle originated from the medial palpebral ligament and inserted
(Henle 1858, 1871; Lightoller 1925; Park et al. 2011; Zufferey into the zygomaticus muscles and the skin of the cheek.
2013; Standring 2016; Watanabe 2016). Its extent varies and it Park et al. (2012) found that it may originate from the fron-
may end at the zygomatic arch, in the cheek region, or at the tal portion of occipitofrontalis without an origin from the
medial palpebral ligament. Park et al. (2012) also found that Comparative Anatomy
the muscle may end above the inferior border of orbicularis Zygomaticus minor has a similar typical presentation in the
oculi. Snider et al. (2017) describe a “malar levator” situ- apes, extending from the zygomatic bone and orbicularis
ated between orbicularis oculi and levator labii superioris oculi to the upper lip and corner of the mouth (Raven 1950;
alaeque nasi that inserted into the malar fat pad. Kampan Miller 1952; Gibbs 1999; Diogo et al. 2010, 2012, 2013a,b,
et al. (2018) found a suspending bundle between the attach- 2017). It may occasionally fuse with zygomaticus major,
ments of malaris (lateral bundle) and the orbitozygomatic forming a muscle mass that decussates with nearby muscles
muscle (medial bundle) and suggest all three bundles are (Gibbs 1999).
part of the same malaris muscle complex.
Variations
Innervation
Description
The orbitozygomatic muscle is innervated by the zygomatic
Zygomaticus minor may receive a slip from risorius or
branches of the facial nerve (Kampan et al. 2018).
orbicularis oculi (Macalister 1875; Watanabe 2016). It may
Prevalence also originate entirely from orbicularis oculi (Macalister
1875; Knott 1883a). It may be fused with zygomaticus
In a sample of 42 hemifaces, Hwang et al. (2002) found the
major (Macalister 1875; Bergman et al. 1988; Watanabe
orbitozygomatic muscle present in 17 cases (40.5%). In a
2016). Macalister (1875) notes that Eustachius described
sample of 22 head halves, Kampan et al. (2018) found medial
a rare case in which zygomaticus minor joined with fron-
bundles of orbicularis oculi in nine cases (40.9%). Snider
talis. Zygomaticus minor may end before it reaches the
et al. (2017) found a malar levator in all 12 cadavers they
upper lip (Macalister 1875). It may have an additional
dissected (100%). In a sample of 61 hemifaces, Park et al.
attachment to the lateral alar region of the nose (Choi
(2012) found the orbitozygomatic muscle present in 40 cases
et al. 2014). It may be doubled or, more rarely, tripled upon
(65.6%). It attached to the frontal portion of occipitofronta-
insertion (Macalister 1875; Bergman et al. 1988; Watanabe
lis without an attachment to the medial palpebral ligament
2016). Zygomaticus minor may also be absent (Macalister
in 14 cases (23%). It extended between the medial palpe-
1875; Sato 1968a; Bergman et al. 1988; Pessa et al. 1998;
bral ligament and the skin of the cheek in 14 cases (23%). It
Watanabe 2016).
inserted into the cheek skin and attached to the frontal por-
tion of occipitofrontalis without an attachment to the medial Prevalence
palpebral ligament in 12 cases (19.7%). In 14 cases (23%),
Sato (1968a) found zygomaticus minor absent in 8 out of
the insertion of the muscle did not pass the inferior border
378 facial sides (2.12%) in Kyushu-Japanese males and
of orbicularis oculi. In 12 cases (19.7%), the insertion of the
absent in 13 out of 242 sides (5.37%) in females. Pessa et al.
muscle was beyond the inferior border of orbicularis oculi.
(1998) found zygomaticus minor present only in 18 out of
Anomalies 50 (36%) hemifacial cadaver dissections. Farahvash et al.
(2010) found zygomaticus minor present in 31 out of 52
N/A
(59.6%) hemifacial cadaver dissections. Choi et al. (2014)
Clinical Implications classified the anatomy of zygomaticus minor into three
types based on dissections of 54 hemifaces from 30 cadav-
As neurovascular structures travel in the deep surface of
ers. Zygomaticus minor attached to the upper lip only in 34
the orbitozygomatic muscle, it may be a useful landmark to
cases (type A, 63%). In 17 of these cases, the muscle was
avoid injury when performing a subciliary incision (Hwang
straight (31.5%), and in the 17 other cases (31.5%), the mus-
et al. 2002).
cle was curved. Zygomaticus minor attached to the upper
lip and the lateral alar region in 15 cases (type B, 27.8%).
Zygomaticus minor (Figure 2.1) Zygomaticus minor was poorly developed or absent in five
cases (type C, 9.2%).
Synonyms
N/A
Anomalies
Typical Presentation Description
Description In a fetus with craniorachischisis, zygomaticus minor was
Zygomaticus minor originates from the zygomatic bone and absent on the right side (Alghamdi et al. 2017). In an infant
ends in the muscles of the upper lip, where it merges with with mandibulofacial dysostosis, zygomaticus minor was
levator labii superioris (Standring 2016). absent and replaced with connective tissue (Herring et al.
1979). In a neonate with trisomy 13, zygomaticus major and
Innervation minor formed a continuous sheet of muscle on the right side
Zygomaticus minor is innervated by the buccal and (Aziz 1980). In two neonates with trisomy 18, a continuous
zygomatic branches of the facial nerve (Standring 2016). sheet of muscle fibers extended from the cheek to the nose
and the fbers of zygomaticus minor, zygomaticus major, and 2016). It may connect with orbicularis oculi or platysma
levator labii superioris could not be differentiated (Aziz 1979, (Macalister 1875).
1981). Similarly, in a female fetus with trisomy 18, zygomati- It may originate from the masseteric fascia (Macalister
cus major, zygomaticus minor, levator labii superioris, levator 1875). Zygomaticus major may fuse with depressor anguli
labii superioris alaeque nasi, and levator anguli oris were fused oris at its insertion (Macalister 1875). It may be absent
bilaterally and formed a single muscle sheet that extended from (Macalister 1875; Sato 1968a; Bergman et al. 1988).
the zygomatic bone and inferior margin of orbicularis oculi to Zygomaticus major may be doubled throughout or at its
the upper lip (Alghamdi et al. 2018). These authors describe insertion, and this latter presentation is often referred to as
this muscle sheet as the “complex triangle.” Individuals with bifd zygomaticus major (Macalister 1875; Bergman et al.
trisomy 21 also show poor differentiation of the midface mus- 1988; Watanabe 2016).
cles, as the zygomaticus muscles and levator labii superioris
are not distinct and are replaced by a muscle sheet that extends Prevalence
between orbicularis oculi and orbicularis oris (Bersu 1980). Sato (1968a) found zygomaticus major present in 376 out
Prevalence present in 245 out of 248 sides (98.79%) in females. In a
Replacement of distinct zygomaticus muscles and levator labii sample of 50 hemifaces, Pessa et al. (1998) found a single
superioris with a single sheet of muscle was found in fve out zygomaticus major in 33 cases (66%) and a double or bifd
of fve (100%) individuals with trisomy 21 (Bersu 1980). zygomaticus major in 17 cases (34%). In a sample of 70
cadavers, Hu et al. (2008) found a bifd zygomaticus major
Clinical Implications in 28 cases (40%). In a sample of 52 hemifaces, Farahvash
The attachment of zygomaticus minor into the nose ala et al. (2010) found a single zygomaticus major in 42 cases
could aggravate nasal faring and action of other muscles in (80.8%) and a bifd zygomaticus major in ten cases (19.2%).
this region and may also form a protrusion next to the nose In a sample of 29 sides from 15 heads, Elvan et al. (2020)
ala during certain facial expressions (Choi et al. 2014). found a bifd zygomaticus major on four sides (13.8%). In
a meta-analysis of seven studies (including some detailed
here), Phan and Onggo (2019) found that the overall preva-
zygOMaticus MajOr (figure 2.1) lence of bifd zygomaticus major is 22.7%.
Synonyms
N/A Anomalies
Description
In a fetus with craniorachischisis, zygomaticus major
Description
was absent on the right side (Alghamdi et al. 2017). In
Zygomaticus major originates from the zygomatic bone and a fetus with trisomy 18 and cyclopia, zygomaticus major
ends at the angle of the mouth, where it merges with levator originated from orbicularis oculi and the zygomatic pro-
anguli oris and orbicularis oris (Standring 2016). cess and spread wider than is typical (Smith et al. 2015).
It also received fbers from platysma. In an infant with
Innervation
mandibulofacial dysostosis, zygomaticus major origi-
Zygomaticus major is innervated by the buccal and nated from the zygomatic process of the frontal bone
zygomatic branches of the facial nerve (Standring 2016). (Herring et al. 1979).
In a neonate with trisomy 13, zygomaticus major and
Comparative Anatomy
minor formed a continuous sheet of muscle on the right
Zygomaticus major has a similar typical presentation in the side (Aziz 1980). In two neonates with trisomy 18, a con-
apes, extending from the zygomatic bone and the tempo- tinuous sheet of muscle fbers extended from the cheek to
ralis fascia to the corner of the mouth (Raven 1950; Miller the nose and the fbers of zygomaticus minor, zygomati-
1952; Gibbs 1999; Diogo et al. 2010, 2012, 2013a,b, 2017). cus major, and levator labii superioris could not be dif-
It may occasionally fuse with zygomaticus minor, forming ferentiated (Aziz 1979, 1981). Similarly, in a female fetus
a muscle mass that decussates with nearby muscles (Gibbs with trisomy 18, zygomaticus major, zygomaticus minor,
1999). Zygomaticus major may be divided into two bundles levator labii superioris, levator labii superioris alaeque
in gibbons, gorillas, and common chimpanzees (Deniker nasi, and levator anguli oris were fused bilaterally and
1885; Sonntag 1923; Raven 1950; Burrows et al. 2006; formed a single muscle sheet that extended from the zygo-
Diogo et al. 2010, 2012, 2013a). matic bone and inferior margin of orbicularis oculi to the
upper lip (Alghamdi et al. 2018). These authors describe
Variations this muscle sheet as the “complex triangle.” Individuals
Description with trisomy 21 also show poor differentiation of the mid-
Zygomaticus major may fuse with zygomaticus minor or face muscles, as the zygomaticus muscles and levator labii
risorius (Macalister 1875; Bergman et al. 1988; Watanabe superioris are not distinct and are replaced by a muscle
sheet that extends between orbicularis oculi and orbicu- Prevalence

laris oris (Bersu 1980). Macalister (1875) states that levator labii superioris may be
fused with levator labii superioris alaeque nasi in about one
Prevalence out of every three subjects. Sato (1968a) found levator labii
superioris present in 230 out of 240 facial sides (95.83%) in
Replacement of distinct zygomaticus muscles and levator
Kyushu-Japanese males and present in 158 out of 162 sides
labii superioris with a single sheet of muscle was found in fve
(97.53%) in females.
out of fve (100%) individuals with trisomy 21 (Bersu 1980).
Anomalies
Clinical Implications Description
Understanding variations of zygomaticus major is impor- In an infant with mandibulofacial dysostosis, levator labii
tant for planning and performing facial reanimation sur- superioris was absent and replaced with connective tissue
gery (Hu et al. 2008). (Herring et al. 1979). In a fetus with craniorachischisis, leva-
tor labii superioris was absent on the right side (Alghamdi
et al. 2017). It was also absent in a neonate with trisomy 13
levatOr labii superiOris (figure 2.1) (Aziz 1980). Bersu and Ramirez-Castro (1977) noted that in
infants with trisomy 18, levator labii superioris and levator
Synonyms labii superioris alaeque nasi were more extensively fused
This muscle may also be referred to as levator labii superi- than usual. In two neonates with trisomy 18, a continuous
oris proprius (Macalister 1875). sheet of muscle fbers extended from the cheek to the nose
and the fbers of zygomaticus minor, zygomaticus major,
Typical Presentation and levator labii superioris could not be differentiated (Aziz
Description 1979, 1981). Similarly, in a female fetus with trisomy 18,
zygomaticus major, zygomaticus minor, levator labii superi-
Levator labii superioris originates from the maxilla and
oris, levator labii superioris alaeque nasi, and levator anguli
zygomatic bone and inserts into the muscles of the upper
oris were fused bilaterally and formed a single muscle sheet
lip (Standring 2016).
that extended from the zygomatic bone and inferior margin
of orbicularis oculi to the upper lip (Alghamdi et al. 2018).
Innervation These authors describe this muscle sheet as the “complex
Levator labii superioris is innervated by the buccal and triangle.” Individuals with trisomy 21 also show poor dif-
zygomatic branches of the facial nerve (Standring 2016). ferentiation of the midface muscles, as the zygomaticus
muscles and levator labii superioris are not distinct and are
Comparative Anatomy replaced by a muscle sheet that extends between orbicularis
Levator labii superioris has a similar typical presentation oculi and orbicularis oris (Bersu 1980).
in the great apes, extending from the infraorbital region
to the upper lip (Miller 1952; Gibbs 1999; Diogo et al. Prevalence
2010, 2013a,b, 2017). In gibbons, the muscle runs pos- Replacement of distinct zygomaticus muscles and levator
teroanteriorly and mediolaterally from the infraorbital labii superioris with a single sheet of muscle was found in
region to the nose, with only a few fbers attaching near fve out of fve (100%) individuals with trisomy 21 (Bersu
the upper lip, thus having a presentation similar to that 1980).
found in non-catarrhine primates (Deniker 1885; Diogo
et al. 2012). Clinical Implications
N/A
Variations
Description levatOr labii superiOris alaeque nasi (figure 2.1)
Levator labii superioris may be fused with levator labii
superioris alaeque nasi (Macalister 1875). It may also be Synonyms
fused in rare cases with zygomaticus minor (Macalister This muscle may also be referred to as levator alae nasi
1875). It may send a slip to nasalis or have a connec- (Pessa et al. 1998).
tion to levator anguli oris (Macalister 1875; Watanabe
2016). Levator labii superioris may be doubled or tripled Typical Presentation
(Macalister 1875). It may also have two or three heads, with Description
the outer head arising from the zygomatic bone or orbicu- Levator labii superioris alaeque nasi originates from the fron-
laris oculi (Macalister 1875; Knott 1883a; Watanabe 2016). tal process of the maxilla (Standring 2016). Its medial slip
Levator labii superioris may be absent (Macalister 1875; inserts into the nose, on the perichondrium, and skin over the
Sato 1968a; Watanabe 2016). lateral wall of the major alar cartilage (Standring 2016). Its
lateral slip ends in the upper lip where it merges with levator (1977) noted that in infants with trisomy 18, levator labii
labii superioris and orbicularis oris (Standring 2016). superioris and levator labii superioris alaeque nasi were
more extensively fused than usual.
Innervation In a fetus with trisomy 18 and cyclopia, levator labii
superioris alaeque nasi seemed to originate from the max-
Levator labii superioris alaeque nasi is innervated by the
illa and insert into orbicularis oris (Smith et al. 2015).
superior buccal and zygomatic branches of the facial nerve
In a female fetus with trisomy 18, zygomaticus major,
(Standring 2016).
zygomaticus minor, levator labii superioris, levator labii
Comparative Anatomy superioris alaeque nasi, and levator anguli oris were fused
bilaterally and formed a single muscle sheet that extended
Levator labii superioris alaeque nasi has a similar typical pre-
from the zygomatic bone and inferior margin of orbicularis
sentation in the apes, extending from the maxilla and region
oculi to the upper lip (Alghamdi et al. 2018). These authors
of the medial palpebral ligament to the ala of the nose and
describe this muscle sheet as the “complex triangle.”
upper lip (Gratiolet and Alix 1866; Miller 1952; Gibbs 1999;
Burrows et al. 2006; Diogo et al. 2010, 2012, 2013a,b, 2017).
Prevalence
Variations N/A
Description
Levator labii superioris alaeque nasi may only have one
N/A
slip (Macalister 1875). The medial slip may be doubled or
separated from the lateral slip (Macalister 1875; Watanabe
2016). Levator labii superioris alaeque nasi may be fused levatOr anguli Oris (figure 2.1)
with levator labii superioris (Macalister 1875). It may
receive a slip from frontalis or procerus (Macalister 1875; Synonyms
Watanabe 2016). It may send fbers to nasalis (Macalister This muscle may also be referred to as musculus caninus
1875; Hur et al. 2010). It may also be absent (Sato 1968a; (Macalister 1875) or levator anguli oris facialis (Diogo and
Watanabe 2016). Abdala 2010).
This muscle is associated with anomalous nasi, also
known as the musculus anomalous of Albinus or the rhom- Typical Presentation
boideus of Santorini (Macalister 1875; Standring 2016).
Description
It originates from the frontal process of the maxilla and
ends near the canine fossa (Macalister 1875; Standring Levator anguli oris originates from the canine fossa of the
2016). Its origin may connect to the medial slip of levator maxilla and inserts into the angle of the mouth, where it
labii superioris alaeque nasi, and its insertion may fuse with blends with other muscles that contribute to the modiolus
levator anguli oris (Macalister 1875). (Standring 2016).
Innervation
Prevalence
Levator anguli oris is innervated by the buccal and zygo-
Macalister (1875) states that levator labii superioris alaeque
matic branches of the facial nerve (Standring 2016).
nasi may be fused with levator labii superioris in about one
out of every three subjects. Sato (1968a) found levator labii Comparative Anatomy
superioris alaeque nasi present in 217 out of 230 facial sides
Levator anguli oris has a similar typical presentation in the
(94.35%) in Kyushu-Japanese males and present in 175 out
apes, extending from the canine fossa to the angle of the
of 184 sides (95.11%) in females. In a sample of 40 cadavers,
mouth, often blending with nearby muscles (Raven 1950;
Hur et al. (2010) found that the medial slip of levator labii
Miller 1952; Gibbs 1999; Diogo et al. 2010, 2012, 2013a,b,
superioris alaeque nasi sent fbers to the transverse part of
2017).
nasalis in 36 cases (90%).
Variations
Anomalies
Description
Description
Levator anguli oris may be divided (Macalister 1875). It
Levator labii superioris alaeque nasi may be absent in indi- may be connected with nasalis (Macalister 1875). It may be
viduals with cleft lip defects (Standring 2016). In an infant continuous with levator labii superioris or depressor anguli
with mandibulofacial dysostosis, levator labii superioris oris (Macalister 1875; Watanabe 2016).
alaeque nasi inserted onto the wing of the nose but not the
upper lip (Herring et al. 1979). In a fetus with craniorachis-
chisis, levator labii superioris alaeque nasi was absent bilat- Prevalence
erally (Alghamdi et al. 2017). Bersu and Ramirez-Castro N/A
Anomalies platysma, fascia over the mastoid process, and/or the zygo-
Description matic arch (Macalister 1875; Knott 1883a; Standring 2016;
Watanabe 2016). The muscle may also arise from the skin
Levator anguli oris was absent bilaterally in a fetus with
over the superior third of sternocleidomastoid or the exter-
craniorachischisis (Alghamdi et al. 2017). In a female fetus
nal ear (Macalister 1875; Knott 1883a; Watanabe 2016). It
with trisomy 18, zygomaticus major, zygomaticus minor,
can also originate from the masseter tendon (Macalister
levator labii superioris, levator labii superioris alaeque
1875; Eisler 1912; Sekine et al. 1988). Risorius may receive
nasi, and levator anguli oris were fused bilaterally and
a slip from or fuse with transversus nuchae (Macalister
formed a single muscle sheet that extended from the zygo-
1875; Knott 1883a; Jovanovski et al. 2020). It may send a
matic bone and inferior margin of orbicularis oculi to the
slip to zygomaticus minor or fuse with zygomaticus major
upper lip (Alghamdi et al. 2018). These authors describe
(Macalister 1875; Watanabe 2016). It may blend with orbi-
this muscle sheet as the “complex triangle.” In an infant
cularis oris (Knott 1883a). At its insertion, it may join with
with mandibulofacial dysostosis, levator anguli oris origi-
depressor anguli oris (Macalister 1875; Sekine et al. 1988;
nated medially, near the medial corner of the eye (Herring
Kim et al. 2015a; Watanabe 2016).
et al. 1979).
Some authors have classified risorius into three types
Prevalence depending on its orientation and direction of insertion:
platysma risorius (inserts into the modiolus horizontally),
N/A
zygomaticus risorius (inserts into the modiolus from above
Clinical Implications horizontal), and triangularis risorius (inserts into the
modiolus from below horizontal) (Lightoller 1925; Loth
N/A
1931; Sekine et al. 1988; Kim et al. 2015a). According to
Kim et al. (2015a), the layer at which risorius inserts into
Risorius (Figure 2.1) the modiolus can also be classified into three types, rang-
ing from superficial to, flush with, or deep to depressor
Synonyms anguli oris.
This muscle may also be referred to as risorius of Santorini
(Macalister 1875). Prevalence
Sato (1968a) found risorius absent in 39 out of 386 facial sides
Typical Presentation (10.1%) in Kyushu-Japanese males and absent in 32 out of
Description 240 sides in females (13.3%). Ito et al. (2006) found risorius
Risorius originates from one or more structures in the absent in 4 out of 10 cadavers (40%). Pessa et al. (1998) found
region near the parotid gland and attaches to the modiolus risorius present in only 3 out of 50 (6%) hemifacial cadaver
(Standring 2016). dissections. Farahvash et al. (2010) found risorius present
in 16 out of 52 (30.8%) hemifacial cadaver dissections. In a
Innervation sample of 80 cadavers, Kim et al. (2015a) found platysma
Risorius is innervated by the buccal branches of the facial risorius in 36 cases (45%), triangularis risorius in 28 cases
nerve (Standring 2016). (35%), and zygomaticus risorius in 16 cases (20%). The inser-
tion level of risorius was superficial in 45 cases (56.3%), flush
Comparative Anatomy in 25 cases (31.2%), and deep in 10 cases (12.5%).
Risorius is typically absent in gibbons (Diogo et al. 2012),
orangutans (Diogo et al. 2013b), and bonobos (Miller 1952; Anomalies
Diogo et al. 2017). It is often present in gorillas (Deniker
Description
1885; Ruge 1887; Raven 1950; Seiler 1976; Diogo et al.
2010) and in common chimpanzees (Sonntag 1923; Seiler Risorius has been found absent bilaterally in both a neonate
1976; Burrows et al. 2006; Diogo et al. 2013a). When pres- with trisomy 18 (Aziz 1979) and a neonate with trisomy
ent, it extends from the superior margin of platysma to the 13 (Aziz 1980). In a fetus with craniorachischisis, the right
angle of the mouth (Gibbs 1999; Diogo et al. 2010, 2013a). risorius was absent (Alghamdi et al. 2017). In individuals
with trisomy 21, Bersu (1980) found that platysma cervicale
merged with risorius upon insertion. In an infant with man-
Variations
dibulofacial dysostosis, risorius originated from the fascia
Description over the masseter and was distinguished completely from
Risorius may present as a wide fan of fibers or one or more platysma (Herring et al. 1979).
fascicles (Macalister 1875; Bergman et al. 1988; Sekine et al.
1988; Standring 2016). Risorius may also be absent (Sato Prevalence
1968a; Bergman et al. 1988; Pessa et al. 1998; Farahvash et In their literature review, Smith et al. (2015) found that riso-
al. 2010; Watanabe 2016). Typical origins of risorius include rius was absent in 1 out of 20 individuals with trisomy 13
the parotid fascia, fascia over the masseter, fascia over (5%) and 1 out of 17 individuals with trisomy 18 (5.9%).
Clinical Implications Anomalies

As risorius can cover different parts of the masseter depend- Description
ing on its attachments, injection of botulinum toxin type Bersu et al. (1976) describe a male infant with Hanhart syn-
A into the masseter may affect risorius (Bae et al. 2014). drome. The buccinator muscles were comprised of fibers
Vector of attachment (platysma risorius, zygomaticus riso- that attached to the lateral surface of the maxilla with some
rius, or triangularis risorius) may affect the function of riso- fibers blended with the anomalous portion of the superior
rius and how it acts on the lips (Kim et al. 2015a). pharyngeal constrictor. In a fetus with craniorachischisis,
both buccinator and masseter were undifferentiated muscle
Buccinatorius (buccinator) (Figure 2.1) tissues situated medial to the ears (Alghamdi et al. 2017).
Synonyms
N/A Prevalence
N/A
Description Clinical Implications
Buccinator originates from the maxilla, mandible, and the The fourth band of buccinator may affect the alveolar bone
pterygomandibular raphe and inserts into the modiolus of the mandible or occlusion during muscular movements
(Standring 2016). (Hur et al. 2011a).
Innervation Orbicularis oris (Figure 2.1)

Buccinator is innervated by the buccal branch of the facial
nerve (Standring 2016). See also: Incisivus labii superior, Incisivus labii inferior
Comparative Anatomy Synonyms

Buccinator has a similar typical presentation in the apes, Some authors have described parts of orbicularis oris as
extending from the maxilla, mandible, and pterygoman- separate muscles: protractor labii superioris, protractor labii
dibular raphe to the angle of the mouth and lips (Raven inferioris, co-angustator labii inferioris, constrictor labii infe-
1950; Miller 1952; Gibbs 1999; Diogo et al. 2010, 2012, rioris, constrictor labii superioris, levator prolabii superioris,
2013a,b, 2017). and nasalis labii superioris (Macalister 1875; Knott 1883a).
Variations
Description
Description
Orbicularis oris encircles the mouth and is comprised of
Buccinator may vary in the extent of its maxillary and man- four parts that each contain a pars peripheralis (super-
dibular attachments (Macalister 1875; Hur et al. 2011a). An ficial or extrinsic) and pars marginalis (deep or intrinsic)
inferior bundle (fourth band) of the buccinator may be pres- (Standring 2016). The fibers of orbicularis oris originate
ent, which extends from the mandible and passes under from the modiolus and receive contributions from buccina-
orbicularis oris, ending beyond the modiolus (Hur et al. tor, levator anguli oris, and depressor anguli oris (Standring
2011a). Macalister (1875) reports that it may be thin and 2016). The fibers have submucosal, fascial, and dermal
underdeveloped in its center. It may be bilaminar (Bergman attachments around the lips (Standring 2016).
et al. 1988). It may fuse with masseter or connect to the
buccopharyngeal part of the superior pharyngeal constric- Innervation
tor (Macalister 1875; Bergman et al. 1988). It may receive a Orbicularis oris is innervated by the buccal and mandibular
slip from tensor veli palatini or the parotid duct (Macalister branches of the facial nerve (Standring 2016).
1875; Knott 1883a; Patel and Loukas 2016).
Ullah and Khan (2006) report the presence of an acces- Comparative Anatomy
sory muscle in the infratemporal fossa that they term the Orbicularis oris has a similar typical presentation in the
zygobuccinator muscle, which originated from the posterior apes, but it is possible that this muscle is not differentiated
surface of the frontal process of the zygomatic bone and into peripheral and marginal portions (Diogo et al. 2010,
inserted into the buccinator. This muscle likely corresponds 2012, 2013a,b, 2017).
to the temporobuccinator band described by Gaughran
(1957) (Buck 2007). Variations
Description
Prevalence Orbicularis oris may vary in its development and in the
In a sample of 40 hemifaces, Hur et al. (2011a) found a amount and extent of accessory bands from the alveolar
fourth band of the buccinator on 14 sides (35%). arches (Macalister 1875). Orbicularis oris is associated
with incisivus labii superioris and incisivus labii inferioris, Comparative Anatomy
two bundles that are considered accessory bundles of the Depressor anguli oris has a similar typical presentation in the
orbicularis oris complex (see the entries for these muscles) apes, extending from the angle of the mouth to the fascia of
(Standring 2016; Iwanaga et al. 2017). platysma, with connections to nearby muscles (Miller 1952;
Gibbs 1999; Diogo et al. 2010, 2012, 2013a,b, 2017). It may
Prevalence be differentiated into two or more bundles in gorillas (Ruge
N/A 1887; Diogo et al. 2010). It is fused with platysma cervicale
and orbicularis oris in bonobos (Miller 1952; Diogo et al.
2017). Transversus menti has been noted as a variation in
Anomalies common chimpanzees (Gratiolet and Alix 1866; Loth 1931;
Description Diogo et al. 2013a) and in one fetal gorilla (Deniker 1885).
Fibers of orbicularis oris are affected by cleft lip defects
(Bersu et al. 1977; De Mey et al. 1989; Wijayaweera Variations
et al. 2000). In incomplete clefts, the intrinsic portion of Description
the muscle ends in the submucosa of the vermilion bor-
Depressor anguli oris may not always have a continuous
der of the lip while the extrinsic portion crosses the cleft
origin from platysma and can originate entirely from the
and is distorted (De Mey et al. 1989; Wijayaweera et al.
mandible (Macalister 1875; Watanabe 2016). It may be split
2000). In complete clefts, the intrinsic portion is again
by the facial artery into three bundles (Macalister 1875).
interrupted, and fibers of the extrinsic portion are hori-
Its fibers may be continuous with levator labii superioris or
zontally oriented on the medial side and deviate along
zygomaticus major (Macalister 1875; Watanabe 2016). Its
the margin of the cleft on the lateral side (De Mey et al.
medial fibers may pass deep to depressor labii inferioris,
1989; Wijayaweera et al. 2000). In an infant with trisomy
enabling connections to incisivus labii inferioris, mentalis,
18 and a bilateral cleft lip, the fibers of orbicularis oris
and/or the inferior border of the mandible (Hur et al. 2014).
ended at the medial border of each maxillary process
It may be absent in some cases (Sato 1968a).
(Bersu et al. 1977).
Transversus menti is a variation that occurs when fibers
In a neonate with trisomy 13, orbicularis oris was
of depressor anguli oris cross the midline and interdigi-
absent (Aziz 1980). In a fetus with trisomy 18 and cyclo-
tate with those of its counterpart below the mental sym-
pia, the inferior part of orbicularis oris was comprised of
physis (Hallett 1848; Macalister 1875; Sato 1968a; Weaver
only the deep (marginal) portion (Smith et al. 2015).
1978; Bergman et al. 1988; Sripanidkulchai et al. 2013;
Standring 2016; Watanabe 2016). It may also be referred to
Prevalence as transversalis menti (Santorini), faisceau sous-symphysien
N/A (Cruveilhier), or the mental sling (Macalister 1875; Knott
1883a; Standring 2016). When transversus menti is present,
Clinical Implications its fibers may interdigitate with fibers of platysma, such that
N/A they may be mistaken for fibers of platysma that cross the
midline (Hallett 1848).
Prevalence
Depressor anguli oris (Figure 2.1)
Hur et al. (2014) studied depressor anguli oris in 40 hemi-
Synonyms faces from cadavers. Medial fibers of depressor anguli oris
This muscle may also be referred to as triangularis (Huber that pass deep to depressor labii inferioris were found in
and Hughson 1926). 18 specimens (45%). These fibers connected to incisivus
labii inferioris and the inferior border of the mandible in
14 specimens (35%), connected to incisivus labii inferioris
Typical Presentation and fibers of mentalis in two specimens (5%), connected to
Description incisivus labii inferioris only in one specimen (2.5%), and
Depressor anguli oris is continuous with platysma, origi- blended with mentalis only in one specimen (2.5%).
nating from the mental tubercle and from along the oblique Sato (1968a) found depressor anguli oris present in 366
line of the mandible (Standring 2016). It ends at the angle of out of 372 facial sides (98.39%) in Kyushu-Japanese males
the mouth, having connections to risorius, orbicularis oris, and present in 230 out of 240 sides (95.83%) in females.
and levator anguli oris (Standring 2016). Sato (1968a) found transversus menti present in 206 out
of 330 facial sides (62.42%) in Kyushu-Japanese males
Innervation and present in 116 out of 206 sides (56.31%) in females.
Depressor anguli oris is innervated by the marginal man- Knott (1883a) found transversus menti in 3 out of 11 cases
dibular branch and the buccal branch of the facial nerve (27.3%). Weaver (1978) found transversus menti in 9 out of
(Standring 2016). 16 (56.25%) cadaveric heads.
Anomalies Variations
Description
Description
Depressor labii inferioris may be divided into two or more
In a fetus with trisomy 18 and cyclopia, depressor anguli fascicles (Macalister 1875).
oris was partially fused with mentalis and depressor labii
inferioris (Smith et al. 2015). It also received fibers from Prevalence
platysma (Smith et al. 2015). In descriptions of other speci- N/A
mens with trisomy 18, both Bersu and Ramirez-Castro
(1977) and Aziz (1979) noted that depressor anguli oris was
Anomalies
poorly differentiated and partially blended with depressor
labii inferioris and platysma. Description
Congenital hypoplasia or agenesis of depressor anguli In a fetus with trisomy 18 and cyclopia, depressor labii
oris is a primary cause of asymmetric crying facies (ACF) inferioris was partially fused with mentalis and depressor
(Pasick et al. 2013; Akcan et al. 2016; Watanabe 2016). This anguli oris (Smith et al. 2015). In the female fetus with tri-
anomaly is often associated with other congenital anoma- somy 18 dissected by Alghamdi et al. (2018), depressor labii
lies throughout the body and has been observed in individu- inferioris was fused with mentalis bilaterally. In descrip-
als with 22q11.2 deletion syndrome and infants born after tions of other specimens with trisomy 18, both Bersu and
in vitro fertilization (Alexiou et al. 1976; Pasick et al. 2013; Ramirez-Castro (1977) and Aziz (1979) noted that depres-
Akcan et al. 2016; Watanabe 2016). sor labii inferioris was poorly differentiated and partially
blended with depressor anguli oris and platysma. In a fetus
Prevalence with craniorachischisis, depressor labii inferioris was also
Alexiou et al. (1976) found hypoplasia of depressor anguli poorly differentiated, primarily connected to fat-like tissue,
oris in 44 out of 6487 neonates (0.68%). and had only a few muscle fibers on the anterior surface of
the chin (Alghamdi et al. 2017).
Prevalence
Understanding variations in depressor anguli oris is help-
N/A
ful for administering botulinum toxin type A therapies
(Hur et al. 2008, 2014). Weaver (1978) suggests that trans- Clinical Implications
versus menti could be used in reconstructive surgery in
N/A
cases where depressor anguli oris and/or depressor labii
inferioris are damaged.
Procerus (Figure 2.1)
Synonyms
Depressor labii inferioris (Figure 2.1)
This muscle may also be referred to as pyramidalis nasi
Synonyms (Macalister 1875).
N/A
Description Procerus originates from the nasal bone, the lateral nasal
Depressor labii inferioris originates from the oblique line cartilage, and the transverse portion of nasalis (Standring
of the mandible and inserts into the mucosa and skin of the 2016). It inserts into the skin of the glabella and is partially
lower lip (Standring 2016). It is continuous with platysma, blended with occipitofrontalis (Standring 2016).
merges with orbicularis oris, and decussates with its coun-
terpart (Standring 2016). Innervation
Procerus is typically innervated by the lower zygomatic
Innervation and temporal branches of the facial nerve but may be inner-
Depressor labii inferioris is innervated by the mandibular vated by the buccal branch of the facial nerve in some cases
branch of the facial nerve (Standring 2016). (Standring 2016).
Comparative Anatomy Comparative Anatomy

Depressor labii inferioris has a similar typical presentation Procerus has a similar typical presentation in the apes,
in the apes, extending from the mandible and platysma to extending from frontalis to the dorsomedial region of the
the lower lip and blending with its counterpart and other nose, blending with frontalis and/or levator labii superioris
nearby muscles (Deniker 1885; Sonntag 1923; Raven 1950; alaeque nasi (Gratiolet and Alix 1866; Miller 1952; Seiler
Miller 1952; Seiler 1976; Pellatt 1979; Gibbs 1999; Burrows 1976; Pellatt 1979; Gibbs 1999; Burrows et al. 2006; Diogo
et al. 2006; Diogo et al. 2010, 2012, 2013a,b, 2017). et al. 2010, 2012, 2013a,b, 2017).
The portion of procerus closest to the orbit or the entire Nasalis may continue into procerus (Bergman et al. 1988;
muscle may be absent (Macalister 1875; Bergman et al. Watanabe 2016). The transverse part of nasalis may be
1988; Watanabe 2016). The procerus muscles may present split into fascicles or upper and lower bands (Macalister
as a single slip that continues onto the bridge of the nose 1875). The transverse parts may present as a single sheet
(Macalister 1875; Watanabe 2016). In rare cases, it may be of muscle (Macalister 1875). The muscle may be poorly
continuous with the inner border of levator labii superioris developed or absent (Macalister 1875). It may be fused
alaeque nasi (Macalister 1875). It may be separate from with levator anguli oris (Macalister 1875). The transverse
occipitofrontalis (Macalister 1875). part may receive fibers from levator labii superioris alae-
que nasi (Macalister 1875; Hur et al. 2010). It may also
Prevalence have an origin from the upper half of the alar facial crease
N/A (Hur et al. 2010). The alar part of nasalis may be absent
(Macalister 1875). It may merge with a slip of orbicularis
Anomalies oris (Macalister 1875).
Description
Procerus may be absent in individuals with cleft lip defects Prevalence
(Standring 2016). Procerus was absent bilaterally in a In a sample of 40 cadavers, Hur et al. (2010) found that
fetus with trisomy 18 and cyclopia (Smith et al. 2015). In the medial slip of levator labii superioris alaeque nasi sent
a fetus with craniorachischisis, procerus was represented fibers to the transverse part of nasalis in 36 cases (90%).
by a thin sheet of fibers between the eyes (Alghamdi The transverse part of nasalis originated from the maxilla
et al. 2017). only in 26 cases (65%) and from both the maxilla and the
alar facial crease in 14 cases (35%).
Prevalence
N/A
Anomalies
Description
N/A
Nasalis may be absent in individuals with cleft lip defects
(Standring 2016). In some specimens with trisomy 18, nasa-
Nasalis (Figure 2.1) lis was not identifiable (Bersu and Ramirez-Castro 1977).
Synonyms In a fetus with trisomy 18 and cyclopia, a muscle possibly
corresponding to nasalis was found on the superior por-
The transverse part of nasalis may be referred to as compres-
tion of the maxilla, situated where the nose typically lies
sor naris or compressor nasi (Macalister 1875; Standring
(Smith et al. 2015). It originated from the lateral maxilla
2016). The alar part of nasalis may be referred to as dilator
and inserted along the midline prominence below the
naris posterior (Macalister 1875; Standring 2016).
orbit, with a partial insertion into levator labii superioris
Typical Presentation alaeque nasi (Smith et al. 2015). Similarly, in a fetus with
craniorachischisis, a possible nasalis was identified on the
Description
left side spanning the region between the nose and mouth
The transverse part of nasalis originates from the maxilla (Alghamdi et al. 2017).
superior and lateral to the incisive fossa and merges with its
counterpart along the bridge of the nose (Standring 2016).
Prevalence
The alar part of nasalis originates from the maxilla superior
to the lateral incisor and canine teeth and inserts into the Nasalis was not identifiable in two out of four cases of tri-
skin and cartilage of the ala (Standring 2016). somy 18 (50%) (Bersu and Ramirez-Castro 1977).
Innervation Clinical Implications

Nasalis is typically innervated by the buccal branch of the N/A
facial nerve but may also be innervated by the zygomatic
branch in some cases (Standring 2016). Depressor septi nasi (Figure 2.1)
Nasalis has a similar typical presentation in the apes, This muscle may be referred to as depressor septi (Standring
extending from the maxilla to the lateral margin and ala of 2016), depressor septi mobilis narium, nasolabialis, depres-
the nose (Raven 1950; Miller 1952; Gibbs 1999; Diogo et al. sor apicis naris, or nasalis labii superioris (Macalister 1875;
2010, 2012, 2013a,b, 2017). Knott 1883a).

Description N/A
Depressor septi nasi may be considered a septal attach-
ment of orbicularis oris (Knott 1883a). It originates from Clinical Implications
the maxilla above the incisors and from orbicularis oris and Hypertrophy of depressor septi nasi may present as a swell-
inserts into the columella, medial crus of the major alar car- ing or mass of the columella (Ohtsuka 2005).
tilage, and nasal septum (Standring 2016).
incisivus labii superiOris (figure 2.1)
Innervation
Synonyms
Depressor septi nasi is typically innervated by the buccal
branch of the facial nerve but may be innervated by the This muscle may be referred to as cuspidator oris or labia-
zygomatic branch in some cases (Standring 2016). lis superior profundus (Seiler 1976; Diogo et al. 2010, 2012,
2013a,b, 2017), accessory fasciculi or skeletal head of orbi-
Comparative Anatomy cularis oris, facial portion of orbicularis oris, incisive bundle
Depressor septi nasi has a similar typical presentation of orbicularis oris, or incisive Cowper (Iwanaga et al. 2017).
in the apes, situated deep to orbicularis oris and extend-
ing from the maxilla to the inferior region of the nose Typical Presentation
(Raven 1950; Burrows et al. 2006; Diogo et al. 2010, 2012, This muscle is present only as a variation or anomaly.
2013a,b, 2017).
Comparative Anatomy
Variations Incisivus labii superioris has a similar typical presentation
Description in the apes, extending between the incisive fossa of the max-
illa and orbicularis oris (Seiler 1976; Iwanaga et al. 2021).
Depressor septi nasi may be absent or poorly developed
(Rohrich et al. 2000; Ebrahimi et al. 2012; Standring 2016;
Kikuta et al. 2020). It may also be hypertrophic (Ohtsuka Variations
2005; Watanabe 2016). It may be indistinguishable from Description
orbicularis oris (Macalister 1875). The origin from the Incisivus labii superioris is considered an accessory mus-
maxilla may be absent (Rohrich et al. 2000; Ebrahimi et al. cle of orbicularis oris (Iwanaga et al. 2017). It arises deep
2012; Kikuta et al. 2020). Kikuta et al. (2020) found that it to the superior portion of orbicularis oris, below the alar
originated from orbicularis oris and incisivus labii superi- portion of nasalis, lateral to depressor septi, and medial
oris, without an origin directly from the maxilla. Slips may to levator anguli oris (Lightoller 1925; Standring 2016;
extend between the medial crura and the tip of the nose Iwanaga et al. 2017; Hur 2018). It originates from the inci-
(Standring 2016). sive fossa of the maxilla above the eminence of the lat-
eral incisor (Standring 2016). Its bony attachment extends
Prevalence mediolaterally between the areas above the central incisor
In a sample of 55 cadavers, Rohrich et al. (2000) found and the canine and superoinferiorly between the attach-
that depressor septi nasi originated completely from orbi- ment of nasalis and the mucogingival junction (Iwanaga
cularis oris in 34 cases (61.8%). It originated from the peri- et al. 2017). Its fbers course laterally, interdigitate with
osteum of the maxilla and partially from orbicularis oris orbicularis oris, and divide into superfcial and deep parts
in 12 cases (21.8%). Depressor septi nasi was rudimentary that attach to the modiolus (Lightoller 1925; Standring
or absent in nine cases (16.4%). In a sample of 82 mus- 2016; Iwanaga et al. 2017). Hur (2018) notes that it may
cles from 41 cadavers, Ebrahimi et al. (2012) found that not divide into superfcial and deep parts in some cases.
depressor septi nasi originated from the periosteum of the It blends with other muscles at the angle of the mouth,
maxilla in 36 cases (44%) and from orbicularis oris in 32 particularly levator anguli oris (Lightoller 1925; Standring
cases (39%). The muscles were diminutive or foating and 2016; Iwanaga et al. 2017; Hur 2018).
comprised of fbrous tissue in 14 cases (17%). In a sample
of 20 sides, Kikuta et al. (2020) found depressor septi nasi
Innervation
present on 18 sides (90%). On 6 of these 18 sides (33%),
depressor septi nasi was diffcult to separate completely N/A
from orbicularis oris.
Prevalence
Anomalies In a sample of 52 facial sides, Hur (2018) found incisi-
Description vus labii superioris present in all cases (100%). It divided
Depressor septi nasi may be absent in individuals with cleft into superfcial and deep parts upon insertion in 48 cases
lip defects (Standring 2016). (92.3%) and did not divide in four cases (7.7%). In a sample
of 20 facial sides, Kikuta et al. (2020) found incisivus labii intermingled with the upper lateral portion of mentalis in
superioris present in all cases (100%). all cases (100%) and with the lower or middle portions of
mentalis in 22 cases (55%). In a sample of 40 hemifaces,
Anomalies Hur et al. (2014) fbers of depressor anguli oris connected to
incisivus labii inferioris and the inferior border of the man-
Description
dible in 14 specimens (35%), connected to incisivus labii
In an adult cadaver with an incomplete isolated cleft palate inferioris and fbers of mentalis in two specimens (5%), and
on the right maxilla, incisivus labii superioris was incom- connected to incisivus labii inferioris only in one specimen
pletely divided by two fstulae (Kikuta et al. 2019). (2.5%).
Anomalies
Prevalence
N/A
N/A
Clinical Implications N/A
Understanding the anatomy of incisivus labii superioris is
important for planning and performing surgical procedures Mentalis (figure 2.1)
and injections in the upper lip region (Iwanaga et al. 2017).
Synonyms
incisivus labii inferiOris (figure 2.1) This muscle may also be referred to as levator menti
Synonyms (Macalister 1873).
This muscle may be referred to as labialis inferior profun-
dus (Seiler 1976; Diogo et al. 2010, 2012, 2013a,b, 2017).

This muscle is present only as a variation. Mentalis originates from the incisive fossa of the mandible
and inserts into the skin of the chin (Standring 2016).
Comparative Anatomy
Incisivus labii inferioris has a similar typical presentation Innervation
in the apes, extending from the incisive fossa of the mandi- Mentalis is innervated by the marginal mandibular branch
ble to orbicularis oris and the skin of the chin region (Seiler of the facial nerve (Standring 2016).
1976; Iwanaga et al. 2021).
Comparative Anatomy
Variations Mentalis has a similar typical presentation in the apes,
Description descending from the region of the anterior teeth into the
Incisivus labii inferioris is considered an accessory muscle skin at the inferior edge of the mandible (Raven 1950;
of orbicularis oris situated lateral to mentalis (Hur et al. Gibbs 1999; Diogo et al. 2010, 2012, 2013a, 2017). In all
2011a; Standring 2016). It originates from the incisive fossa apes, it may interdigitate with its counterpart on the other
of the mandible below the eminence of the lateral incisor side of the body (Lightoller 1928; Raven 1950; Gibbs 1999;
(Hur et al. 2011a; Standring 2016). Its fbers course later- Diogo et al. 2010, 2012, 2013a,b, 2017). It may also blend
ally, interdigitate with orbicularis oris, and divide into with platysma or depressor labii inferioris in some cases
superfcial and deep bundles that attach to the modiolus (Gibbs 1999; Diogo et al. 2012, 2013a).
(Lightoller 1925; Hur et al. 2011a; Standring 2016). It may
also have connections to buccinator, mentalis, or depressor Variations
anguli oris (Hur et al. 2011a, 2013, 2014). Description
Mentalis may be connected with platysma (Bergman et al.
Innervation 1988; Lee and Yang 2016). It may be split into two bundles
N/A (Watanabe 2016). The direction of its lateral fbers may
vary (Hur et al. 2013). It may have connections to orbicu-
Prevalence laris oris, depressor anguli oris, or incisivus labii inferioris
In a sample of 40 hemifaces, Hur et al. (2011a) found that (Hur et al. 2013, 2014).
incisivus labii inferioris was present on 39 sides (97.5%).
It blended with mentalis on 10 sides (25%), merged into Prevalence
buccinator on 36 sides (90%), and merged into orbicularis In a sample of 40 hemifaces, Hur et al. (2013) found that
oris on three sides (7.5%). In a sample of 40 hemifaces, Hur the lateral fbers of mentalis coursed inferomedially in
et al. (2013) found that fbers of incisivus labii inferioris 38 cases (95%) and inferolaterally in two cases (5%).
The upper fbers of mentalis interlaced with the inferior Variations

border of orbicularis oris in all cases (100%). Fibers of Description
incisivus labii inferioris intermingled with the upper lateral
The development of platysma may vary, presenting in some
portion of mentalis in all cases (100%) and with the lower
cases as a well-developed sheet of muscle and in other cases
or middle portions of mentalis in 22 cases (55%). In a sam-
as multiple detached bundles (Macalister 1875; Bergman
ple of 40 hemifaces, Hur et al. (2014) found that fbers of
et al. 1988; Standring 2016; Lee and Yang 2016; Watanabe
depressor anguli oris connected to incisivus labii inferioris
2016). It may also be absent unilaterally, bilaterally, or in
and fbers of mentalis in two specimens (5%) and blended
the lower half only (Macalister 1875; Sato 1968b; Bergman
with mentalis only in one specimen (2.5%).
et al. 1988; Lee and Yang 2016). It may have a high origin
directly from the clavicle or a low origin from the level of
Anomalies the third or fourth rib (Macalister 1875).
Description Platysma may have connections to risorius, mentalis,
In a fetus with trisomy 18 and cyclopia, mentalis was par- depressor anguli oris, depressor labii inferioris, zygomati-
tially fused with depressor anguli oris and depressor labii cus minor, zygomaticus major, auricularis anterior, orbicu-
inferioris (Smith et al. 2015). In a female fetus with trisomy laris oculi, or transversus nuchae (Macalister 1875; Bergman
18, mentalis was fused with depressor labii inferioris bilat- et al. 1988; Bakkum and Miller 2016; Lee and Yang 2016;
erally (Alghamdi et al. 2018). In a fetus with craniorachis- Watanabe 2016). Its fbers can extend to the mastoid process
chisis, mentalis presented as a thin muscle layer deep to or the concha of the auricle (Macalister 1875). Some fbers
a poorly differentiated depressor labii inferioris (Alghamdi can also join with serratus anterior, infraspinatus, latissimus
et al. 2017). dorsi, or trapezius (Macalister 1875). It may connect via slips
to the occipital bone, the skin of the forehead, the midline
Prevalence of the back of the neck, the skin over the upper lobes of
N/A the mammary gland, the tendon of latissimus dorsi, or the
skin over deltoideus (Macalister 1875; Lee and Yang 2016;
Clinical Implications Watanabe 2016). The degree to which the fbers of both pla-
N/A tysma muscles interlace may vary (Mori 1964; de Castro
1980; Kim et al. 2001; Lee and Yang 2016; Watanabe 2016).
When transversus menti is present, its fbers may interdigi-
platysMa MyOiDes (platysMa) (figure 2.1) tate with fbers of platysma, such that they may be mistaken
for fbers of platysma that cross the midline (Hallett 1848).
Synonyms Sphincter colli profundus is a variable sheet of muscle
N/A that may originate from the clavicle deep to platysma and
insert into the fascia of the auricle (Bergman et al. 1988;
Typical Presentation Lee and Yang 2016). Beattie and Horsfall (1930) describe
Description a variant muscle situated immediately below the skin that
Platysma originates from the fascia over pectoralis major and originated from the posterior border of the mandible, the
deltoideus (Standring 2016). It courses over the clavicle and sheath of the masseter, and the sheath of the medial ptery-
along the anterior aspect of the neck to connect to the lower goid muscle. This muscle inserted into the superfcial fascia
border of the mandible, skin and subcutaneous tissue of the and skin over the mastoid. They conclude that this muscle
lower face, the lower lip, and the muscles near the mouth was derived from sphincter colli profundus.
(Standring 2016). Some fbers decussate across the midline
(Bergman et al. 1988; Standring 2016; Watanabe 2016). Prevalence
Sato (1968b) found platysma absent on 2 out of 168 sides
Innervation
(1.19%) in Kyushu-Japanese females. In a sample of 128
Platysma is innervated by the cervical branch of the facial cadavers, Mori (1964) found that the lower border of pla-
nerve (Standring 2016). tysma ended on the frst intercostal space in 76 cadavers
(59.4%) and on the second intercostal space in 52 cadavers
Comparative Anatomy (40.6%). The fbers of the platysma muscles did not inter-
Platysma has a similar typical presentation in the apes, sect in 16 cadavers (12.5%). The fbers of the platysma mus-
arising from the pectoral fascia and inserting onto the man- cles intersected under the chin in 84 cadavers (65.6%), in
dible and near the mouth (Diogo et al. 2010, 2012, 2013a, the neck above the laryngeal prominence in three cadavers
b, 2017). Sphincter colli profundus is typically not present (2.3%), in the neck below the laryngeal prominence in 19
in the apes, but Diogo et al. (2017) identify some fbers cadavers (14.8%), and at the sternum in six cadavers (4.6%).
in one adolescent bonobo that run ventrally from the ear de Castro (1980) studied platysma in 50 cadavers. Fibers
region and may correspond to a vestigial auricular portion of the platysma muscles interlaced just below the chin
of sphincter colli profundus. (within 1–2 cm) in 75% of cadavers and at the level of the
thyroid cartilage in 15% of cadavers. The fbers did not and Blatt 1966), occipital platysma (Gasser 1967), occipita-
interlace in the remaining 10% of cases. Kim et al. (2001) lis minor or the occipital transverse muscle (Bergman et al.
studied platysma in 70 Korean cadavers. Fibers of the pla- 1988), or the querer Halsmuskel (Bakkum and Miller 2016).
tysma muscles interlaced just below the mandibular border
(within 20 mm) in 30 cadavers (42.8%) and interlaced more
than 20 mm from the border in 30 cadavers (42.8%), The Typical Presentation
fbers did not interlace in the remaining 10 cases (14.3%). In This muscle is only present as a variation or anomaly.
29 cadavers (41.4%), both the left and right fbers interlaced
with each other. In 20 cases (28.6%), the right platysma Comparative Anatomy
fbers covered the left fbers. In 11 cadavers (15.7%), the left Platysma cervicale is often present in adult nonhuman mam-
platysma fbers covered the right fbers. mals (Smith et al. 2015). Among nonhuman apes, platysma
cervicale is only present as a distinct muscle in gibbons and
orangutans, sometimes absent in gorillas, and typically absent
Anomalies in common chimpanzees and bonobos. When the muscle
Description is present in common chimpanzees, bonobos, and gorillas,
In a female anencephalic fetus, platysma was abnormally it is often diminutive and resembles the presentation of this
well-developed and extended along the surface of the thorax muscle in humans (Diogo et al. 2010, 2012, 2013a,b, 2017).
(Windle 1893). In a fetus with craniorachischisis, platysma
was a thin layer of fbers that passed distally from the latero- Variations
inferior borders of the mouth (Alghamdi et al. 2017). On the
Description
left side of one infant with trisomy 13, sternocleidomastoid
sent slips to platysma (Pettersen et al. 1979). In a neonate with Platysma cervicale is present in normal human develop-
trisomy 13, platysma was comprised of two small bundles ment, as the occipital lamina that gives rise to this muscle
on both sides of the chin that merged anteriorly (Aziz 1980). is established during the sixth week of embryonic growth
Summarizing four cases of trisomy 18, Bersu and Ramirez- (Lewis 1910; Sataloff and Selber 2003). It typically dis-
Castro (1977) found that the facial muscles were poorly differ- appears in early embryonic developmental stages and is
entiated and depressor anguli oris, depressor labii inferioris, therefore often absent in normal adult humans (Futamura
and platysma were more fused than usual at the angle of the 1906; Lewis 1910; Gasser 1967; Smith et al. 2015). Gasser
mouth. In another infant with trisomy 18, platysma originated (1967) notes that between CR58 and CR80 mm (crown-
above the clavicle bilaterally (Aziz 1979). In a fetus with tri- rump length of 58–80 mm) “occipital platysma” is present
somy 18 and cyclopia, platysma had a strong fascial connec- as a distinct band that runs between the platysma muscle
tion to sternocleidomastoid and sent fbers to zygomaticus and the occipital region, but this muscle was not identi-
major and depressor anguli oris (Smith et al. 2015). fed in a CR 142 mm fetus. Gasser (1967) also observes the
occasional presence of “transversus nuchae” laying dor-
sal to auricularis posterior in the lower occipital or upper
Prevalence
cervical region between CR58 and CR80 mm. Later, these
In their literature review, Smith et al. (2015) found that pla- muscles were still observed in 210 mm and 270 fetuses and
tysma was anomalous in 2 out of 20 (10%) individuals with were blended with auricularis posterior (Gasser 1967).
trisomy 13 (left and right sides separated, absent cervical When present, platysma cervicale often extends from
portion). In 1 out of 17 individuals with trisomy 18 (5.9%), the region of auricularis posterior to the external occipi-
platysma originated above the clavicles. tal protuberance or superior nuchal line (Macalister 1875;
Clinical Implications Knott 1883a; Blodget and Blatt 1966; Sato 1968a; Bergman
et al. 1988; Bakkum and Miller 2016; Standring 2016;
N/A
Watanabe 2016). It may connect with auricularis posterior,
auricularis superior, splenius capitis, or sternocleidomas-
toid (Macalister 1875; Knott 1883a; Blodget and Blatt 1966;
platysMa cervicale (figure 2.3)
Bergman et al. 1988; Bakkum and Miller 2016; Standring
Entry adapted by permission from Springer Nature Customer 2016). Blodget and Blatt (1966) describe a “pencil-shaped”
Service Centre GmbH: Springer Current Molecular Biology transversalis nuchae muscle that arises from the surface
Reports, Muscles Lost in Our Adult Primate Ancestors of splenius capitis and inserts onto the anterior border of
Still Imprint in Us: on Muscle Evolution, Development, sternocleidomastoid.
Variations, and Pathologies. E. Boyle, V. Mahon, R. Diogo, Hallett (1848) describes what is likely platysma cervi-
2020. cale as a variation of the lower fascicle of auricularis pos-
terior. Hallett (1848) states that the lower fascicle formed
Synonyms a round-bellied muscle that attached to the occipital pro-
This muscle is also referred to as the transversus nuchae tuberance and superior nuchal line at one end and the
(Futamura 1906; Lewis 1910), transversalis nuchae (Blodget concha auris at the other end. The muscle was sheathed
and situated between the tendons of occipitofrontalis and the mastoid process, or it can originate from the corner of the
sternocleidomastoid. mouth and/or parotid fascia and insert with trapezius onto the
Its presentation can also resemble the condition seen in occipital bone (Bersu and Ramirez-Castro 1977; Colacino and
anomalous cases, in which the muscle can originate from Pettersen 1978; Pettersen 1979; Pettersen et al. 1979; Bersu
near the mouth or parotid fascia to insert near the occipital 1980; Aziz 1981; Urban and Bersu 1987; Smith et al. 2015).
region (Wood 1864; Schmidt 1982; Bergman et al. 1988; It was also present bilaterally in both a female fetus with
Lei et al. 2010; Bakkum and Miller 2016; Jovanovski triploidy (Moen et al. 1984) and a male neonate with Meckel
et al. 2020). Wood (1864) notes that it may extend from the syndrome (Pettersen 1984). Mieden (1982) describes an
parotid fascia and risorius to trapezius and occipitofronta- infant with median cleft lip, hypotelorism, and alobar holo-
lis. Jovanovski et al. (2020) found that transversus nuchae prosencephaly. Transversus nuchae was present on the left
fused with risorius bilaterally. Lei et al. (2010) observed side, extending over trapezius in between the occipital pro-
fibers extending to the zygomatic region. Platysma cervi- tuberance and the posterior border of sternocleidomastoid
cale may have two bellies (Schmidt 1982; Lei et al. 2010; (Mieden 1982).
Jovanovski et al. 2020).
Prevalence
Innervation In their literature review, Smith et al. (2015) found that pla-
Platysma cervicale may be innervated by a branch of the tysma cervicale was present in 5 out of 20 individuals with
facial nerve (Blodget and Blatt 1966; Jovanovski et al. trisomy 13 (25%), 13 out of 17 individuals with trisomy 18
2020), or from anastomoses of facial nerve branches and (76.5%), and 5 out of 5 individuals with trisomy 21 (100%).
great auricular nerve branches (Schmidt 1982).
Platysma cervicale may present as a painful swelling or
Prevalence
mass in the neck (Blodget and Blatt 1966).
The variation described by Hallett (1848) was observed in 3
out of 105 subjects (2.9%). In a sample of 28 bodies, Knott
(1883a) found transversus nuchae bilaterally in five bodies MUSCLES OF MASTICATION
(17.9%) and unilaterally in two bodies (7.1%). Sato (1968a)
Masseter (Figure 2.2)
found that transversus nuchae was present in 105 out of 300
facial sides (35%) in Kyushu-Japanese males and present in Synonyms
62 out of 196 sides (31.6%) in females. Bergman et al. (1988) N/A
cite studies that state that “occipitalis minor” is present fre-
quently in Malaysian individuals, present in 56% of Black Typical Presentation
individuals, present in 50% of Japanese individuals, pres-
ent in 36% of European individuals, and typically absent in Description
Khoisan peoples and Melanesians. Lei et al. (2010) found Masseter is comprised of three layers (Standring 2016). The
“transversus nuchae” on 12 out of 20 sides (60%) from a total superficial layer originates from the maxillary process of
sample of 10 cadavers (seven male and three female). The the zygomatic bone and from the zygomatic arch (Standring
muscle was absent in the females. Standring (2016) suggests 2016). The middle and deep layers also originate from the
that “transversus nuchae” is present in 25% of individuals. zygomatic arch (Standring 2016). The superficial layer inserts
Watanabe et al. (2017) found “transversus nuchae” in 40 onto the lower mandibular ramus and angle of the mandible,
out of 124 sides (32.2%) from a total sample of 62 cadavers the middle layer inserts into the middle of the mandibular
(present in 26 cadavers, bilaterally in 14 and unilaterally in 12). ramus, and the deep layer inserts into the upper portion of the
Watanabe et al. (2017) observed that this muscle mandibular ramus and the coronoid process (Standring 2016).
extended from the external occipital protuberance to insert
onto the mastoid process (43% of cases) or originated from
Innervation
the external occipital protuberance and curved around the
mastoid process to join with platysma (58% of cases). Masseter is innervated by the masseteric nerve, a branch
of the anterior division of the mandibular nerve (Standring
2016).
Anomalies
Description Comparative Anatomy
Platysma cervicale frequently presents as an anomaly in Masseter has a similar typical presentation in the apes,
individuals with trisomy 13, 18, or 21. When present, pla- extending from the zygomatic arch to the mandible (Raven
tysma cervicale is a discrete bundle of the platysma com- 1950; Miller 1952; Gibbs 1999; Diogo et al. 2010, 2012,
plex that extends from the mouth to the nuchal region (Smith 2013a,b, 2017).
et al. 2015). It can either originate from the posterior pla- Masseter can send fibers to the tendon of temporalis in
tysma myoides and insert with sternocleidomastoideus onto gorillas (Diogo et al. 2010) and to the medial pterygoid in
FIGURE 2.2 Muscles of mastication in lateral view. Masseter is only partially shown.
gibbons (Diogo et al. 2012). The superfcial and deep por- buccinator (Macalister 1875). The deep layer of the masse-
tions are well separated in chimpanzees (Sonntag 1923; ter may originate in part from the temporomandibular liga-
Miller 1952), while there is no strong fascia or aponeuro- ment (Macalister 1875). There is no consensus regarding
sis between these layers in the other apes (Sonntag 1924; whether masseter sends fbers to the articular disc of the
Boyer 1939; Gibbs 1999; Diogo et al. 2010, 2012, 2013b). temporomandibular joint (Standring 2016). Bergman et al.
Zygomatico-mandibularis—an accessory muscle superf- (1988) note that both the masseter and temporalis muscles
cial to temporalis that extends from the deep surface of the may send fbers to this articular disc.
zygomatic process to the coronoid process of the mandi-
ble—has been found as an accessory bundle of the masseter Prevalence
in common chimpanzees (Gratiolet and Alix 1866; Starck
N/A
1973; Göllner 1982; Diogo et al. 2013a) and in a few orang-
utans (Boyer 1939; Saban 1968) and gorillas (Göllner 1982). Anomalies
Description
Variations
The presentation of the masseter muscle is often affected by
Description congenital malformations, cephalic disorders, and various
Macalister (1875) notes that the muscles of mastication have syndromes. Macalister (1875) notes that Dumeril found the
few variations aside from individual variations in size. Most masseter muscles absent in a specimen with phocomelia.
authors describe masseter as having a superfcial and deep The masseter can be underdeveloped in cases of hemifacial
head (e.g., Macalister 1875; Bergman et al. 1988; Watanabe microsomia (e.g., Takashima et al. 2003) or hypertrophic in
2016), not the three-layered confguration deemed typical cases of congenital hemifacial hyperplasia (e.g., Rončević
by Standring (2016). Connections between the muscles of 1986; Tsuneki et al. 2019).
mastication may occur and refect these muscles’ shared On the right side of a male infant with Hanhart syn-
origin from the same muscular mass (Bergman et al. 1988). drome, Bersu et al. (1976) found that the right mandible
For example, the deep layer of the masseter may send fbers was represented by a small bone. The fbers of the masse-
to temporalis and masseter may receive a slip from the ter were oriented obliquely downward and posteriorly and
medial pterygoid (Macalister 1875; Knott 1883a; Bergman had a restricted insertion on the angle of the mandible. In
et al. 1988; Watanabe 2016). Masseter may also fuse with an infant with mandibulofacial dysostosis, there was one
superfcial layer and three deeper layers of the masseter Variations

muscle (Herring et al. 1979). In a neonate with trisomy 13, Description
the right anterior belly of the digastric was quadrupled, with
Macalister (1875) notes that the muscles of mastication
the fourth belly inserting onto the masseter (Aziz 1980).
have few variations aside from individual variations in
Mieden (1982) describes two male fetuses with cyclo-
size. Temporalis can vary in the extent of its origin on the
pia and alobar holoprosencephaly. In one specimen, the
cranium and its insertion along the mandible (Macalister
muscles of mastication were absent on the right side and
1875; Bergman et al. 1988; Watanabe 2016). Temporalis
temporalis and masseter were small on the left side. In
may send fbers to the articular disc of the temporoman-
an otocephalic fetus examined by Lawrence and Bersu
dibular joint (Zenker 1955; Bergman et al. 1988; Standring
(1984), the mandible was represented by two separate bony
2016). Connections between the muscles of mastication
masses located within the middle ear cavities. Due to this
may occur and refect these muscles’ shared origin from
anomaly, temporalis and masseter fused at the midline
the same muscular mass (Bergman et al. 1988). For exam-
into a muscle mass that formed the foor of the oral cavity.
ple, temporalis may receive fbers from the deep layer
In a fetus with craniorachischisis, Alghamdi et al. (2017)
of the masseter and temporalis may be connected to the
found that both masseter and buccinator were undifferen-
lateral pterygoid (Macalister 1875; Bergman et al. 1988;
tiated muscle tissues situated medial to the ears.
Watanabe 2016).
Temporalis minor was termed by Henle (1871) to refer to
Prevalence an accessory muscle that originated from the anterior por-
N/A tion of temporalis and inserted onto the mandibular notch
(Bergman et al. 1988; Watanabe 2016). Ullah and Khan
(2006) report the presence of an accessory muscle in the
infratemporal fossa that they term the zygobuccinator mus-
N/A cle, which originated from the posterior surface of the frontal
process of the zygomatic bone and inserted into the buccina-
tor. This muscle likely corresponds to the temporobuccinator
teMpOralis (figure 2.2) band described by Gaughran (1957) (Buck 2007).
Synonyms
Some authors have referred to the anteromedial/deep Prevalence
portion of temporalis as a separate muscle, sphenoman- N/A
dibularis (Dunn et al. 1996), but there is little evidence to
support designating this portion as an independent muscle Anomalies
(e.g., Türp et al. 1997; Schön Ybarra and Bauer 2001; Geers Description
et al. 2005; Akita et al. 2019). The presentation of the temporalis may be affected by con-
genital malformations and cephalic disorders. Temporalis
Typical Presentation can be underdeveloped in cases of hemifacial microsomia
Description (e.g., Takashima et al. 2003). Mieden (1982) describes two
Temporalis originates from the deep temporal fascia and male fetuses with cyclopia and alobar holoprosencephaly.
from the temporal fossa (Standring 2016). It ends in a ten- In one specimen, the muscles of mastication were absent on
don that inserts onto the coronoid process and anterior mar- the right side and temporalis and masseter were small on
gin of the mandibular ramus (Standring 2016). the left side. In an otocephalic fetus examined by Lawrence
and Bersu (1984), the mandible was represented by two
Innervation separate bony masses located within the middle ear cavi-
Temporalis is innervated by the deep temporal branches of the ties. Due to this anomaly, temporalis and masseter fused at
anterior division of the mandibular nerve (Standring 2016). the midline into a muscle mass that formed the foor of the
oral cavity. Temporalis also sent fbers to the bony masses
Comparative Anatomy that represented the coronoid process of each mandible. In a
Temporalis has a similar typical presentation in the apes, fetus with craniorachischisis, Alghamdi et al. (2017) found
extending from the temporal fossa and temporal fascia to that temporalis was absent on the right side and was repre-
the coronoid process and anterior margin of the mandibular sented by undifferentiated muscle tissue between the eye
ramus (Boyer 1939; Raven 1950; Miller 1952; Gibbs 1999; and the ear on the left side.
Diogo et al. 2010, 2012, 2013a,b, 2017). A “pars suprazy- Bersu et al. (1976) describe a male infant with Hanhart
gomatica” of the temporalis is typically present in many syndrome in which the right mandible was represented by
primates including several genera of monkeys (Diogo and a small bone. On the right side, the insertion of temporalis
Wood 2011) and has been observed in some gorillas (Raven was diffuse and most of the muscle attached to the coro-
1950; Göllner 1982; Diogo et al. 2010), common chimpan- noid process. Smaller portions of the muscle inserted onto
zees, and bonobos (Göllner 1982; Diogo et al. 2017). the articular mass and condylar process or blended with the
superior fascicle of pterygoideus lateralis. In an infant with size. The degree of separation between the upper and lower
mandibulofacial dysostosis, temporalis was divided into heads may vary (Macalister 1875; Bergman et al. 1988).
superfcial, middle, and deep parts (Herring et al. 1979). The lateral pterygoid may alternatively be comprised of
On both sides of a fetus with trisomy 18 and cyclopia, the one or three heads (Bertilsson and Ström 1995; Fujita et al.
insertion of temporalis reached the angle of the mandible 2001; Watanabe 2016). When present, the third head may
(Smith et al. 2015). be referred to as an inner or medial head that originates
from the greater wing of the sphenoid and inserts into the
Prevalence pterygoid fovea and/or the medial part of the temporoman-
N/A dibular joint disc-capsule complex (Bertilsson and Ström
1995; Fujita et al. 2001; Pompei Filho et al. 2009; Kiliç
Clinical Implications et al. 2010; Dergin et al. 2012; Antonopoulou et al. 2013).
Variations in the dimensions of the medial portion of tem- Connections between the muscles of mastication may
poralis may contribute to maxillary nerve compression, occur and refect these muscles’ shared origin from the
headaches, and tic douloureux (Schön Ybarra and Bauer same muscular mass (Bergman et al. 1988). For example,
2001). Temporalis hypertrophy may occur in tandem with the lateral pterygoid may join with temporalis or digastricus
masseter hypotrophy (e.g., Legg 1880; Graziano et al. 2016) (Macalister 1875; Bergman et al. 1988; Watanabe 2016). A
or without associated hypertrophy of the masseter (e.g., portion of the upper head may insert into the articular cap-
Wilson and Brown 1990; Wang et al. 2013; Watanabe 2016; sule and/or articular disc of the temporomandibular joint
Tuncel et al. 2017). (Bertilsson and Ström 1995; Naidoo 1996; Fujita et al. 2001;
Mazza et al. 2009; Kiliç et al. 2010; Dergin et al. 2012;
Antonopoulou et al. 2013; Standring 2016; Watanabe 2016).
pterygOiDeus lateralis (lateral The upper head may exclusively insert into the articular
pterygOiD) (figure 2.2) capsule and disc (Naidoo 1996; Mazza et al. 2009; Kiliç
et al. 2010; Dergin et al. 2012; Antonopoulou et al. 2013).
The lower head may also send fbers to the disc or joint
Synonyms
capsule (Naidoo 1996; Kiliç et al. 2010; Fujita et al. 2001).
This muscle may be referred to as pterygoideus externus The pterygoid muscles are associated with several
(Macalister 1875). accessory muscles. Sphenotemporalis was termed by Mack
Typical Presentation (1984) to describe a small muscle in the infratemporal fossa
situated superior and deep to the upper head of the lateral
Description pterygoid (see the entry for this muscle). The pterygospinous
The lateral pterygoid has two heads (Standring 2016). The or pterygospinsosus muscle extends between the posterior
upper head originates from the infratemporal crest of the margin of the lateral pterygoid plate and the spine of the
greater wing of the sphenoid, and the lower head originates sphenoid (see the entry for this muscle) (Macalister 1875;
from the lateral pterygoid plate of the sphenoid (Standring Poland 1890; Nathan 1989; von Lüdinghausen et al. 2006;
2016). The two heads merge and insert onto the pterygoid Watanabe 2016). Pterygoideus proprius extends between
fovea on the neck of the mandible (Standring 2016). the infratemporal crest and the tuber palati or the lateral
pterygoid plate (see the entry for this muscle) (Macalister
Innervation 1875; Poland 1890; Tubbs et al. 2007; Watanabe 2016).
The upper head and the lateral portion of the lower head
of lateral pterygoid are innervated by a branch of the buc- Prevalence
cal nerve (Standring 2016). The medial portion of the lower Bertilsson and Ström (1995) conducted a comprehensive
head is innervated by a branch of the anterior division of the review of 89 research articles on the anatomy of the lat-
mandibular nerve (Standring 2016). eral pterygoid. These authors found that 65% of the arti-
cles report lateral pterygoid having two heads, 20% of the
Comparative Anatomy articles report lateral pterygoid presenting as a single head,
The lateral pterygoid has a similar typical presentation in the and 15% of the articles report lateral pterygoid having three
apes, arising via two heads from the lateral pterygoid plate and heads. Furthermore, 60% of the articles report an insertion
sphenoid to the capsule of the temporomandibular joint and onto the mandible, articular disc, and joint capsule of the
the neck of the mandible (Gratiolet and Alix 1866; Sonntag temporomandibular joint; 30% report an insertion into the
1923, 1924; Boyer 1939; Raven 1950; Miller 1952; Wall et al. mandible with only a few fbers inserting into the disc; and
1994; Gibbs 1999; Diogo et al. 2010, 2012, 2013a,b, 2017). 10% found an insertion exclusively into the mandible.
Naidoo (1996) studied the insertions of the lateral ptery-
Variations goid in 40 cadaveric specimens. In 26 specimens (65%), the
Description upper head inserted into the pterygoid fovea, capsule, and
Macalister (1875) notes that the muscles of mastication disc. In 11 specimens, the upper head inserted into the con-
have few variations aside from individual variations in dyle only. In two cases, it inserted into the disc only (5%). In
one case, the lower head had an additional insertion into the Anomalies
meniscus (2.5%). Fujita et al. (2001) studied the lateral pter- Description
ygoid in 20 cadaveric specimens. Two heads of the muscle
Bersu et al. (1976) describe a male infant with Hanhart syn-
(upper and lower) were found in 13 specimens (65%), while
drome in which the right mandible was represented by a
seven specimens had a third inner head (35%). All 20 upper
small bone. On the right side, the pterygoid muscles were
heads (100%), all seven inner heads (100%), and two lower
partially fused and diffcult to separate from one another.
heads (10%) had an insertion into the disc-capsule complex
The well-developed superior fascicle of the lateral ptery-
of the temporomandibular joint.
goid received a contribution from temporalis and inserted
Mazza et al. (2009) studied the insertion of the upper
completely onto the temporomandibular articular mass.
head in 193 temporomandibular joint specimens. In 63
Fibers from the inferior fascicle extended beyond the con-
cases (32.6%), the upper head inserted via a single bundle
dylar process on its medial side and attached along the
onto the joint capsule and mandible. In 76 cases (39.4%)
medial aspect of the sphenomandibular ligament and to the
the upper head inserted via two bundles, one onto the
petrotympanic fssure.
disc and one onto the mandible. In 54 cases (28%), the
Mieden (1982) describes two male fetuses with cyclopia
upper head inserted via a single bundle onto the disc.
and alobar holoprosencephaly. In one specimen, the mus-
Pompei Filho et al. (2009) evaluated 178 MRI to deter-
cles of mastication were absent on the right side, and the
mine the prevalence of the third head of the lateral ptery-
medial and lateral pterygoid muscles were fused on the left
goid. The third head was found in 36 cases (20.2%), and
side. In the second specimen, the medial and lateral ptery-
these heads inserted entirely onto the disc of the tem-
goid muscles were fused bilaterally. In an otocephalic fetus
poromandibular joint.
examined by Lawrence and Bersu (1984), the mandible was
Kiliç et al. (2010) examined the lateral pterygoid in 49
represented by two separate bony masses located within the
temporomandibular joint specimens from 26 cadavers. Two
middle ear cavities. The lateral pterygoid was represented
heads were present in 32 specimens (65.3%), and three
by a fused muscle mass on both sides.
heads were present in 17 specimens (34.7%). In 18 speci-
In an infant with mandibulofacial dysostosis, the lateral
mens (36.7%), the upper head inserted into the disc-capsule
pterygoid had three heads of origin that joined to insert
complex and the mandible, and the lower head inserted
onto the anterior and lateral aspects of the medial condyle
into the mandible. In 14 specimens (28.6%), the upper head
(Herring et al. 1979). In a fetus with craniorachischisis,
inserted into the disc-capsule complex and the lower head
the lateral pterygoid muscles were missing on both sides
inserted into the mandible. In 13 specimens (26.5%), both
(Alghamdi et al. 2017).
the upper head and the lower head inserted into the mandi-
In one neonate with trisomy 13, an accessory muscle slip
ble only. In four specimens (8.2%), the upper head inserted
extended from the upper head of the lateral pterygoid to the
into the disc-capsule complex and the lower head inserted
lower part of the medial pterygoid near its insertion point
into the complex and into the mandible.
(Pettersen et al. 1979). On both sides of one neonate with tri-
Dergin et al. (2012) examined the lateral pterygoid in 98
somy 18, the lateral pterygoid inserted onto the deep aspect of
temporomandibular joint specimens from 49 patients using
the temporomandibular joint and adjacent part of the tempo-
MRI. In 29 cases (29.6%), the upper head inserted into the
ral bone, with no fbers extending to the mandible (Aziz 1979).
disc and the lower head inserted into the mandible. In 40
In a fetus with trisomy 18 and cyclopia, the lateral pterygoid
cases (40.8%), the upper head inserted into the disc and the
was mostly normal but had an origin that extended more supe-
mandible, and the lower head inserted into the mandible. In
riorly and posteriorly on the cranium (Smith et al. 2015).
29 cases (29.6%), the upper head inserted into the disc and
both the lower head and third head inserted into the man- Prevalence
dible. Antonopoulou et al. (2013) studied the attachment of In their literature review, Smith et al. (2015) found that a
the upper head of the lateral pterygoid in 36 sides from 18 slip from the lateral pterygoid to the medial pterygoid was
cadavers. These authors found that the upper head inserted only present in the case described by Pettersen et al. (1979),
into the pterygoid fovea and the disc-capsule complex of thus having a prevalence of 1 out of 20 individuals (5%)
the temporomandibular joint in 20 cases (55.5%). It inserted with trisomy 13. These authors also found that an insert
into the pterygoid fovea only in 10 cases (27.8%) and into exclusively onto the temporomandibular joint and adjacent
the disc-capsule complex only in six cases (16.7%). A third temporal bone was only observed in the case described by
medial head was found in eight cases (22.2%). Aziz (1979), thus having a prevalence of 1 out of 17 indi-
Stöckle et al. (2019) conducted a comprehensive review viduals (5.9%) with trisomy 18.
of 11 research articles on the anatomy of the lateral pter-
ygoid. In an overall sample of 521 subjects, the relative
frequency of one-headed lateral pterygoid muscles was Clinical Implications
between 7.7% and 26.7%, the relative frequency of two- It is unclear whether an insertion of the upper head entirely
headed lateral pterygoid muscles was between 61.4% and into the articular disc increases risk of anterior disc dis-
91.1%, and the relative frequency of three-headed lateral placement (Taskaya-Yilmaz et al. 2005; Mazza et al. 2009;
pterygoid muscles was between 4% and 35%. Dergin et al. 2012; Antonopoulou et al. 2013). Valenzuela
et al. (2020) suggest that cases where the upper belly inserts to the anterior margin of the carotid canal, medial to the
only into the disc and a third head is present may have the foramen spinosum and foramen ovale. The muscle coursed
most risk of developing symptoms related to temporoman- inferiorly and narrowed to an insertion onto the medial
dibular dysfunction. surface and/or the superior border of the medial pterygoid
muscle about a centimeter below its exit from the pterygoid
fossa. A small, independent accessory muscle bundle may
pterygOiDeus MeDialis (MeDial pterygOiD) (figure 2.2) be present with attachments below or posterior to the retro-
molar pad, or into the fascia of the mylohyoid muscle (Abe
Synonyms et al. 1997; Sakamoto and Akita 2004).
This muscle may be referred to as pterygoideus internus The pterygoid muscles are associated with several other
(Macalister 1875). accessory muscles. Sphenotemporalis was termed by Mack
(1984) to describe a small muscle in the infratemporal fossa
Typical Presentation situated superior and deep to the upper head of the lateral
pterygoid (see the entry for this muscle). The pterygospinous
Description or pterygospinsosus muscle extends between the posterior
The medial pterygoid has two heads (Standring 2016). The margin of the lateral pterygoid plate and the spine of the
deep head originates from the medial surface of the lat- sphenoid (see the entry for this muscle) (Macalister 1875;
eral pterygoid plate of the sphenoid (Standring 2016). The Poland 1890; Nathan 1989; von Lüdinghausen et al. 2006;
superfcial head originates from the maxillary tuberosity Watanabe 2016). Pterygoideus proprius extends between
and the pyramidal process of the palatine (Standring 2016). the infratemporal crest and the tuber palati or the lateral
It inserts onto the medial surface of the mandibular ramus pterygoid plate (see the entry for this muscle) (Macalister
and angle of the mandible (Standring 2016). 1875; Poland 1890; Penhall et al. 1998; Tubbs et al. 2007;
Watanabe 2016).
Innervation
The medial pterygoid is innervated by the medial pterygoid
Prevalence
branch of the mandibular nerve (Standring 2016).
In a sample of 39 head halves, Bhojwani et al. (2017) found
Comparative Anatomy that the deep head of the medial pterygoid originated from
The medial pterygoid has a similar typical presentation in the medial surface of the lateral pterygoid plate and the
the apes, originating from the medial surface of the lateral pterygoid fossa in seven specimens (17.9%). In 28 specimens
pterygoid plate and sometimes the pterygoid fossa to insert (71.8%), the origin from these two structures was expanded
onto the medial side of the mandible (Raven 1950; Miller onto the lateral surface of the medial pterygoid plate. In one
1952; Gibbs 1999; Diogo et al. 2010, 2012, 2013a,b, 2017). specimen (2.6%), the origin was from the medial pterygoid
It may also have origins from the pyramidal process of plate and the pterygoid fossa.
the palatine, the maxillary tuberosity, and/or the pterygo- In a sample of 84 medial pterygoid muscle specimens
mandibular raphe in orangutans (Boyer 1939; Gibbs 1999; from 42 cadavers, Abe et al. (1997) found a small, inde-
Diogo et al. 2013b). pendent accessory muscle bundle in ten specimens (11.9%)
from fve cadavers. The bundle inserted posterior to the
Variations retromolar pad in fve specimens (6%), below the retromo-
Description lar pad in three specimens (3.6%), and into the fascia of
mylohyoid in two specimens (2.4%). In a sample of 24 head
Macalister (1875) notes that the muscles of mastication
halves from 12 cadavers, Sakamoto and Akita (2004) found
have few variations aside from individual variations in size.
an accessory muscle bundle of the medial pterygoid in ten
Connections between the muscles of mastication may occur
head halves (41.7%) from fve cadavers.
and refect these muscles’ shared origin from the same mus-
cular mass (Bergman et al. 1988). For example, the medial
pterygoid may send a slip to the masseter (Bergman et al. Anomalies
1988; Watanabe 2016). It may also give origin to styloglos- Description
sus or send a slip to tensor veli palatini (Macalister 1875; Bersu et al. (1976) describe a male infant with Hanhart syn-
Bergman et al. 1988; Watanabe 2016). The medial pterygoid drome in which the right mandible was represented by a
may originate in part from the pterygoid fossa or the lateral small bone. On the right side, the pterygoid muscles were
surface of the medial pterygoid plate (Bhojwani et al. 2017). partially fused and diffcult to separate from one another.
An accessory medial pterygoid muscle or accessory bun- The attachments of the medial pterygoid were extensive,
dle of the muscle may be present (Koritzer and Suarez 1980; with some fbers inserting onto the angle of the mandible,
Abe et al. 1997; Sakamoto and Akita 2004). Koritzer and others blending with the lateral pterygoid fbers that joined
Suarez (1980) report the presence of an accessory medial with the sphenomandibular ligament, and other fbers
pterygoid muscle that had a broad origin spanning from passed into the neck deep to styloglossus and blended with
the posterior superior margin of the lateral pterygoid plate the tongue muscles and middle pharyngeal constrictor.
Mieden (1982) describes two male fetuses with cyclopia on the squamous portion of the temporal bone (Mack 1984;
and alobar holoprosencephaly. In one specimen, the mus- Watanabe 2016). Mack (1984) notes that sphenotemporalis
cles of mastication were absent on the right side, and the is distinct from the lateral pterygoid and should not be con-
medial and lateral pterygoid muscles were fused on the left sidered a slip of this muscle.
side. In the second specimen, the medial and lateral ptery-
goid muscles were fused bilaterally. In an infant with man- Innervation
dibulofacial dysostosis, the medial pterygoid arose from Sphenotemporalis is innervated by a nerve arising directly
the lower orbital bar and pterygoid process and inserted from the trigeminal ganglion (Mack 1984).
just above the mylohyoid line and onto the medial condyle
and the skull (Herring et al. 1979). In an otocephalic fetus Prevalence
examined by Lawrence and Bersu (1984), the mandible was Mack (1984) found sphenotemporalis in 2 out of 30 half
represented by two separate bony masses located within the heads (6.7%).
middle ear cavities. The medial pterygoid was represented
by a fused muscle mass on both sides.
In a fetus with craniorachischisis, the medial pterygoid Anomalies
was absent on the right side (Alghamdi et al. 2017). On the N/A
left side, it seemed to be represented by fbers that extended
between the maxilla and the mandible. In one neonate with Clinical Implications
trisomy 13, an accessory muscle slip extended from the upper N/A
head of the lateral pterygoid to the lower part of the medial
pterygoid near its insertion point (Pettersen et al. 1979).
pterygOspinOus (figure 2.2)
Prevalence
See also: Pterygoideus lateralis, Pterygoideus medialis
In their literature review, Smith et al. (2015) found that a slip
from the lateral pterygoid to the medial pterygoid was only Synonyms
present in the case described by Pettersen et al. (1979), thus This muscle was originally named the pterygospinosus mus-
having a prevalence of 1 out of 20 individuals (5%) with tri- cle (Thane) (Poland 1890). This muscle may also be referred
somy 13. to as pterygotympanicus (Saban) (Mack 1984). A variant of
pterygospinous is the pterygofascialis muscle (Poland 1890).
N/A Typical Presentation
This muscle is present only as a variation.
sphenOteMpOralis (figure 2.2)
Comparative Anatomy
Diogo et al. (2013a,b, 2017) note that pterygotympanicus
Synonyms has not been observed in the apes.
N/A
Variations
This muscle is present only as a variation. The pterygospinous muscle may replace the pterygospinous
ligament or course along the ligament (Macalister 1875;
Comparative Anatomy Poland 1890; Standring 2016; Watanabe 2016). When pres-
Mack (1984) states that it is possible that sphenotemporalis is a ent, it often replaces the pterygospinous ligament, originat-
vestigial remnant of a muscle situated between the alisphenoid ing from the posterior margin of the lateral pterygoid plate
and squamosal, which is found in some bats and wombats. between the two pterygoid muscles and inserting onto the
spine of the sphenoid (Macalister 1875; Poland 1890; Nathan
Variations 1989; von Lüdinghausen et al. 2006; Watanabe 2016). von
Description Lüdinghausen et al. (2006) also observed an additional
Sphenotemporalis was termed by Mack (1984) to describe insertion into the capsule and articular disc of the temporo-
a small muscle in the infratemporal fossa situated superior mandibular joint in some cases. These authors suggest that
and deep to the upper head of the lateral pterygoid. It is due to its innervation and location, it could be designated
attached to the cranium on either side of the sphenotem- as a third head of the lateral pterygoid. In cases where the
poral suture. It originated from the greater wing of the pterygospinous muscle is present with a fbrous or ossifed
sphenoid, with its attachment extending as far laterally as pterygospinous ligament, the upper attachment of the mus-
the infratemporal crest. The muscle coursed horizontally cle varies and is associated with an abnormal arrangement
to insert onto the anterior aspect of the articular eminence of the sphenomandibular ligament (Poland 1890). In some
cases, muscle may insert into the petrotympanic fissure and Variations
blend with the sphenomandibular ligament (Poland 1890). Description
Van Dongen (1968) observed that the pterygospinous
When present, the pterygoideus proprius (Henle 1858) typi-
muscle was present in a fetus at CR80 mm (crown-rump
cally originates from the infratemporal crest of the greater
length 80 mm). Mérida Velasco et al. (1994) examined this
wing of the sphenoid, courses superficially over the lateral
muscle in five human fetuses and found that it originated
pterygoid muscle, and inserts onto the lower part of pos-
from the posterior border of the lateral pterygoid plate and
terior edge of the lateral pterygoid plate (Wagstaffe 1871;
inserted into Meckel’s cartilage. These authors suggest that
Macalister 1875; Knott 1880, 1883a; Poland 1890; Eisler
it is a remnant of the masticatory muscle group.
1912; Barker 1981; Bergman et al. 1988; Penhall et al.
Poland (1890) considers pterygofascialis to be a variant
1998; Akita et al. 2001; Tubbs et al. 2007; Watanabe 2016).
of the pterygospinous muscle. In one case, pterygofascialis
Macalister (1875) describes it as a nearly vertical slip. The
arose from the sphenomandibular ligament and attached to
muscle may also originate from the anteromedial portion of
the posterior edge and outer aspect of the medial pterygoid
temporalis (Poland 1890; Akita et al. 2001). In some cases,
(Poland 1890; Watanabe 2016). In a second case, it had the
it inserts onto the tuber palati (Macalister 1875), either refer-
same origin and attached onto both the posterior edge and
ring to the pyramidal process of the palatine bone (Tubbs
outer aspect of the medial pterygoid and the lateral ptery-
et al. 2007) or the maxillary tuberosity (Knott 1880).
goid plate (Poland 1890). In these cases, the sphenoman-
Pterygoideus proprius may be entirely tendinous or
dibular ligament did not attach to the spine of the sphenoid
be comprised of both muscular and tendinous fibers
but to the petrotympanic fissure (Poland 1890).
(Wagstaffe 1871; Poland 1890; Tubbs et al. 2007;
Innervation Watanabe 2016). The posterior fibers of the muscle
Nathan (1989) observed that the pterygospinous muscle is may join with those of the medial pterygoid muscle
innervated by a branch of the inferior alveolar nerve. von (Wagstaffe 1871; Tubbs et al. 2007). The lateral ptery-
Lüdinghausen et al. (2006) observed that this muscle is goid may partially originate from the pterygoideus pro-
innervated by a small twig of the pterygoid branch of the prius (Wagstaffe 1871; Macalister 1875; Poland 1890;
mandibular nerve. Penhall et al. 1998; Watanabe 2016).
Penhall et al. (1998) propose three potential scenarios
Prevalence for the development of pterygoideus proprius. First, some
In a sample of ten sides from five fetuses, Mérida Velasco deep muscle fibers of the temporalis anlage may attach to
et al. (1994) found the pterygospinous muscle on eight the lateral pterygoid plate before temporalis attaches onto
sides (80%). In a sample of 54 cadaveric specimens, von the coronoid, thus persisting as a separate, vertical band of
Lüdinghausen et al. (2006) found a pterygospinous mus- fibers. Second, this muscle may be formed by fibers of the
cle in five cases (9.3%). In three cases, the muscle had an lateral pterygoid that fail to attach to the articular disc and/
attachment into the medial surface of the temporomandibu- or the mandible and elongate with growth into a vertical slip
lar joint capsule and articular disc. The muscle was pres- that retains an attachment to the sphenoid. Third, the muscle
ent alongside a pterygospinous osseous bar in one case and could be formed by fibers of lateral pterygoid that fail to join
alongside a pterygospinous ligament in two cases. the upper or lower head of this muscle during development.
Anomalies Innervation
N/A Akita et al. (2001) found that pterygoideus proprius is inner-
vated by twigs of the anterior deep temporal nerve.
Prevalence
When the pterygospinous muscle is present, it may affect
the distribution pattern of the mandibular nerve (von Knott (1883a) found three cases of pterygoideus proprius in
Lüdinghausen et al. 2006). a sample of 112 bodies (2.7%). Penhall et al. (1998) found
three cases of pterygoideus proprius in a sample of about
150 cadavers (~2%). Akita et al. (2001) found the muscle in
Pterygoideus proprius (Figure 2.2)
3 out of 66 head halves (3%).
Anomalies
Synonyms N/A
N/A
Typical Presentation Tubbs et al. (2007) suggest that pterygoideus proprius has
This muscle is present only as a variation. the potential to impinge upon the lingual or alveolar nerve
branches of the trigeminal nerve or compress the maxil-
Comparative Anatomy
lary artery.
N/A
EXTRINSIC MUSCLES OF THE EAR Variations

AND MIDDLE EAR MUSCLES Description
Auricularis anterior and superior may blend to form a con-
auricularis anteriOr (figure 2.3)
tinuous sheet of muscle (Macalister 1875; Knott 1883a).
Synonyms Auricularis anterior may originate from the zygomatic arch
This muscle may also be referred to as attrahens aurem and insert into the tragus, and in this case would be referred
(Macalister 1875). to as musculus auricularis anterior profundus (Macalister
1875; Knott 1883a). It may be doubled, with one part insert-
Typical Presentation ing into the helix and the other inserting into the concha
Description (Macalister 1875). Its lower border may connect to platysma
(Macalister 1875). Auricularis anterior may be absent more
Auricularis anterior originates from the epicranial aponeu-
often than the other extrinsic ear muscles (Macalister 1875;
rosis and attaches to the auricle at the spine of the helix
Bergman et al. 1988; Watanabe 2016). It may be reduced
(Standring 2006).
and comprised of a single bundle or numerous unconnected
Innervation fasciculi (Macalister 1875; Watanabe 2016).
Auricularis anterior is innervated by a temporal branch of Prevalence
the facial nerve (Standring 2016).
N/A
Comparative Anatomy
Anomalies
Diogo et al. (2010, 2012, 2013a,b, 2017) consider auricularis
anterior to be a bundle of the auriculo-orbitalis muscle pres- Description
ent in the apes, which extends from the anterior aspect of On the left side of a fetus with trisomy 18 and cyclopia,
the ear to the region of frontalis. auricularis anterior was present and partially fused with
auricularis superior and orbicularis oculi (Smith et al.
FIGURE 2.3 Extrinsic muscles of the ear and posterolateral neck muscles in lateral view.
2015). On the right side, auricularis anterior was not a dis- trisomy 18 (Aziz 1979). In a fetus with craniorachischisis,
tinct muscle. Auricularis anterior was absent bilaterally in auricularis superior was found only on the left side, present-
a neonate with trisomy 18 (Aziz 1979) and in a fetus with ing as a thin sheet of muscle extending between the supe-
craniorachischisis (Alghamdi et al. 2017). rior aspect of the ear and the superior side of the left eye
(Alghamdi et al. 2017). In an infant with mandibulofacial
Prevalence
dysostosis, auricularis superior had two bellies that fused
In their literature review, Smith et al. (2015) found that into an intermediate tendon (Herring et al. 1979). The ten-
auricularis anterior was absent in 1 out of 17 cases of tri- don became muscular again and the muscle ended in the
somy 18 (5.9%). fascia of the neck.
Prevalence
N/A
N/A
auricularis superiOr (figure 2.3) Clinical Implications
Synonyms N/A
This muscle may also be referred to as attollens aurem
(Macalister 1875). auricularis pOsteriOr (figure 2.3)
Typical Presentation Synonyms
Description This muscle may also be referred to as retrahens aurem
Auricularis superior originates from the epicranial apo- (Macalister 1875).
neurosis and attaches to the superior part of the auricle
(Standring 2006). Typical Presentation
Innervation Description
Auricularis superior is innervated by a temporal branch of Auricularis posterior originates from the mastoid portion of
the facial nerve (Standring 2016). the temporal bone and attaches to the auricle at the ponticu-
lus (Standring 2016).
Comparative Anatomy
Auricularis superior has a similar typical presentation in Innervation
the apes, extending from the epicranial aponeurosis to the Auricularis posterior is innervated by the posterior auricu-
superior aspect of the ear, with occasional origins from, lar branch of the facial nerve (Standring 2016).
or connections to, the temporal fascia in orangutans and
gorillas (Raven 1950; Miller 1952; Gibbs 1999; Diogo et al. Comparative Anatomy
2010, 2012, 2013a,b, 2017). Auricularis posterior is typically not present as a distinct
Variations muscle in orangutans (Ruge 1887; Sullivan and Osgood
1925; Lightoller 1928; Miller 1952; Diogo et al. 2013b). In
Description bonobos, it extends from the mastoid region of the temporal
Auricularis superior and anterior may blend to form a con- bone to the posterior aspect of the ear, as in humans (Gibbs
tinuous sheet of muscle (Macalister 1875; Knott 1883a). 1999; Miller 1952; Diogo et al. 2017). In gibbons, common
Auricularis superior may connect with transversus nuchae chimpanzees, and gorillas, it extends from the occipital
(Macalister 1875; Bergman et al. 1988). It may be divided region to the posterior aspect of the ear (Raven 1950; Gibbs
into multiple fascicles (Macalister 1875). It may have a 1999; Diogo et al. 2010, 2012, 2013a).
lower origin, from the temporal aponeurosis (Macalister
1875). Its fbers may not reach the auricle and instead end Variations
in the temporal aponeurosis (Macalister 1875). It may be
Description
absent in rare cases (Macalister 1875; Bergman et al. 1988;
Hoogbergen et al. 1996; Watanabe 2016). Auricularis posterior may be comprised of two or three fas-
cicles (Macalister 1875; Knott 1883a; Standring 2016). In
Prevalence rare cases, it may have four or more fascicles, or present as
N/A a continuous sheet of muscle (Macalister 1875). One of the
fascicles may be replaced by a tendinous band (Macalister
Anomalies 1875; Knott 1883a). The lower fascicle may arise from the
Description cervical fascia (Macalister 1875). Hallett (1848) describes
On the left side of a fetus with trisomy 18 and cyclopia, a variation of the lower fascicle that is likely platysma cer-
auricularis superior was present and partially fused with vicale or transversus nuchae (see the entry for this muscle).
auricularis anterior (Smith et al. 2015). Auricularis supe- Auricularis posterior may have connections to sterno-
rior presented as a thin sheet of muscle in a neonate with cleidomastoid, occipitofrontalis, platysma, or transversus
nuchae (Macalister 1875; Knott 1883a; Bergman et al. (Barsoumian et al. 1998; Abe et al. 2004; Rodríguez-
1988). It may be doubled (Macalister 1875). It may also Vázquez 2016; Standring 2016). Tensor tympani may be
be absent in some cases (Macalister 1875; Bergman et al. absent and replaced by a fbrous band (Kelemen 1943;
1988; Sato 1968a; Hoogbergen et al. 1996; Watanabe 2016). Rodríguez-Vázquez 2016).
Prevalence
Prevalence
Sato (1968a) found auricularis posterior present in 312 out
In a sample of 496 ears with normal development, Wright
and Etholm (1973) found that tensor tympani was split into
present in 194 out of 226 sides (85.84%) in females.
medial and lateral parts in one ear (0.2%).
Anomalies
Anomalies
Description
Description
In a fetus with craniorachischisis, auricularis posterior was
Tensor tympani has been noted as absent bilaterally in a
found only on the left side, presenting as a thin band of
fetus with craniorachischisis (Alghamdi et al. 2017) and
fbers extending from the posterior-superior aspect of the
in an otocephalic fetus (Lawrence and Bersu 1984). In one
ear to the posterior neural tissue (Alghamdi et al. 2017).
infant with trisomy 13–15 and another infant with trisomy
Auricularis posterior was absent in an infant with mandibu-
18, tensor tympani bifurcated into medial and lateral parts
lofacial dysostosis (Herring et al. 1979).
unilaterally (Wright and Etholm 1973). In both cases, the
medial part traveled with the facial nerve in the facial canal,
Prevalence to just above the oval window, where it exited the canal and
N/A blended with stapedius.
On the right side of a male infant with Hanhart syn-
Clinical Implications drome, some lateral pterygoid fbers attached to the pet-
N/A rotympanic fssure and formed a separate muscle bundle
that was about half the size of a typical tensor tympani
tensOr tyMpani (nOt illustrateD) (Bersu et al. 1976). It passed through the fssure and
inserted via a tendon onto the malleus. On the left side of
Synonyms an infant with CHARGE syndrome, the manubrium of the
N/A malleus was absent and tensor tympani instead attached
to the head of the malleus (Wright et al. 1986). In the right
Typical Presentation ear of a fetus with trisomy 18, tensor tympani was anoma-
Description lous and its tendon inserted onto one side of the malleus
Tensor tympani originates from the greater wing of the (Tadaki et al. 2003).
sphenoid, the pharyngotympanic tube, and from the bony
canal it occupies (Standring 2016). It inserts via a tendon Prevalence
onto the manubrium of the malleus (Standring 2016). In their literature review, Tadaki et al. (2003) found that
anomalies of tensor tympani or its tendon were found in
Innervation 5 out of 16 cases of trisomy 18 (31.25%). In a sample of
Tensor tympani is innervated by a branch of the trigeminal 13 temporal bones from children with the Pierre Robin
nerve (Standring 2016). sequence, Gruen et al. (2005) found that the middle ear
muscles were anomalous in fve bones.
Comparative Anatomy Clinical Implications
Tensor tympani has a similar typical presentation in the Bifurcation of tensor tympani into medial and lateral parts
apes, extending from the pharyngotympanic tube and adja- was found in an individual who also had chronic otitis
cent regions of the neurocranium to insert onto the malleus media (Wright and Etholm 1973).
(Diogo et al. 2010, 2012, 2013a,b, 2017).
stapeDius (nOt illustrateD)

Variations
Description Synonyms
Tensor tympani may be doubled (Macalister 1875; Le N/A
Double 1897; Rodríguez-Vázquez 2016). It may also be
split into lateral and medial parts, with the lateral belly Typical Presentation
having a typical presentation and the medial belly travel- Description
ing with the facial nerve in the facial canal (Wright and Stapedius originates from the pyramidal eminence of the
Etholm 1973; Rodríguez-Vázquez 2016). Some fbers of tympanic cavity and inserts via a tendon onto the neck of
tensor veli palatini may be continuous with tensor tympani the stapes (Standring 2016).
Innervation bones from children with the Pierre Robin sequence, Gruen
Stapedius is innervated by a branch of the facial nerve et al. (2005) found that the middle ear muscles were anoma-
(Standring 2016). lous in fve bones.

Information about stapedius in the apes is scarce, but it likely Ectopic muscle tissue in the middle ear has been found in
has a typical presentation similar to that in humans and individuals who also exhibit chronic otitis media, presbycu-
inserts on the stapes (Diogo et al. 2010, 2012, 2013a,b, 2017). sis, or sensorineural deafness (Wright and Etholm 1973). A
shortened stapedius tendon may cause conductive hearing
Variations loss (Zawawi et al. 2014).
Description
Stapedius may be doubled (Macalister 1875; Le Double MUSCLES OF THE TONGUE
1897; Wright and Etholm 1973; Bergman et al. 1988;
intrinsic Muscles Of the tOngue (nOt illustrateD)
Rodríguez-Vázquez 2016). The length of the stapedius ten-
don may vary (Hough 1958; Zawawi et al. 2014; Rodríguez- Synonyms
Vázquez 2016). Ectopic muscle tissue may be present in the N/A
middle ear and is often closely associated with the facial
nerve (Hoshino and Paparella 1971; Wright and Etholm Typical Presentation
1973; Rodríguez-Vázquez 2016). Hypoplasia or absence Description
of stapedius, and/or absence of its tendon, may occur There are four paired intrinsic muscles of the tongue (Standring
(Kelemen 1943; Hough 1958; Hoshino and Paparella 1971; 2016). The superior longitudinal muscle courses from the sub-
Wright and Etholm 1973; Bergman et al. 1988; Kopuz et al. mucous fbrous tissue of the posterior tongue and the medial
2006; Rodríguez-Vázquez et al. 2010; Rodríguez-Vázquez lingual septum to the margins of the tongue (Standring 2016).
2016; Dalmia and Behera 2017). The inferior longitudinal muscle courses from the root of the
tongue and the body of the hyoid to the apex of the tongue,
Prevalence
blending with styloglossus (Standring 2016). The transverse
Out of 500 middle ear surgeries, Hough (1958) found sta- muscle blends with palatopharyngeus as it extends from the
pedius completely absent in fve cases (1%). In a sample median fbrous septum to the submucous tissue at the lateral
of 195 temporal bones from 141 individuals, hypoplasia lingual margin (Standring 2016). The vertical muscle is situ-
of stapedius and absence of its tendon was found in two ated just under the superior longitudinal muscle with fbers
bones (1%) and ectopic muscle tissue was present in 12 that extend dorsoventrally (Standring 2016).
bones (6.2%) (Hoshino and Paparella 1971). In a sample of
496 ears with normal development, stapedius was doubled Innervation
in two ears (0.4%) and absent in two ears (0.4%), the sta- The intrinsic muscles of the tongue are innervated by the
pedius tendon was absent in four ears (0.8%), and ectopic hypoglossal nerve (Standring 2016).
muscle tissue was present in 18 ears (3.6%) (Wright and
Etholm 1973). Comparative Anatomy
N/A
Anomalies
Stapedius has been noted as absent bilaterally in a fetus Description
with craniorachischisis (Alghamdi et al. 2017) and in an Patel and Loukas (2016) note that there are no variations of
otocephalic fetus (Lawrence and Bersu 1984). In two the intrinsic muscles of the tongue. However, variable slips
infants with CHARGE syndrome, stapedius was absent associated with the extrinsic muscles may affect the exter-
bilaterally (Wright et al. 1986). In an infant with man- nal anatomy of the tongue (see the entries for these muscles
dibulofacial dysostosis, stapedius was located posterosu- below) (Macalister 1875; Patel and Loukas 2016).
perior to the jaw articulation (Herring et al. 1979). In the
right ear of a fetus with trisomy 18, stapedius lacked a Prevalence
tendon and connected to the head of the stapes via funicu- N/A
lar tissue (Tadaki et al. 2003). Wright and Etholm (1973)
found ectopic muscle tissue in the middle ear of an infant
Anomalies
with 13–15 trisomy.
Description
Prevalence Congenital malformations that affect the intrinsic muscles
In their literature review, Tadaki et al. (2003) found that of the tongue are often associated with other birth defects or
anomalies of stapedius or its tendon were found in 5 out of clinical syndromes and include aglossia (complete absence),
18 cases of trisomy 18 (27.8%). In a sample of 13 temporal microglossia or hypoglossia (reduction), and macroglossia
FIGURE 2.4 Muscles of the tongue in lateral view.
(enlargement), bifd tongue (anterior separation of the tongue Prevalence

along the midline), and an accessory tongue (Stevenson In a sample of 2301 patients from the United States,
2006; Kumar et al. 2009; Patel and Loukas 2016). Agrawal McCarthy (1941) found macroglossia in one individual. In
et al. (2014) describe an individual with a congenital trilobe a sample of 1,906 Chilean individuals, Witkop and Barros
tongue associated with partial cleft palate. (1963) found macroglossia in one individual (0.05%) and a
Other anomalies of the tongue include occupying a bifd tongue in two individuals (0.1%). In a sample of 956
position in the pharynx and an extended connection with African American patients, Schaumann et al. (1970) found
the mandible (Mieden 1982; Lawrence and Bersu 1984; macroglossia in one individual (0.1%) and a bifd tongue in
Pettersen 1984). Mieden (1982) describes two male fetuses another individual (0.1%).
with cyclopia and alobar holoprosencephaly. In one fetus,
the oral cavity was reduced and separated from the pharynx Clinical Implications
by the buccopharyngeal membrane. A small tongue flled
N/A
the pharynx and compressed the epiglottis. In an otoce-
phalic fetus examined by Lawrence and Bersu (1984), the
geniOglOssus (figure 2.4)
oral cavity was reduced and separated from the pharynx by
a membrane deemed to be the buccopharyngeal membrane. Synonyms
A small tongue was situated posterior to the membrane and This muscle may also be referred to as geniohyoglossus
flled the pharynx. The dorsal surface of the tongue faced (Macalister 1875).
the posterior pharyngeal wall, and the tip of the tongue was
oriented inferiorly. The tongue was connected via muscle Typical Presentation
fbers to the body of the hyoid. In a male neonate with
Meckel syndrome, Pettersen (1984) found that the anterior Description
two-thirds of the tongue were malformed, and the tongue Genioglossus originates from the superior mental spine on
was united with the mandible on the left side. The tongue the posterior surface of the mental symphysis (Standring
was very thin anteriorly along the midline. The posterior 2016). Its superior fbers insert along the length of the
third of the tongue was normal. ventral surface of the tongue and blend with the intrinsic
tongue muscles (Standring 2016). Its intermediate fbers contribute to ankyloglossia, or tongue-tie (Choi et al. 2011;
insert into the posterior tongue (Standring 2016). Its infe- Sharma et al. 2013; Patel and Loukas 2016).
rior fbers insert into the body of the hyoid and have an Bersu et al. (1976) describe a male infant with Hanhart
additional attachment into constrictor pharyngis medius syndrome in which the right mandible was represented by
(Standring 2016). Genioglossus muscles on each side of the a small bone. In this infant, the right genioglossus muscle
body decussate across the midline anteriorly (Standring was absent. It was replaced by a muscle mass that passed
2016). from the inferior margin of the right mandible to the deep
surface of the left hyoglossus muscle. On the left side of a
Innervation fetus with craniorachischisis, genioglossus was fused with
Genioglossus is innervated by the hypoglossal nerve geniohyoid (Alghamdi et al. 2017).
(Standring 2016). In a male neonate with Meckel syndrome, auriculoglos-
sus was present on each side of the body, originating from the
Comparative Anatomy region of the external auditory meatus and blending with hyo-
Genioglossus has a similar typical presentation in the apes, glossus at its insertion (Pettersen 1984). The left muscle was
extending from the mandible to the tongue and the hyoid more prominent and had a slip that continued from the associ-
(Deniker 1885; Sonntag 1923; Gibbs 1999; Diogo et al. ation with hyoglossus to blend with genioglossus. Additionally,
2010; 2012, 2013a,b, 2017). In a gorilla dissected by Diogo the genioglossus muscles were separated a bit more than usual
et al. (2010), the connection of genioglossus to the hyoid in the midline which gave the impression of two distinct
seemed to be loose or absent. tongue halves. An accessory muscle slip was also present in
the frenulum between the two genioglossus muscles.
Variations
Description Prevalence
Genioglossus may fuse with geniohyoid or receive fbers In a sample of 956 African American patients, Schaumann
from styloglossus (Macalister 1875; Knott 1883a; Lee and et al. (1970) found partial ankyloglossia in 21 individuals
Yang 2016; Standring 2016). Genioglossus may send a (2.2%). In 17 of the cases, the degree of ankyloglossia was
slip to the stylohyoid ligament (Macalister 1875; Bergman minor. In four of the cases, both the frenulum and the fbers
et al. 1988). The muscle may also be tripled (Macalister of genioglossus were markedly fbrosed.
1875). Genioglossus is also associated with several named
accessory slips. A slip from genioglossus to the epiglottis Clinical Implications
is referred to as levator epiglottidis (Bergman et al. 1988; Partial myotomy of the genioglossus muscle can be used
Patel and Loukas 2016) or genioepiglotticus (Luschka) with other surgical methods to correct ankyloglossia (Choi
(Macalister 1875). The central muscle of Bochdalek refers et al. 2011).
to a small muscle in the posterior quarter of the tongue
where the genioglossus muscles meet along the midline hyOglOssus (figure 2.4)
(Macalister 1875). A slip from genioglossus to constric-
tor pharyngis superior is referred to as geniopharyngeus Synonyms
(Winslow) (Macalister 1875; Knott 1883a; Bergman et al. N/A
1988; Patel and Loukas 2016). An isolated bundle of genio-
glossus that passes between the superior mental spine Typical Presentation
and the apex of the tongue is referred to as longitudinalis Description
linguae inferior medius (Bergman et al. 1988; Patel and Hyoglossus originates from the greater horn and the body
Loukas 2016). A separate deep slip of genioglossus that of the hyoid and inserts into the lateral aspect of the tongue
originates from the most inferior part of the genial tuber- between styloglossus and the inferior longitudinal muscle
cle and inserts into the hyoid is referred to as genioglossus (Standring 2016).
accessorius (Luschka) (Macalister 1875; Knott 1883a).
Innervation
Prevalence Hyoglossus is innervated by the hypoglossal nerve
Knott (1883a) found genioglossus accessorius separated (Standring 2016).
from the other fbers of genioglossus in one out of seven
subjects (14.3%). Comparative Anatomy
In the apes, hyoglossus is differentiated into ceratoglossus,
Anomalies which connects the greater horn of the hyoid to the tongue,
Description and chondroglossus, which primarily connects the body of
the hyoid to the tongue (the lesser horn is often reduced or
Shortness of genioglossus, a highly attached genioglossus,
absent) (Kohlbrügge 1890–1892; Saban 1968; Diogo et al.
or fbrosis of the genioglossus along with the frenulum can
2010, 2012, 2013a,b, 2017).
Variations and the hyoglossus muscles. Furthermore, the right genio-

Description glossus muscle was absent and replaced by a muscle mass
that passed from the inferior margin of the right mandible
The origin of hyoglossus may be continuous with a slip from
to the deep surface of the left hyoglossus muscle.
thyrohyoid (Macalister 1875). Geniohyoid may originate from
Mieden (1982) describes two male fetuses with cyclopia
hyoglossus (Bergman et al. 1988; Patel and Loukas 2016).
and alobar holoprosencephaly. In one case, auriculoglossus
Wood (1868) observed a case in which the superior belly of
was present on the left side, originating from the tympanic
the omohyoid was doubled and the upper belly fused with
bone and attaching to hyoglossus (Mieden 1982). In a male
hyoglossus and the middle pharyngeal constrictor. The bun-
neonate with Meckel syndrome, Pettersen (1984) found an
dles that comprise hyoglossus may vary in their development
auriculoglossus muscle on each side of the body originat-
and how separated they are from one another (Macalister
ing from the region of the external auditory meatus and
1875; Bergman et al. 1988; Ogata et al. 2002; Standring
blending with hyoglossus at its insertion. The left muscle
2016). When the fbers of hyoglossus that originate from the
was more prominent and had a slip that continued from the
greater horn are distinct from the fbers that originate from
association with hyoglossus to blend with genioglossus.
the body of the hyoid, the former comprise a bundle referred
In an otocephalic fetus examined by Lawrence and
to as ceratoglossus and the latter form a bundle referred to
Bersu (1984), the oral cavity was reduced and separated
as basioglossus (Bergman et al. 1988; Patel and Loukas
from the pharynx by a membrane deemed to be the buc-
2016). Chondroglossus is a bundle that originates from the
copharyngeal membrane. A small tongue was situated pos-
lesser horn and body of the hyoid and inserts into the intrin-
terior to the membrane and flled the pharynx. The dorsal
sic muscles of the tongue (Bergman et al. 1988; Ogata et al.
surface of the tongue faced the posterior pharyngeal wall
2002; Standring 2016). It may be considered as the third part
and the tip of the tongue was oriented inferiorly. The tongue
of hyoglossus but is typically partitioned from it by fbers of
was connected via muscle fbers to the body of the hyoid.
genioglossus (Le Double 1897; Ogata et al. 2002; Standring
Hyoglossus was the only extrinsic tongue muscle present in
2016). Triticeoglossus (Bochdalek) is a bundle that originates
this specimen. In a fetus with craniorachischisis, the attach-
from the cartilago triticea and inserts into the tongue with the
ment of hyoglossus was more medial than normal on the
posterior fbers of hyoglossus (Macalister 1875; Knott 1883a;
right side, and hyoglossus was fused with styloglossus on
Bergman et al. 1988; Patel and Loukas 2016; Standring 2016).
the left side (Alghamdi et al. 2017).
It may be considered as the fourth part of hyoglossus and is
often present when the posterior fbers of hyoglossus lie pos- Prevalence
terior to the lingual artery (Macalister 1875).
In their literature review, Smith et al. (2015) found that hyo-
glossus was anomalous in only 1 out of 20 individuals with
Prevalence trisomy 13 (5%).
Macalister (1875) notes that Bochdalek found triticeoglos-
sus in 8 out of 22 subjects (36.4%). Macalister (1875) found Clinical Implications
triticeoglossus as a bundle with connections to hyoglossus N/A
in one out of six subjects (16.7%) and as a completely dis-
tinct muscle in 1 out of 30 subjects (3.3%). Knott (1883a)
found triticeoglossus in 4 out of 44 cases (11.4%). In a stylOglOssus (figure 2.4)
sample of 100 tongue halves from 50 cadavers, Ogata et al. Synonyms
(2002) found chondroglossus in all specimens (100%) but N/A
observed that the chondroglossus muscle fbers were sparse
and poorly developed in 14% of specimens. Typical Presentation
Description
Anomalies Styloglossus originates from the styloid process and from
Description the stylomandibular ligament (Standring 2016). It inserts
In a fetus with trisomy 13, the right posterior belly of the along the lateral length of the tongue, blending with the
digastric inserted into the hyoid bone and had an extended inferior longitudinal muscle, and has an oblique insertion
attachment into the posterior aspect of hyoglossus (Pettersen that blends with hyoglossus (Standring 2016).
et al. 1979). Furthermore, the right hyoglossus muscle was
comprised of two distinct muscle sheets that were separated Innervation
by a gap. Bersu et al. (1976) describe a male infant with Styloglossus is innervated by the hypoglossal nerve
Hanhart syndrome in which the right mandible was repre- (Standring 2016).
sented by a small bone. In this infant, the right hyoglossus
was substantially reduced. An accessory muscle was pres- Comparative Anatomy
ent that sent a slip between the geniohyoid muscles to have a Styloglossus has a similar typical presentation in the apes,
diffuse insertion onto the deep aspect of the left geniohyoid originating from the styloid process and/or the temporal
bone and inserting into the tongue with some fbers blend- connected to the angle of the mandible by fbrous tracts. On
ing with hyoglossus (Kohlbrügge 1890–1892; Dean 1984; two sides (11.1%) of one specimen, an accessory bundle was
Gibbs 1999; Diogo et al. 2010, 2012, 2013a,b, 2017). In present that originated from the fbrous tract connecting the
one bonobo, Diogo et al. (2017) found that styloglossus angle of the mandible to Reichert’s cartilage.
was fused with the superior pharyngeal constrictor and
stylopharyngeus. Anomalies
Description
Variations Bersu et al. (1976) describe a male infant with Hanhart syn-
Description drome. In this infant, the right styloglossus was substan-
Styloglossus may be doubled, comprised of either two partially reduced. Mieden (1982) describes two male fetuses
allel bellies or two superimposed layers (Macalister 1875; with cyclopia and alobar holoprosencephaly. On the left
Bergman et al. 1988; Patel and Loukas 2016). It may origi- side of one specimen, styloglossus was absent.
nate from the medial pterygoid (Macalister 1875; Bergman On the right side of one infant with trisomy 18, an acces-
et al. 1988; Patel and Loukas 2016). It may send fbers to sory styloglossus muscle originated from the temporal bone
genioglossus or to the pharynx (Macalister 1875; Patel and (Bersu and Ramirez-Castro 1977). The accessory styloglos-
Loukas 2016). An accessory head of styloglossus may orig- sus muscle blended with hyoglossus just below the fbers
inate from the stylomandibular ligament (Macalister 1875; from the normal styloglossus muscle. In a male neonate
Knott 1883a; Patel and Loukas 2016). with Meckel syndrome, Pettersen (1984) found that both
Styloglossus may also be absent (Macalister 1875; styloglossus muscles originated anterior to the styloid pro-
Knott 1883a; Bergman et al. 1988; Patel and Loukas 2016). cess. On the right side of this specimen, the muscle had
When absent, styloglossus may be replaced by myloglos- an additional origin from an aponeurotic attachment to
sus (Wood), which originates from the inner aspect of the the stylohyoid ligament. In a fetus with craniorachischi-
angle of the mandible and inserts into the lateral aspect of sis, Alghamdi et al. (2017) found that the right styloglos-
the tongue (see the entry for this muscle) (Wood 1867b; sus originated from an enlarged styloid process and from
Macalister 1875; Knott 1883a; Valenti 1926; Bergman et al. the “temporo-occipital” bone. It inserted into the outer side
1988; Nakajima and Nakamura 2008; Patel and Loukas of the mandible lateral to hyoglossus. The left styloglossus
2016; Buffoli et al. 2017). Myloglossus can present as a slip was absent.
or second head of styloglossus instead of a distinct muscle
(Macalister 1875; Patel and Loukas 2016). Prevalence
Styloglossus may connect to the accessory muscle stylo- N/A
auricularis (Hyrtl) which originates from the styloid process
and attaches to the cartilage of the external auditory meatus Clinical Implications
(see the entry for this muscle) (Macalister 1875; Knott N/A
1883a; Patel and Loukas 2016; Touré and Anzouan-Kacou
2016). Styloglossus may also connect to auriculoglossus
(Gruber), a slip that may pass from the ear to the tongue MylOglOssus (figure 2.4)
(Macalister 1875). A slip from genioglossus to the epiglottis See also: Styloglossus
is referred to as levator epiglottidis (Bergman et al. 1988)
or genioepiglotticus (Luschka) and may be continuous with Synonyms
styloglossus (Macalister 1875; Patel and Loukas 2016). This muscle may also be referred to as mandibuloglossus
(Valenti 1926; Bergman et al. 1988).
Prevalence
As noted in Mérida‐Velasco et al. (2006), Barnwell (1977) Typical Presentation
studied the origin of styloglossus in 14 fetuses. Styloglossus This muscle is only present as a variation.
originated from the styloid process and the stylomandibu-
lar ligament in 10 fetuses (71.4%) and from the styloman- Comparative Anatomy
dibular ligament only in one fetus (7.1%). Mérida‐Velasco N/A
et al. (2006) studied the origin of styloglossus in 18 sides
of nine fetuses. In all cases (100%), styloglossus originated Variations
in Reichert’s cartilage, the structure from which the styloid Description
process arises during development. On 12 sides (66.7%), Myloglossus originates from the inner surface of the angle
styloglossus had an accessory head that originated from of the mandible and inserts into the lateral aspect of the
the angle of the mandible. The authors note that this acces- tongue (Wood 1867b; Macalister 1875; Knott 1883a; Valenti
sory head likely does not correspond to myloglossus as it 1926; Bergman et al. 1988; Nakajima and Nakamura
attaches to the main belly of styloglossus before reach- 2008; Patel and Loukas 2016; Buffoli et al. 2017). Knott
ing the tongue. On four sides (22.2%), styloglossus was (1883a) attributes its name to Rolfncius but other authors
(e.g., Valenti 1926; Bergman et al. 1988) credit Wood. It may note that this accessory head likely does not correspond to
present as a slip or second head of styloglossus instead of a myloglossus as it attaches to the main belly of styloglossus
distinct muscle (Macalister 1875; Patel and Loukas 2016). In before reaching the tongue. However, this accessory head
these cases, it often joins with styloglossus before reaching would be considered a myloglossal structure according
the tongue (Buffoli et al. 2017). Myloglossus may be associ- to the classifcation used by Buffoli et al. (2017). On four
ated with a rudimentary styloglossus or replace styloglossus sides (22.2%), styloglossus was connected to the angle of
when it is absent (Macalister 1875; Nakajima and Nakamura the mandible by fbrous tracts. On two sides (11.1%) of one
2008; Patel and Loukas 2016; Buffoli et al. 2017). specimen, an accessory bundle was present that originated
Myloglossus may join with hyoglossus upon reaching from the fbrous tract connecting the angle of the mandible
the tongue (Wood 1867b; Valenti 1926; Buffoli et al. 2017). to Reichert’s cartilage.
In the cases observed by Nakajima and Nakamura (2008),
myloglossus bifurcated around hyoglossus just before Anomalies
inserting into the tongue. One bundle passed to the apex
N/A
of the tongue while the other blended with hyoglossus, thus
having an insertion similar to a normal styloglossus muscle.
According to Nakajima and Nakamura (2008), styloglos- Clinical Implications
sus typically develops as a bicipital muscle with the head N/A
from the angle of the mandible disappearing before birth,
so myloglossus represents the persistence of this muscular
stylOauricularis (figure 2.4)
head. Buffoli et al. (2017) suggest a different mechanism of
development and argue that myloglossus forms from some See also: Styloglossus
bundles of the medial pterygoid muscle that fuse with the
inferolateral portion of styloglossus. Synonyms
This muscle may also be referred to as depressor auriculae
Innervation (Lauth) (Macalister 1875; Knott 1883a).
In the bilateral presentation of myloglossus described by
Nakajima and Nakamura (2008), one side was innervated Typical Presentation
by both the hypoglossal and the mandibular nerve while This muscle is only present as a variation or anomaly.
the other side was innervated by the hypoglossal nerve only.
Innervated was supplied by both the hypoglossal nerve Comparative Anatomy
and/or the mandibular nerve (via the buccal or mylohyoid Styloauricularis may be the homologue of the mandibulo-
nerves) in the cases described by Buffoli et al. (2017). auricularis muscle, which is typically absent in most pri-
mates but found in other mammals (Huber and Hughson
Prevalence 1926; Diogo et al. 2018).
In a sample of 21 sides from 11 heads, Buffoli et al. (2017)
found “myloglossal structures” on 13 sides (61.9%). Out of Variations
these 13 cases, a myloglossus muscle was found in fve cases Description
(38.5%), a myloglossal ligament was found in three cases Styloauricularis (Hyrtl) originates from the styloid process
(23%), and a myloglossal ligament that had some muscu- and inserts into the cartilage of the external auditory meatus
lar fbers was found in fve cases (38.5%). The myloglossus (Macalister 1875; Knott 1883a; Patel and Loukas 2016;
muscles were associated with a normal-sized styloglossus Touré and Anzouan-Kacou 2016). It is described here along
muscle in one out of fve cases (20%) and a rudimentary with the tongue muscles as it is considered an auricular slip
styloglossus in four cases (80%). The myloglossal ligaments of styloglossus (Macalister 1875). A variant of styloauricu-
were associated with a normal-sized styloglossus muscle in laris is auriculoglossus (Gruber), a slip that may pass from
seven out of eight cases (87.5%) and a rudimentary stylo- the ear to the tongue (Macalister 1875). Styloauricularis
glossus muscle in one case (12.5%). Out of the 13 cases, may have tendinous or feshy attachments (Macalister 1875;
the myloglossal structures inserted into hyoglossus (lingual Touré and Anzouan-Kacou 2016). It may be comprised of
insertion) in four cases (30.8%) and into styloglossus in one or two bellies, have two heads, have two tendons, or
nine cases (69.2%). All the myloglossal ligaments inserted may be replaced by a fbrous cord (Macalister 1875; Knott
into styloglossus (100%) while four out of fve myloglossus 1883a).
muscles (80%) inserted into hyoglossus.
Mérida‐Velasco et al. (2006) studied the origin of sty- Innervation
loglossus in 18 sides of nine fetuses. In all cases (100%), Styloauricularis is innervated by the facial nerve (Touré
styloglossus originated in Reichert’s cartilage, the structure and Anzouan-Kacou 2016).
from which the styloid process arises during development.
On 12 sides (66.7%), styloglossus had an accessory head Prevalence
that originated from the angle of the mandible. The authors N/A
Anomalies MUSCLES OF THE SOFT PALATE

Description
tensOr veli palatini (figure 2.5)
Styloauricularis is present in the form of auriculoglos-
sus in some individuals with congenital malformations. Synonyms
Mieden (1982) describes two male fetuses with cyclopia This muscle may also be referred to as tensor palati
and alobar holoprosencephaly. In one case, auriculoglos- (Macalister 1875).
sus was present on the left side, originating from the tym-
panic bone and attaching to hyoglossus (Mieden 1982). Typical Presentation
In a male neonate with Meckel syndrome, Pettersen Description
(1984) found an auriculoglossus muscle on each side of Tensor veli palatini originates from the scaphoid fossa of
the body originating from the region of the external audi- the pterygoid process and from the medial surface of the
tory meatus and blending with hyoglossus at its insertion. sphenoidal spine (Standring 2016). It also has an origin
The left muscle was more prominent and had a slip that from the pharyngotympanic tube (Eustachian tube or audi-
continued from the association with hyoglossus to blend tory tube) (Standring 2016). Its fbers end in a tendon that
with genioglossus. courses around the pterygoid hamulus to insert into the pal-
atine aponeurosis (Standring 2016). Some researchers may
Prevalence
use the name dilator tubae to refer to the portion of tensor
N/A veli palatini that originates from the pterygoid hamulus and
has attachments to the pharyngotympanic cartilage, con-
nective tissue lateral to the tubal wall, and Ostmann’s fat
Styloauricularis can be a landmark for the facial nerve pad (Standring 2016).
(Touré and Anzouan-Kacou 2016).
Innervation
Tensor veli palatini is innervated by the medial pterygoid
branch of the mandibular nerve (Standring 2016).
FIGURE 2.5 Muscles of the soft palate in posterior view.

Comparative Anatomy et al. (2004) found additional insertions into the maxil-
Tensor veli palatini has a similar typical presentation in lary tuberosity on 40 sides (33.6%) and into the submuco-
the apes, extending from the Eustachian tube cartilage and sal tissue near the palatoglossal arch on 45 sides (37.8%).
adjacent regions of the cranium (e.g., scaphoid fossa) to the Overall, a palatine insertion only was found on 54 sides
pterygoid hamulus and soft palate (Gratiolet and Alix 1866; (45.4%), a palatine and pharyngeal insertion were found
Sonntag 1923; Dean 1985; Gibbs 1999; Diogo et al. 2010, on 25 sides (21%), a palatine and maxillary insertion were
2012, 2013a,b, 2017). As the palate is closer to the roof of found on 20 sides (16.8%), and a palatine insertion with
the nasopharynx in adult apes than it is in adult humans, both secondary insertions was found on 20 sides (16.8%).
tensor veli palatini in the apes does not pass as distinctly
downward to reach the palate as it does in humans (Aiello Anomalies
and Dean 1990; Gibbs 1999; Diogo et al. 2010, 2012, Description
2013a,b, 2017). The morphology of tensor veli palatini often varies in indi-
viduals with cleft palate (e.g., Matsune et al. 1991; Arnold
Variations et al. 2005; Heidsieck et al. 2016; George et al. 2018).
Matsune et al. (1991) measured the ratio of the length of
Description
the Eustachian tube cartilage with insertion of tensor veli
Tensor veli palatini may be longitudinally divided palatini to its total length (nasopharyngeal end to tubal
(Macalister 1875; Patel and Loukas 2016). The portion isthmus). In cleft palate cases, this ratio was smaller than
designated as dilator tubae may vary in its development in control cases and the insertions had fewer muscular
and connections to the rest of the muscle (Barsoumian and tendinous fbers (Matsune et al. 1991; Heidsieck et al.
et al. 1998). The attachment to the pharyngotympanic 2016). These results demonstrate an abnormal insertion
tube cartilage varies with age (Suzuki et al. 2003). Tensor into the Eustachian tube cartilage in cleft palate patients.
veli palatini may have an insertion into the palatine bone Arnold et al. (2005) examined a fetus with bilateral cleft
(Macalister 1875; Standring 2016). It may also have addi- palate and found that tensor veli palatini was situated
tional insertions into the maxillary tuberosity, or the sub- under the auditory tube and inserted at the lateral wall
mucosal tissue close to the palatoglossal arch (Abe et al. of the auditory tube. George et al. (2018) found that ten-
2004; Patel and Loukas 2016). Muscular fbers may be sor veli palatini was shorter and less voluminous in adults
present in the tendon near its insertion, giving tensor veli with repaired cleft palate than in adults without cleft
palatini a digastric appearance (Macalister 1875). Tensor palate.
veli palatini may receive an accessory slip from the medial It can also vary in individuals with other congenital
pterygoid or from the outer margin of the scaphoid fossa defects (Herring et al. 1979; Lawrence and Bersu 1984;
(Macalister 1875; Patel and Loukas 2016). It may send a Pettersen 1984; Alghamdi et al. 2017; Perry et al. 2020).
slip to the buccinator muscle (Macalister 1875; Patel and In an infant with mandibulofacial dysostosis, tensor veli
Loukas 2016). Some fbers of tensor veli palatini may be palatini was enlarged (Herring et al. 1979). In an otoce-
continuous with fbers of tensor tympani (Barsoumian phalic fetus examined by Lawrence and Bersu (1984),
et al. 1998; Standring 2016). muscle fbers were present in the soft palate, but indi-
vidual palatal muscles could not be identifed. In a male
Prevalence
neonate with Meckel syndrome, Pettersen (1984) found
Barsoumian et al. (1998) examined tensor veli palatini and that the right tensor veli palatini traveled lateral to the lat-
dilator tubae on 20 sides from 16 heads. On 13 sides (65%) eral pterygoid plate and its tendon blended with the lower
from 11 heads, most of dilator tubae was distinct from ten- portion of it. Tensor veli palatini was absent in the fetus
sor veli palatini and the two only shared connective tis- with craniorachischisis described by Alghamdi et al.
sue and some fbers. On fve sides (25%) from fve heads, (2017). Perry et al. (2020) found that tensor veli palatini
dilator tubae was strongly intermingled with tensor veli was shorter, thinner, and less voluminous in children with
palatini. On two sides (10%), dilator tubae was defcient. 22q11.2 deletion syndrome (DiGeorge syndrome) relative
Abe et al. (2004) studied tensor veli palatini on 119 to control subjects.
sides from 77 heads. The muscle originated from the audi-
tory tube and cranial base in all cases. On 29 sides (24.4%),
the auditory tube and cranial base origins were equal in Prevalence
width. On eight sides (6.7%), the auditory tube origin was Using histology, Matsune et al. (1991) measured the ratio of
wider than the cranial base origin. On 41 sides (34.5%), the the length of the Eustachian tube cartilage with insertion of
auditory tube origin was narrower than the cranial base tensor veli palatini to its total length (nasopharygneal end
origin. On 37 sides (31.1%), the auditory tube origin was to tubal isthmus) in 20 control cases and in ten individu-
narrower than the cranial base origin and the latter was als with cleft palate. In all 20 controls (100%) and six cleft
shifted anteromedially toward the pharyngeal opening of palate cases (60%), tensor veli palatini inserted into the tip
the auditory tube. On four sides (3.4%), the origins did not of the lateral lamina of the Eustachian tube cartilage. Four
overlap and were separated posterolaterally. In addition to cleft palate cases (40%) lacked an insertion. Overall, the
the primary insertion into the palatine aponeurosis, Abe ratio of tensor veli palatini insertion on the Eustachian tube
cartilage to the total length of this cartilage was smaller in split at its origin (Macalister 1875; Patel and Loukas 2016).
the cleft palate cases. Levator veli palatini may also have a single origin with a
doubled insertion (Macalister 1875; Patel and Loukas 2016).
An abnormal attachment or variation in size or length of Prevalence
tensor veli palatini in individuals with cleft palate likely
N/A
contributes to recurrent otitis media with effusion (Matsune
et al. 1991; Arnold et al. 2005; Heidsieck et al. 2016; George Anomalies
et al. 2018).
Description
The morphology of levator veli palatini may vary in
levatOr veli palatini (figure 2.5) individuals with cleft palate or other palatal anomalies
(e.g., Pettersen 1984; Lindman et al. 2001; Senoo et al. 2001;
Synonyms
Arnold et al. 2005; Kotlarek et al. 2017; Trudel et al. 2018).
This muscle may also be referred to as levator palati In a male neonate with Meckel syndrome, Pettersen (1984)
(Macalister 1875). observed that the right levator veli palatini split around the
Typical Presentation hamulus of the medial pterygoid plate and attached onto the
periosteum of the partial palatal shelf. The left muscle traveled
Description medial to the hamulus and attached to the small fap of the soft
Levator veli palatini originates via a tendon from the infe- palate. Kotlarek et al. (2017) found that levator veli palatini
rior surface of the petrous temporal bone, the cartilage had smaller circumference, diameter, and volume in adults
of the pharyngotympanic tube (Eustachian tube or audi- with repaired cleft palate than in adults without cleft palate.
tory tube), and from the vaginal process of the sphenoid Lindman et al. (2001) found that levator veli palatini had
(Standring 2016). It inserts into the palatine aponeurosis a smaller mean muscle fber diameter and greater variability
between the two fasciculi of palatopharyngeus (Standring in fber size and form in sample of infants with cleft palate
2016). Levator veli palatini intermingles with its counter- compared to normal adults. Arnold et al. (2005) examined
part at the midline (Standring 2016). a fetus with bilateral cleft palate and found that levator veli
palatini originated only from the auditory tube, ran parallel
Innervation and medial to the auditory tube, and inserted into the dorso-
medial edge of the palatal cleft.
Levator veli palatini is innervated by the pharyngeal plexus
Senoo et al. (2001) describe a case of congenital unilat-
(Standring 2016).
eral soft palate aplasia in which palatoglossus and levator veli
Comparative Anatomy palatini were absent on the right side. In a child with palatal
anomaly associated with 8q22.1–22.2 microduplication, leva-
Levator veli palatini has a similar typical presentation in
tor veli palatini had an insertion into the hard palate bilaterally
the apes, extending from the Eustachian tube and the apex
(Trudel et al. 2018). There were also thin, membranous gaps
of the petrous temporal bone to the soft palate (Gratiolet
between levator veli palatini and palatoglossus on both sides.
and Alix 1866; Kohlbrügge 1890–1892; Sonntag 1923;
Dean 1985; Gibbs 1999; Diogo et al. 2010, 2012, 2013a,b,
2017). As the palate is closer to the roof of the nasophar- Prevalence
ynx in adult apes than it is in adult humans, levator veli N/A
palatini is more horizontal in the apes and does not pass as
distinctly downward to reach the palate as it does in humans Clinical Implications
(Gratiolet and Alix 1866; Sonntag 1923; Aiello and Dean An aberrant insertion of levator veli palatini may contribute
1990; Gibbs 1999; Diogo et al. 2010, 2012, 2013a,b, 2017). to hypernasal speech (Seif and Dellon 1978).
Variations Musculus uvulae (figure 2.5)

Levator veli palatini may be longitudinally divided with This muscle may also be referred to as azygos uvulae or
the fbers from the Eustachian tube and the fbers from the motor uvulae (Harrison 1848; Macalister 1875).
petrous bone remaining separated (Macalister 1875; Patel
and Loukas 2016). The portion of the muscle that originates Typical Presentation
from the Eustachian tube may be referred to as salpingo- Description
staphylinus (Macalister 1875; Patel and Loukas 2016). Musculus uvulae originates from the posterior nasal spine
Levator veli palatini may entirely arise from the cartilage of the palatine bone and from the palatine aponeurosis
of the Eustachian tube or this tubal origin may be reduced (Standring 2016). It courses posteriorly to insert beneath the
(Macalister 1875; Patel and Loukas 2016). The portion of uvular mucosa (Standring 2016). It merges with its counter-
the muscle that originates from the petrous bone may be part along the majority of its length (Standring 2016).

Musculus uvulae is innervated by the pharyngeal plexus N/A
(Standring 2016).
palatOglOssus (figure 2.5)
Comparative Anatomy See also: Muscles of the tongue
There is no information on musculus uvulae in bono-
bos (Diogo et al. 2017). It is well-developed in gibbons Synonyms
(Kohlbrügge 1890–1892; Diogo et al. 2012). In gorillas, N/A
it originates from the soft palate and courses posterior
to insert onto or near to the uvula (Diogo et al. 2010). In Typical Presentation
orangutans and common chimpanzees, it is present but may
end in a membrane without attaching to the uvula (Gratiolet Description
and Alix 1866; Chapman 1880; Sonntag 1923, 1924; Gibbs Palatoglossus originates from the palatine aponeurosis of
1999; Diogo et al. 2013a,b). the soft palate (Standring 2016). It inserts into the side of
the posterior tongue, with some fbers attaching to the dor-
Variations sal surface of the tongue and other fbers blending with the
Description intrinsic transverse muscle (Standring 2016). At its origin,
The degree of separation between the two uvular muscles palatoglossus is continuous with its counterpart on the other
on either side of the midline can vary from complete fusion side of the body (Standring 2016). Though it is considered
to complete separation (Macalister 1875). Macalister (1875) one of the extrinsic muscles of the tongue, palatoglossus is
describes a patient that exhibited an unusual action of mus- described in this section as its innervation and function are
culus uvulae on the uvula and concludes that the muscle more similar to the soft palate muscles.
was inserted into the extreme tip of the uvula.
Innervation
Prevalence Palatoglossus is innervated by the pharyngeal plexus
N/A (Standring 2016).
Comparative Anatomy
Anomalies
There is no information on palatoglossus in orangutans
Description (Gibbs 1999; Diogo et al. 2013b). In the other apes, palato-
The morphology of musculus uvulae may vary in individuals glossus corresponds to muscle fbers within the palatoglos-
with cleft palate (Lewin et al. 1980; Shprintzen et al. 1985; sal fold that connect the soft palate to the supero-posterior
Todd and Krueger 1992; Perry et al. 2019). In cases of clas- portion of the tongue (Gibbs 1999; Diogo et al. 2010, 2012,
sic or occult submucous cleft palate, musculus uvulae is often 2013a, 2017). The muscle is most distinct in common chim-
absent, is defcient, or shows some degree of diastasis (Lewin panzees and bonobos (Diogo et al. 2013a, 2017).
et al. 1980; Shprintzen et al. 1985). Todd and Krueger (1992)
described cases of what they termed miniscule submucous Variations
cleft palate that exhibited reduced bulk of musculus uvulae. Description
Perry et al. (2019) found that musculus uvulae was reduced in Palatoglossus may send some fbers to palatopharyngeus or
volume in adults with repaired cleft palate than in adults with- be connected with this muscle by a slip (Macalister 1875).
out cleft palate. In an otocephalic fetus examined by Lawrence Its origin may vary, with potential attachments closer to the
and Bersu (1984), muscle fbers were present in the soft palate uvula or the rim of the hard palate (Kuehn and Azzam 1978;
but most individual palatal muscles could not be identifed. Patel and Loukas 2016). Palatoglossus is in close proximity
The only muscle clearly present was musculus uvulae. to an accessory slip, the amygdaloglossal muscle (Broca),
which extends between the muscles of the tongue and the
Prevalence
capsule of the palatine tonsil (Patel and Loukas 2016).
In 29 patients with occult submucous cleft palate, Lewin
et al. (1980) found that ten patients (34.5%) had an absent Prevalence
or hypoplastic musculus uvulae and 19 patients (65.5%) N/A
showed signs of defcient muscular fusion. In a sample
of 25 children with bifd uvulae, Shprintzen et al. (1985)
Anomalies
found that musculus uvulae was absent in 19 children
(76%). Of the remaining six children, two exhibited a Description
musculus uvulae with a deep midline groove, two had a Senoo et al. (2001) describe a case of congenital unilat-
small uvular muscle mass, and two had a moderate uvular eral soft palate aplasia in which palatoglossus and levator
muscle mass. veli palatini were absent on the right side. In a fetus with
craniorachischisis, palatoglossus was the only soft palate/ have a similar typical presentation as it connects the pala-
pharyngeal muscle that was present (Alghamdi et al. 2017). tine region to the pharyngeal wall (Kohlbrügge 1890–1892;
Bersu et al. (1976) describe a male infant with Hanhart syn- Diogo et al. 2010, 2012, 2013a, 2017). In a gorilla dissected
drome. On the right side of this infant, the muscles of the by Diogo et al. (2010), palatopharyngeus was blended with
soft palate were underdeveloped and displaced. A muscle the inferior pharyngeal constrictor.
descended from the cartilage of the auditory tube and split
into two portions at the level of the soft palate. The anterior Variations
portion of this muscle inserted onto the side of the tongue Description
forming the right palatoglossal arch. The posterior portion
The extent of the origin from the nasal side of the palatine
blended with the superior pharyngeal constrictor and the
aponeurosis may vary (Okuda et al. 2008; Sumida et al. 2012).
stylopharyngeus muscles forming the palatopharyngeal
Sumida et al. (2012) argued that palatopharyngeus has two
arch. In an otocephalic fetus examined by Lawrence and
additional fasciculi of origin, the first being a posterior fas-
Bersu (1984), muscle fibers were present in the soft palate,
ciculus that originated from the uvula and the second being
but most individual palatal muscles could not be identified.
salpingopharyngeus (see the entry for this muscle). The
In a male neonate with Meckel syndrome, Pettersen (1984)
nature of its insertions into the thyroid cartilage, hypophar-
found that palatoglossus and palatopharyngeus originated
ynx, and posterior pharynx can also vary (Sumida et al. 2012).
from the medial aspect of the medial pterygoid plate on
Palatopharyngeus may receive some fibers from palatoglossus
each side, instead of originating from the soft palate.
or be connected with this muscle by a slip (Macalister 1875).
Prevalence Palatopharyngeus may be split into two or three parts
N/A (Macalister 1875). Macalister (1875) notes that Winslow
found the muscle divided into three portions including
Clinical Implications peristaphylo-pharyngeus (velum to pharynx), pharyngo-
Ma et al. (1999) suggest that a short palatoglossus is associ- staphylinus (uvula to pharynx), and thyro-staphylinus (uvula
ated with hypernasal or nasal emissive speech. to the posterior margin of the thyroid cartilage). Macalister
(1875) also notes that Luschka divided the muscle into a pars
thyreo-palatina and a pars pharyngo-palatina.
Palatopharyngeus (Figure 2.5) Palatopharyngeus is associated with Passavant’s muscle
See also: Salpingopharyngeus (palatopharyngeal sphincter) (Whillis 1930; Standring
2016; Sakamoto 2016a; Sumida et al. 2017). Its existence
Synonyms is debated in the literature (Standring 2016; Sumida et al.
This muscle may also be referred to as pharyngopalatinus 2017). Some consider Passavant’s muscle to be a portion
(Whillis 1930). of the superior pharyngeal constrictor and/or palatopha-
ryngeus muscles, perhaps due to the difficulty in distin-
Typical Presentation guishing it from the former (Whillis 1930; Sumida et al.
Description 2012; Sakamoto 2015; Standring 2016; Sumida et al. 2017).
Palatopharyngeus is comprised of two fasciculi (Standring Others recognize it as a distinct, sphincter-like muscle that
2016). The anterior fasciculus originates from the posterior originates from the nasal aspect of the lateral palatine apo-
margin of the hard palate and from the palatine aponeuro- neurosis and passes dorsolateral to levator veli palatini to
sis (Standring 2016). The posterior fasciculus has a connec- surround the pharynx (Standring 2016; Sumida et al. 2017).
tion with the mucosa of the pharyngeal region of the palate Palatopharyngeus is also in close proximity to several
(Standring 2016). The two fasciculi join in the posterolateral rare accessory pharyngeal muscles, including petropharyn-
margin of the soft palate and receive a contribution from geus (see the entry for this muscle), occipitopharyngeus,
salpingopharyngeus (Standring 2016). Palatopharyngeus mastoidopharyngeus, sphenopharyngeus, pterygopharyn-
inserts with stylopharyngeus into the posterior border of the geus externus, azygospharyngeus, and tympanopharyngeus
thyroid cartilage (Standring 2016). It also partially inserts (Bergman et al. 1988; Knott 1883a; Sakamoto 2016a). Some
into pharyngeal fibrous tissue and decussates with the fibers of these muscles may be considered variants of the same
of its counterpart (Standring 2016). accessory muscle, cephalo-pharyngeus (Macalister 1875;
Knott 1883a).
Innervation
Palatopharyngeus is innervated by the pharyngeal plexus Prevalence
(Standring 2016). Okuda et al. (2008) studied palatopharyngeus in 20 Japanese
cadavers. In all cases (100%), an origin was present from
Comparative Anatomy the oral tendinous part of the posterior palatine aponeuro-
There is no information on this muscle in orangutans sis, and this oral side origin interlaced with the fibers of the
(Diogo et al. 2013b). Though there is also little information contralateral palatopharyngeal muscle. In 14 cases (70%),
on this muscle in the other apes, palatopharyngeus seems to an origin was present from the nasal tendinous part of the
posterior palatine aponeurosis. In 17 cases (85%), this nasal palatoglossus and palatopharyngeus originated from the
side origin interlaced with the fibers of the contralateral medial aspect of the medial pterygoid plate on each side,
palatopharyngeal muscle. instead of originating from the soft palate (Pettersen 1984).
Sumida et al. (2012) studied palatopharyngeus on 100
sides from 50 Japanese cadavers. These authors found four
fasciculi of origin, the typical oral and nasal fasciculi, a Prevalence
posterior fasciculus that originated from the uvula, and N/A
salpingopharyngeus. On all sides, the oral fasciculus origi-
nated from the posterior border of the palatine aponeurosis. Clinical Implications
On 86 sides (86%), the oral fasciculus had an additional ori- Lindman and Stål (2002) found that patients with sleep-
gin from the median line of the soft palate. In a subsample disordered breathing demonstrated abnormalities in pala-
of 72 sides, the posterior fasciculus was present only on topharyngeus including an increase in connective tissue, an
40 sides (55.6%). In a subsample of 77 sides, the nasal fas- increase in the proportion of smaller-sized muscle fibers,
ciculus was present only on 59 sides (76.6%). From these and other variations of fiber type.
59 sides, the nasal fasciculus was well-developed on five
sides (8.4%), extended toward the palatine aponeurosis on Salpingopharyngeus (Not Illustrated)
27 sides (45.8%), and ended in the nasal mucosa of the soft
palate before reaching the aponeurosis on 27 sides (45.8%). Synonyms
Palatopharyngeus, together with contributions from sal- N/A
pingopharyngeus and stylopharyngeus, also had variable
insertions (Sumida et al. 2012). Muscle fibers ending at the Typical Presentation
posterior pharyngeal insertion were found on only 18 out of Description
the 100 sides (18%). In a subsample of 87 sides, there was Salpingopharyngeus originates from the cartilage of the
an insertion into the thyroid cartilage via two bundles on 37 pharyngotympanic tube and merges with palatopharyngeus
sides (42.5%) and via one bundle on 50 sides (57.5%). In a (Standring 2016).
subsample of 94 sides, the muscle fibers ended at the level
of the cricopharyngeal part of the inferior pharyngeal con- Innervation
strictor and formed an aponeurosis on 63 sides (67%), with Salpingopharyngeus is innervated by the pharyngeal plexus
two of these sides extending to the upper end of the esopha- (Standring 2016).
gus. On 11 of these 94 sides (11.7%), the muscle fibers ended
at the same level but did not form an aponeurosis, with one Comparative Anatomy
of these sides extending to the upper end of the esophagus. There is no information on this muscle in orangutans (Diogo
In the remaining 20 sides (21.3%), the fibers did not reach et al. 2013b). In gibbons, the pharyngeal muscles have an
the cricopharyngeal part of the inferior pharyngeal con- origin from the pharyngotympanic tube, which may contain
strictor and did not form an aponeurosis. fibers that correspond to salpingopharyngeus (Kohlbrügge
1890–1892; Diogo et al. 2012). In one gorilla, Diogo et al.
Anomalies (2010) were not able to identify fleshy fibers within the salpin-
gopharyngeal fold. Diogo et al. (2013a) state that salpingopha-
Description
ryngeus in common chimpanzees is similar to that of humans.
Hypertrophy of Passavant’s muscle may occur in individu- Diogo et al. (2017) were not able to determine if salpingopha-
als with complete cleft palate (Standring 2016). In an infant ryngeus was present as a distinct muscle in bonobos.
with mandibulofacial dysostosis, palatopharyngeus origi-
nated from the edge of the cleft soft palate and was closely Variations
associated with levator veli palatini (Herring et al. 1979).
Description
Furthermore, the superior constrictor and palatopharyn-
geus were continuous at their posterior and inferior attach- Macalister (1875) considers salpingopharyngeus a detached
ments and inserted together into the posterior raphe. slip of palatopharyngeus. Knott (1883a) considered it a vari-
Bersu et al. (1976) describe a male infant with Hanhart ant of an accessory cephalo-pharyngeus muscle. Sumida
syndrome. On the right side of this infant, the muscles of et al. (2012) consider it a fourth fasciculus of origin for pala-
the soft palate were underdeveloped and displaced. A mus- topharyngeus. Salpingopharyngeus may be absent (Sumida
cle descended from the cartilage of the auditory tube and et al. 2012). It may be partially or fully replaced by connec-
split into two portions at the level of the soft palate. The tive and/or adipose tissue (Sumida et al. 2012).
anterior portion of this muscle inserted onto the side of the
tongue forming the right palatoglossal arch. The posterior Prevalence
portion blended with the superior pharyngeal constrictor In a sample of 98 sides from 49 Japanese cadavers, Sumida
and the stylopharyngeus muscles forming the palatopha- et al. (2012) found that salpingopharyngeus was present as
ryngeal arch. In a male neonate with Meckel syndrome, a distinct muscle bundle on 59 sides (60.2%).
Description Constrictor pharyngis superior is innervated by the pharyn-
In a male neonate with Meckel syndrome, salpingo- geal plexus (Standring 2016).
pharyngeus was absent (Pettersen 1984). It was absent
bilaterally in an otocephalic fetus (Lawrence and Bersu Comparative Anatomy
1984). In an individual with a completely bony left pha- Constrictor pharyngis superior may be only comprised
ryngotympanic tube that communicated with the sphe- of the glossopharyngeal part in gibbons and orangutans
noid sinus, salpingopharyngeus was hypoplastic (Khan (Kohlbrügge 1890–1892; Diogo et al. 2012, 2013b). All
et al. 2017). four parts of the superior constrictor and an insertion into
the pharyngeal raphe have been found in gorillas, common
Prevalence chimpanzees, and bonobos (Gibbs 1999; Diogo et al. 2010,
N/A 2013a, 2017). Pterygopharyngeus is present as a muscle
distinct from the superior constrictor in gibbons, extend-
Clinical Implications ing between the pterygoid hamulus and the pharyngeal wall
N/A (Kohlbrügge 1890–1892; Saban 1968; Diogo et al. 2012).
Variations
PHARYNGEAL MUSCLES
Description
Constrictor pharyngis superior (Superior Macalister (1875) notes that Luschka considered constrictor
pharyngeal constrictor) (Figure 2.6) pharyngis superior a composite muscle comprised of four
parts: pterygo-pharyngeus, bucco-pharyngeus, mylo-pha-
See also: Petropharyngeus
ryngeus, and glosso-pharyngeus. The buccopharyngeal part
Synonyms may connect with buccinator (Macalister 1875; Shimada
and Gasser 1989). The extent of the origin from the ptery-
N/A
goid hamulus may vary (Macalister 1875). It may have an
origin from the medial pterygoid plate (Standring 2016)
Typical Presentation or the petrous part of the temporal bone (Bergman et al.
1988; Sakamoto 2009, 2016a). Stylopharyngeus may course
Description transversely and join the superior pharyngeal constrictor
Constrictor pharyngis superior originates from the ptery- (Choi et al. 2020). A portion of the superior constrictor
goid hamulus, pterygomandibular raphe, mylohyoid line may run longitudinally and pass between the superior and
of the mandible, and side of the tongue (Standring 2016). middle constrictors, or merge with the middle constrictor
It inserts into the pharyngeal raphe, which attaches to the and the contralateral constrictor muscles (Choi et al. 2020).
pharyngeal tubercle of the occipital bone (Standring 2016). An accessory muscle, petropharyngeus, may also be joined
FIGURE 2.6 Pharyngeal muscles in posterior view.

with the superior constrictor (Macalister 1875; Knott 1883a; five sides (11.4%), a portion of the superior constrictor ran
see the entry for this muscle). A slip from genioglossus to longitudinally and either passed between the superior and
constrictor pharyngis superior is referred to as geniopharyn- middle constrictors or merged with the middle constrictor
geus (Winslow) (Macalister 1875; Knott 1883a; Bergman and the contralateral constrictor muscles.
et al. 1988; Patel and Loukas 2016). Pterygopharyngeus
externus, which attaches to the pterygoid hamulus, may be Anomalies
present as a distinct muscle (Sakamoto 2016a). Description
Constrictor pharyngis superior is also associated with azy- Bersu et al. (1976) describe a male infant with Hanhart
gos-pharyngeus (azygos pharyngis or solitarius pharyngis), syndrome. In this infant, the inferior parts of both superior
which originates from the pharyngeal tubercle and inserts pharyngeal constrictors were abnormally thickened. On the
into the pharyngeal raphe (Macalister 1875; Knott 1883a; right side, some of its fibers inserted into a pterygoman-
Bergman et al. 1988; Sakamoto 2016a). When present, it over- dibular raphe while the rest of the fibers inserted into the
lies the superior and middle constrictors along the midline, skull near the pterygoid laminae. On the left side, all fibers
and it may be fused with the superior constrictor (Macalister inserted onto the skull near the pterygoid laminae. In an
1875; Knott 1883a). Teixeira et al. (2019) describe a longi- infant with mandibulofacial dysostosis, the superior con-
tudinal bundle that originates bilaterally from the occipital strictor and palatopharyngeus were continuous at their pos-
bone near the pharyngeal tubercle and descends to blend with terior and inferior attachments and inserted together into
the inferior constrictor. The authors suggest that these are the posterior raphe (Herring et al. 1979).
aberrant slips of the superior constrictor. The description of In an infant with trisomy 18, the posterior belly of
these slips is similar to that of azygos-pharyngeus. Sakamoto the digastric sent a slip to the superior constrictor bilat-
(2009) describes longitudinal bundles situated medial to sty- erally (Bersu and Ramirez-Castro 1977). Mieden (1982)
lopharyngeus that passed over the dorsal surface of the supe- describes an infant with median cleft lip, hypotelorism,
rior constrictor and blended with middle constrictor. Teixeira and alobar holoprosencephaly. This infant had a bilateral
et al. (2019) state that the bundles found by Sakamoto (2009) accessory slip extending between the intermediate tendon
do not resemble the bundles described by them. of digastricus and the superior pharyngeal constrictor
Additionally, constrictor pharyngis superior is associ- (Mieden 1982).
ated with Passavant’s muscle (palatopharyngeal sphincter) In a male neonate with Meckel syndrome, Pettersen
(Whillis 1930; Sakamoto 2016a; Standring 2016; Sumida (1984) observed that the superior pharyngeal constrictor
et al. 2017). Its existence is debated in the literature attached to an aberrant medial extension of the left man-
(Standring 2016; Sumida et al. 2017). Some consider dible. Itoh et al. (1991) describe fetal akinesia/hypokine-
Passavant’s muscle to be a portion of the superior pha- sia sequence in one male and one female infant, each with
ryngeal constrictor and/or palatopharyngeus muscles, suite of anatomical anomalies. In the male, the constrictor
perhaps due to the difficulty in distinguishing it from the pharyngis muscles showed irregular but mild atrophy. In an
former (Whillis 1930; Sumida et al. 2012; Sakamoto 2015; infant with craniorachischisis, the pharyngeal constrictor
Standring 2016; Sumida et al. 2017). Others recognize it muscles were absent (Alghamdi et al. 2017).
as a distinct, sphincter-like muscle that originates from the
nasal aspect of the lateral palatine aponeurosis and passes Prevalence
dorsolateral to levator veli palatini to surround the pharynx In a sample of 16 sides from eight infants with trisomy 18,
(Sakamoto 2016a; Standring 2016; Sumida et al. 2017). Bersu and Ramirez-Castro (1977) found a slip between the
posterior belly of the digastricus and the superior or middle
Prevalence pharyngeal constrictors on four sides (25%).
Knott (1883a) found azygos-pharyngeus in 4 out of 87
subjects (4.6%). In a sample of 110 cadavers, the ptery- Clinical Implications
gomandibular raphe was absent on 40 sides (36%), which N/A
allowed for complete continuity of the superior constric-
tor and buccinator muscles. In a sample of 44 sides from Stylopharyngeus (Figure 2.6)
22 cadavers, Sakamoto (2009) found a longitudinal bundle
that descended over the superior constrictor to blend with Synonyms
the middle pharyngeal constrictor on six sides (13.6%). N/A
Sakamoto (2009) also found that fibers originated from
the petrous temporal bone, which passed dorsally to merge Typical Presentation
with the superior constrictor on eight sides (18.2%) and
passed ventrally to attach to the soft palate with the ori- Description
gin of the superior constrictor on five sides (11.4%). In a Stylopharyngeus originates from the base of the styloid
sample of 44 sides, Choi et al. (2020) found that stylopha- process and ends with attachments to the pharyngeal
ryngeus passed transversely and merged with the superior constrictors, palatopharyngeus, and the thyroid cartilage
pharyngeal constrictor in one case (2.3%). Furthermore, in (Standring 2016).
Innervation side and inserted deep into hyoglossus (Pettersen 1979). In an

Stylopharyngeus is innervated by the glossopharyngeal infant with trisomy 18, an abnormal belly of stylopharyngeus
nerve (Standring 2016). coursed posteriorly behind the styloid process and inserted
over the superfcial surface of the superior constrictor (Bersu
Comparative Anatomy and Ramirez-Castro 1977). Stylopharyngeus was absent in a
Stylopharyngeus has a similar typical presentation in the fetus with craniorachischisis (Alghamdi et al. 2017).
apes, extending from the styloid process to the pharyngeal
wall, passing between the superior and middle pharyngeal Prevalence
constrictors (Kohlbrügge 1890–1892; Dean 1984; Gibbs In their literature review, Smith et al. (2015) found that a
1999; Diogo et al. 2010, 2012, 2013a,b, 2017). In one gib- doubled or extra stylopharyngeus muscle was present in 2
bon, it originated from the tympanic region as no styloid out of 20 individuals with trisomy 13 (10%).
process was present (Diogo et al. 2012). It may be indirectly
connected via connective tissue to the greater horn of the Clinical Implications
hyoid bone in gorillas (Diogo et al. 2010). It may have an N/A
insertion into the hyoid bone and/or the thyroid cartilage
in common chimpanzees (Gratiolet and Alix 1866; Diogo petrOpharyngeus (figure 2.6)
et al. 2013a). In one bonobo, Diogo et al. (2017) found that Synonyms
stylopharyngeus was fused with styloglossus and the mid-
Macalister (1875) and Knott (1883a) consider petropha-
dle constrictor.
ryngeus to be a variant of cephalo-pharyngeus. Sakamoto
Variations (2016a) suggests that petropharyngeus with an origin from
the vaginal process of the temporal bone would be more
Description
aptly named tympanopharyngeus.
Stylopharyngeus may be doubled or split into two or three
slips (Wood 1867b; Macalister 1875; Bergman et al. 1988; Typical Presentation
Sakamoto 2016a). It may course transversely and join the This muscle is present only as a variation or anomaly.
superior pharyngeal constrictor (Choi et al. 2020). An
accessory bundle of stylopharyngeus may either descend Comparative Anatomy
parallel to stylopharyngeus and pass between the middle Petropharyngeus is not present as a distinct muscle in the
and inferior pharyngeal constrictors or insert into the pha- apes (Diogo et al. 2010, 2012, 2013a,b, 2017).
ryngeal raphe along the midline with the superior constric-
tor (Choi et al. 2020). A bundle may also connect it to the Variations
intermediate tendon of the digastric (Sakamoto 2009). Description
Prevalence Petropharyngeus originates from the petrous temporal
bone and can insert into any of the pharyngeal constric-
In a sample of 44 sides from 22 cadavers, Sakamoto (2009)
tors (Macalister 1875; Knott 1883a; Shimada et al. 1991;
found a bundle connecting the intermediate tendon of the
Sakamoto 2009, 2016a; Siddiqui et al. 2017; Choi et al. 2020).
digastric and stylopharyngeus on one side (2.3%). In a
It may also originate from the vaginal process of the tempo-
sample of 44 sides, Choi et al. (2020) found that stylopha-
ral bone (Knott 1883a; Bergman et al. 1988; Sakamoto 2009,
ryngeus passed transversely and merged with the superior
2016a). It is situated medially, superiorly, and posteriorly to
pharyngeal constrictor in one case (2.3%). An accessory
stylopharyngeus (Siddiqui et al. 2017; Choi et al. 2020).
bundle of stylopharyngeus was present in eight cases
Petropharyngeus may be present bilaterally (Shimada
(18.2%). In four cases (9.1%), the bundle passed between the
et al. 1991; Sakamoto 2009; Siddiqui et al. 2017). It may be
middle and inferior pharyngeal constrictors. In four cases
divided into two bundles (Choi et al. 2020). It may present
(9.1%), the bundle inserted into the pharyngeal raphe with
as a thin and small fascicle, a fbromuscular structure, or
the superior constrictor.
a well-developed muscular slip (Siddiqui et al. 2017; Choi
et al. 2020).
Anomalies Petropharyngeus often inserts into the superior con-
Description strictor (Macalister 1875; Knott 1883a; Sakamoto 2009;
In one neonate with trisomy 13, the right posterior belly Siddiqui et al. 2017; Choi et al. 2020). Shimada et al. (1991)
of the digastricus split and sent a deep bundle to stylopha- observed that petropharyngeus inserted into the outer sur-
ryngeus (Colacino and Pettersen 1978). In another neonate face of the middle pharyngeal constrictor. In the case of a
with trisomy 13, stylopharyngeus was doubled bilaterally bifurcated petropharyngeus, Choi et al. (2020) found that
(Pettersen et al. 1979). The extra muscles coursed trans- one bundle passed deep to the superior constrictor and the
versely to the superior pharyngeal constrictor and inserted other inserted into the middle constrictor. Sakamoto (2009)
just lateral to the pharyngeal raphe. In a boy with trisomy found that petropharyngeus may end over the dorsal surface
13q, an extra stylopharyngeus muscle was present on the left of the inferior constrictor.
Innervation angle between the greater and lesser horns of the hyoid in
Petropharyngeus may be innervated by the glossopharyn- common chimpanzees and bonobos (Gratiolet and Alix
geal nerve (Shimada et al. 1991) or the pharyngeal plexus 1866; Sonntag 1923; Gibbs 1999; Diogo et al. 2013a, 2017).
(Sakamoto 2009). Therefore, it appears that only the ceratopharyngeal part of
the middle constrictor is present in the apes. It was fused
Prevalence with stylopharyngeus in one bonobo (Diogo et al. 2017).
In a sample of 614 Japanese cadavers, Shimada et al.
(1991) found petropharyngeus in seven cadavers (1.1%). In Variations
a sample of 44 sides from 22 cadavers, Sakamoto (2009)
found bundles that correspond to petropharyngeus on both Description
sides of one cadaver (4.5%). In a sample of 24 sides from Constrictor pharyngis medius may extend to the base of the
12 cadavers, Tubbs et al. (2010) found petropharyngeus on cranium along the stylohyoid ligament (Macalister 1875).
one side (4.2%). In a sample of 44 heads, Siddiqui et al. The origin from the greater horn of the hyoid (ceratopha-
(2017) found petropharyngeus in three cases (6.8%). In a ryngeal part) and the rest of the muscle (chondropharyngeal
sample of 44 sides, Choi et al. (2020) found petropharyn- part) may be distinct (Bergman et al. 1988). If the center
geus in 11 cases (25%). portion is defcient, it may appear bipartite (Macalister
1875). Macalister (1875) reports that it may have three
Anomalies parts, with origins from the hyoid, the triticeal cartilage,
and the thyrohyoid membrane.
Description Constrictor pharyngis medius may be fused with the
In a child with trisomy 21, petropharyngeus was present superior belly of omohyoid or sternothyroid (Wood 1868).
bilaterally (Bersu 1980). It extended between the petrous Its fbers may also connect with hyoglossus, stylohyoid, thy-
part of the temporal bone and the middle pharyngeal rohyoid, the posterior belly of the digastric, or the triticeal
constrictor. cartilage (Macalister 1875; Sakamoto 2009, 2014). Its fbers
may interdigitate with those of stylopharyngeus (Sakamoto
Prevalence
2014). Accessory muscles, such as sphenopharyngeus
Bersu (1980) found petropharyngeus in one out of fve indi- (Wood 1868) or petropharyngeus (Macalister 1875; see the
viduals with trisomy 21 (20%). entry for this muscle), may insert into the middle constric-
tor. Sakamoto (2009) describes longitudinal bundles situ-
ated medial to stylopharyngeus that passed over the dorsal
N/A surface of the superior constrictor and blended with the
middle constrictor.
cOnstrictOr pharyngis MeDius (MiDDle
Prevalence
pharyngeal cOnstrictOr) (figure 2.6)
In a sample of 44 sides from 22 cadavers, the superior fbers
See also: Petropharyngeus of constrictor pharyngis medius joined with hyoglossus on
ten sides (22.7%) and with stylohyoid on fve sides (11.4%)
Synonyms (Sakamoto 2009). A longitudinal bundle that descended
N/A over the superior constrictor to blend with the middle pha-
ryngeal constrictor was present on six sides (13.6%). In
Typical Presentation a sample of 82 sides from 41 cadavers, Sakamoto (2014)
Description found fbers of the middle constrictor attached to hyoglos-
Constrictor pharyngis medius originates from the lesser sus on 29 sides (35.4%), thyrohyoid on two sides (2.4%), sty-
and greater horns of the hyoid, and from the stylohyoid lohyoid and the posterior belly of the digastric on 33 sides
ligament, and inserts into the pharyngeal raphe (Standring (40.2%), only stylohyoid on seven sides (8.5%), and the
2016). triticeal cartilage on one side (1.2%). Fibers of the middle
constrictor interdigitated with those of stylopharyngeus on
Innervation four sides (4.9%).
Constrictor pharyngis medius is innervated by the pharyn-
geal plexus (Standring 2016). Anomalies
Comparative Anatomy Description

Constrictor pharyngis medius has an origin only from the In an infant with Hanhart syndrome, the right medial ptery-
greater horn of the hyoid in gibbons and gorillas (Kohlbrügge goid had extensive attachments and sent fbers to the mid-
1890–1892; Diogo et al. 2010, 2012). It originates from the dle pharyngeal constrictor (Bersu et al. 1976). In an infant
hyoid in orangutans (Sonntag 1924). It originates from the with trisomy 18, the posterior belly of the digastric sent a
slip to the middle constrictor on the right side (Bersu and
Ramirez-Castro 1977). Itoh et al. (1991) describe fetal akine- cricothyroid, or sternothyroid (Macalister 1875; Bergman
sia/hypokinesia sequence in one male and one female infant, et al. 1988; Sakamoto 2009, 2013, 2016a). Fibers of the les-
each with a suite of anatomical anomalies. In the male, the sor constrictor may originate from a tendinous cord that
constrictor pharyngis muscles showed irregular but mild loops over cricothyroid and connects the cricoid cartilage
atrophy. In an infant with craniorachischisis, the pharyngeal with the inferior thyroid tubercle (Sakamoto 2013, 2016a;
constrictor muscles were absent (Alghamdi et al. 2017). Standring 2016). When this tendinous cord is absent or def-
cient, the inferior constrictor may connect with cricothyroid
Prevalence (Sakamoto 2013, 2016a).
In a sample of 16 sides from eight infants with trisomy 18, Constrictor pharyngis inferior may receive a slip from
Bersu and Ramirez-Castro (1977) found a slip between the the lateral thyrohyoid ligament, which may be referred to as
posterior belly of the digastricus and the superior or middle syndesmopharyngeus (Knott 1883a; Bergman et al. 1988;
pharyngeal constrictors on four sides (25%). Sakamoto 2016a). Petropharyngeus may end over the dorsal
surface of the inferior constrictor (Sakamoto 2009; see the
Clinical Implications entry for this muscle). Teixeira et al. (2019) describe a lon-
N/A gitudinal bundle that originates bilaterally from the occipi-
tal bone near the pharyngeal tubercle and descend to blend
with the inferior constrictor. The authors suggest that these
cOnstrictOr pharyngis inferiOr (inferiOr
are aberrant slips of the superior constrictor. The descrip-
pharyngeal cOnstrictOr) (figure 2.6)
tion of these slips is similar to that of the azygos-pharyn-
See also: Petropharyngeus geus muscle (Macalister 1875; Knott 1883a; Bergman et al.
1988; Sakamoto 2016a).
Synonyms
N/A
Prevalence
Typical Presentation Knott (1883a) found syndesmopharyngeus in 2 out of 47
subjects (4.3%). In a sample of 44 sides from 22 cadavers,
Description the tendinous cord connected the two origins of the inferior
Constrictor pharyngis inferior has thyropharyngeal and cri- constrictor on 10 sides (22.7%) (Sakamoto 2009). Fibers of
copharyngeal parts (Standring 2016). The former originates the inferior constrictor connected with thyrohyoid on 13
from the thyroid cartilage and inserts into the pharyngeal sides (29.5%), sternothyroid on 29 sides (65.9%), and crico-
raphe (Standring 2016). The latter originates from the cri- thyroid on 17 sides (38.6%). In a sample of 60 sides from 30
coid cartilage and merges inferiorly with some fbers of the cadavers, the tendinous cord was present on 36 sides (60%)
esophagus (Standring 2016). (Sakamoto 2013).
Innervation Anomalies
Constrictor pharyngis inferior is innervated by the pha- Description
ryngeal plexus (Standring 2016). It may receive additional Itoh et al. (1991) describe fetal akinesia/hypokinesia
Innervation from one or more of the laryngeal nerves sequence in one male and one female infant, each with a
(Sakamoto 2013). suite of anatomical anomalies. In the male, the constrictor
pharyngis muscles showed irregular but mild atrophy. In an
Comparative Anatomy infant with craniorachischisis, the pharyngeal constrictor
Constrictor pharyngis inferior has a similar typical presen- muscles were absent (Alghamdi et al. 2017).
tation in the apes, as both the thyropharyngeal and crico-
pharyngeal parts are typically present with insertions into Prevalence
the pharyngeal raphe (Kohlbrügge 1890–1892; Hosokawa N/A
and Kamiya 1961–1962; Gibbs 1999; Diogo et al. 2010,
2012, 2013a,b, 2017). It may have an origin from the frst Clinical Implications
tracheal ring in common chimpanzees (Gratiolet and Alix N/A
1866; Gibbs 1999).
Variations LARYNGEAL MUSCLES
Description cricOthyrOiDeus (cricOthyrOiD) (figure 2.7)

Constrictor pharyngis inferior may have an origin from the See also: Ceratocricoid, Thyrotrachealis
trachea (Macalister 1875). The thyropharyngeal and cri-
copharyngeal parts may be distinct (Bergman et al. 1988). Synonyms
The inferior constrictor may connect with thyrohyoid, N/A
FIGURE 2.7 Anterior laryngeal muscles in anterior view.
Typical Presentation third, horizontal belly situated deep to the oblique belly.
Cricothyroid may have an origin from the frst tracheal
Description ring (Macalister 1875; Maranillo and Sanudo 2016). It may
Cricothyroid originates from the cricoid cartilage and connect with thyrohyoid, sternothyroid, or the inferior pha-
inserts onto the inferior cornu (oblique part) and lower mar- ryngeal constrictor (Macalister 1875; Bergman et al. 1988;
gin (straight part) of the thyroid cartilage (Standring 2016). Sakamoto 2009, 2013, 2016a; Maranillo and Sanudo 2016).
The cricothyroid muscles may decussate across the midline
Innervation (Bergman et al. 1988). If the straight part is separate from
Cricothyroid is innervated by the external branch of the the rest of the muscle, it may be referred to as cricothy-
superior laryngeal nerve (Standring 2016). roideus superior (Macalister 1875; Bergman et al. 1988;
Maranillo and Sanudo 2016).
Comparative Anatomy Cricothyroid is associated with several named accessory
Cricothyroid has a similar typical presentation in the apes, muscles. Cricothyroid may connect with ceratocricoid, a
extending from the cricoid cartilage to the thyroid cartilage small bundle that originates from the cricoid cartilage and
(Kohlbrügge 1890–1892; Saban 1968; Gibbs 1999; Diogo inserts into the inferior cornu of the thyroid cartilage (see
et al. 2010, 2012, 2013a,b, 2017). Some authors state that in the entry for this muscle). It may connect with thyrotrachea-
addition to a straight part and oblique part, a pars interna lis, which extends from the thyroid cartilage to the trachea
is also often present in the apes (Gratiolet and Alix 1866; (see the entry for this muscle). It may also connect with the
Kohlbrügge 1890–1892; Starck and Schneider 1960; Saban internal cricoid muscle, which is situated medial to crico-
1968; Gibbs 1999; Diogo et al. 2010, 2012, 2013a,b, 2017). thyroid and has attachments to the cricoid cartilage and
Macalister (1875) states that Eschricht found cricothyroi- thyroid cartilage (Le Double 1897; Maranillo and Sanudo
deus superior and a muscle similar to thyroideus transver- 2016). The deep levator of the thyroid gland may originate
sus anomalus in gibbons. from the anteromedial aspect of cricothyroid (Maranillo
and Sanudo 2016).
Variations Incisurae mediae obliquus (Gruber) may be present as a
small muscle on the inferolateral aspect of the thyroid car-
Description tilage (Gruber 1868a; Bergman et al. 1988; Maranillo and
Cricothyroid may be bilaminar (Macalister 1875). Mu Sanudo 2016). It may be present bilaterally (Bergman et al.
and Sanders (2008) suggest that cricothyroid has a 1988). It may have a second head that originates from the
sternothyroid, and in this case, the muscle would be referred It may have medial and lateral heads (Macalister 1875;
to as incisurae mediae obliquus bicaudatus (Bergman et al. Bergman et al. 1988; Maranillo and Sanudo 2016). It may
1988; Maranillo and Sanudo 2016). Thyroideus transversus be a detached slip of sternothyroid (Macalister 1875). It
anomalus (Gruber) extends across the superior portion of may merge with cricothyroid or the inferior pharyngeal
the cricothyroid membrane from one side of the inferior constrictor (Sujata et al. 2013). Sujata et al. (2013) found an
border of the thyroid cartilage to another (Macalister 1875; insertion into the ffth through eighth tracheal rings.
Knott 1883a; Maranillo and Sanudo 2016). It may also be
referred to as thyroideus marginalis inferior, incisurae car- Innervation
tilaginis thyroideae mediae transversus, or the transverse Thyrotrachealis is innervated by a small twig from the ansa
thyroid muscle (Macalister 1875; Knott 1883a; Maranillo cervicalis (Sujata et al. 2013).
and Sanudo 2016). Cricohyoid (Zagorsky) extends from the
cricoid cartilage to the greater horn or body of the hyoid Prevalence
(Macalister 1875; Bergman et al. 1988; Maranillo and Gruber (1868c) found thyrotrachealis in 21 out of 80 cases
Sanudo 2016). Cricotrachealis originates from the lower (26.3%). Macalister (1875) found fve cases of thyrotrachea-
margin of the cricoid cartilage, courses behind the isthmus lis in 80 subjects (6.3%). Knott (1883a) found thyrotrachea-
of the thyroid gland, and inserts into the fourth and/or ffth lis in 3 out of 28 subjects (10.7%). Le Double (1897) found
tracheal ring (Macalister 1875; Knott 1883a; Maranillo and thyrotrachealis in 4 out of 60 cases (6.7%).
Sanudo 2016).
Anomalies
Prevalence N/A
In a sample of 44 sides from 22 cadavers, Sakamoto (2009) Clinical Implications
found fbers of the inferior constrictor connected with
Thyrotrachealis may lead to the misdiagnosis of thyroid
cricothyroid on 17 sides (38.6%).
swellings (Sujata et al. 2013).
Anomalies ceratOcricOiD (figure 2.8)

N/A
Synonyms
Clinical Implications This muscle may also be referred to as crico-corniculatus
N/A (Tourtual), kerato-cricoid (Merkel), Merkel’s muscle, or pos-
terior cricothyroid [crico-thyreoideus posticus (Bochdalek)]
(Turner 1860; Macalister 1875; Knott 1883a; Hetherington
thyrOtrachealis (figure 2.7) 1934).
Synonyms Typical Presentation

This muscle may also be referred to as thyrotrachealis pro- This muscle is present only as a variation or anomaly.
fundus (Krause) or thyrotrachealis biceps (Gruber) (Knott
1883a; Bergman et al. 1988). Comparative Anatomy
Among the primates, ceratocricoid has been observed in
Typical Presentation guenons (Hetherington 1934). It is not present in the apes
This muscle is present only as a variation. (Diogo et al. 2010, 2012, 2013a,b, 2017). In one bonobo, the
posterior cricoarytenoid muscle had an attachment onto the
Comparative Anatomy inferior cornu of the thyroid cartilage, but these fbers did
Gratiolet and Alix (1866) describe an additional bundle of not form a distinct ceratocricoid muscle (Diogo et al. 2017).
cricothyroid in one common chimpanzee that could cor-
Variations
respond to thyrotrachealis. It originated from the inferior
margin of the thyroid cartilage, passed superfcial to crico- Description
thyroid, and ended in the more anterior tracheal cartilages. Ceratocricoid originates from the posterior surface of
the cricoid cartilage, behind and below the cricothyroid
Variations joint, and inserts into the posterior surface of the inferior
cornu of the thyroid cartilage (Turner 1860; Macalister
Description 1875; Hetherington 1934; Bergman et al. 1988; Sharp
Thyrotrachealis originates from the lower border of the thy- 1990; Maranillo et al. 2009; Maranillo and Sanudo 2016;
roid cartilage, courses over cricothyroid and the isthmus of Standring 2016). It is situated lateral to, and near the
the thyroid gland, and inserts into the third, fourth, and/or lower border of, cricoarytenoideus posterior (Bergman
ffth tracheal rings (Macalister 1875; Knott 1883a; Bergman et al. 1988; Hetherington 1934; Maranillo and Sanudo 2016;
et al. 1988; Sujata et al. 2013; Maranillo and Sanudo 2016). Standring 2016). When present, it courses over the recurrent
FIGURE 2.8 Posterior laryngeal muscles in posterior view.
laryngeal nerve (Turner 1860; Hetherington 1934; Sharp Anomalies

1990; Maranillo et al. 2009; Maranillo and Sanudo 2016). Description
Ceratocricoid is present as a condensation of cells at embry-
Ceratocricoid was present in an infant with mandibulofa-
onic stages 22 and 23, at which point it is joined to the poste-
cial dysostosis (Herring et al. 1979).
rior cricoarytenoid muscle but extends toward the chondrifed
inferior cornu (Maranillo et al. 2009). It becomes completely Prevalence
distinct from the posterior cricoarytenoid muscle in the elev- N/A
enth week of fetal development (Maranillo et al. 2009).
Ceratocricoid may be fused with cricothyroid or the pos- Clinical Implications
terior cricoarytenoid muscle (Macalister 1875; Maranillo and Ceratocricoid may compress the recurrent laryngeal nerve,
Sanudo 2016). It may also have an attachment into the capsule and thus be a potential cause of recurrent laryngeal nerve
of the cricothyroid joint (Hetherington 1934; Sharp 1990; palsy (Maranillo et al. 2009).
Maranillo et al. 2009). It may have two bellies (Hetherington
1934; Maranillo et al. 2009). It may also be divided only at
its insertion into the thyroid cartilage (Hetherington 1934). cricOarytenOiDeus pOsteriOr (pOsteriOr
It may be present bilaterally (Turner 1860; Macalister 1875; cricOarytenOiD) (figure 2.8)
Knott 1883a; Hetherington 1934; Maranillo et al. 2009).
See also: Ceratocricoid
Innervation
Synonyms
Ceratocricoid is innervated by the recurrent laryngeal nerve
(Sharp 1990; Maranillo et al. 2009). This muscle may also be referred to as cricoarytenoideus
posticus (Macalister 1875).
Prevalence
Turner (1860) found seven cases of ceratocricoid in 32 sub-
jects (21.9%). Le Double (1897) found fve cases of cer- Description
atocricoid out of 28 specimens (17.9%). In a review of the Cricoarytenoideus posterior originates from the posterior
literature, Hetherington (1934) reports that the prevalence surface of the cricoid cartilage and inserts into the muscular
of ceratocricoid ranges from 10% to 25%. In a sample of process of the arytenoid cartilage (Standring 2016). It has
132 subjects, Hetherington (1934) found ceratocricoid in 22 three muscular compartments: horizontal (medial), oblique,
subjects (16.7%). In a sample of 134 hemilarynges, Sharp and vertical (lateral) (Sanders et al. 1994; Maranillo and
(1990) found nine cases of ceratocricoid (6.7%). Schweizer Sanudo 2016; Standring 2016).
and Dörf (1997) found ceratocricoid in 4 out of 21 larynges
(19%). Maranillo et al. (2009) found ceratocricoid present in Innervation
8 out of 34 fetal larynges (23.5%) and in 13 out of 90 adult Cricoarytenoideus posterior is innervated by the recurrent
larynges (14.4%). laryngeal nerve (Standring 2016).
Comparative Anatomy In a fetus with craniorachischisis, the posterior cricoary-

Cricoarytenoideus posterior has a similar typical pre- tenoid muscles were attached at the midline (Alghamdi
sentation in the apes, extending from the dorsal portion et al. 2017).
of the cricoid cartilage to the arytenoid cartilage (Diogo
et al. 2010, 2012, 2013a,b, 2017). The posterior cricoary- Prevalence
tenoid muscles may meet at the midline in some gibbons N/A
(Kohlbrügge 1890–1892; Starck and Schneider 1960; Diogo
et al. 2012), orangutans (Sonntag 1924), and common chim-
panzees (Jordan 1971a,b,c; Diogo et al. 2013a). The cerato- N/A
arytenoideus lateralis muscle (ceratoarytenoid) is distinctly
present in some common chimpanzees (Macalister 1871; cricOarytenOiDeus lateralis
Saban 1968). Bundles that potentially correspond to cera- (lateral cricOarytenOiD) (figure 2.9)
toarytenoid have been described in gibbons (Kohlbrügge Synonyms
1890–1892; Starck and Schneider 1960; Diogo et al. 2012).
N/A
Variations Typical Presentation

Cricoarytenoideus posterior (the posterior cricoarytenoid Cricoarytenoideus lateralis originates from the upper border
muscle) may be divided (Macalister 1875). It may also be of the arch of the cricoid cartilage and inserts into the mus-
bilaminar (Macalister 1875). It may send a slip to the cri- cular process of the arytenoid cartilage (Standring 2016).
cothyroid joint (Macalister 1875; Maranillo and Sanudo
2016). The posterior cricoarytenoid muscle is associated Innervation
with a few accessory muscles. It may be joined to the Cricoarytenoideus lateralis is innervated by the recurrent
ceratocricoid muscle (Macalister 1875; Maranillo and laryngeal nerve (Standring 2016).
Sanudo 2016; see the entry for this muscle). It may also
be associated with ceratoarytenoid (also referred to as Comparative Anatomy
posterior thyroarytenoid, accessory thyroarytenoid, or Cricoarytenoideus lateralis has a similar typical presenta-
ceratoarytenoideus lateralis), which extends between tion in the apes, extending from the anterior portion of the
the inferior cornu of the thyroid cartilage and the mus- cricoid cartilage to the muscular process of the arytenoid
cular process of the arytenoid cartilage (Gruber 1868b; cartilage (Gibbs 1999; Diogo et al. 2010, 2012, 2013a,b,
Macalister 1867b, 1875; Knott 1883a; Le Double 1897; 2017). There may be an additional insertion onto the thy-
Hetherington 1934; Saban 1968; Maranillo and Sanudo roid cartilage in orangutans and common chimpanzees
2016). A cricocorniculate muscle may be present that (Jordan 1971a; Gibbs 1999).
originates from the upper margin of the cricoid cartilage
and inserts into the corniculate cartilage (Maranillo and
Variations
Sanudo 2016).
Wells and Thomas (1927) describe an accessory muscle Description
of posterior cricoarytenoid that is referred to as the superior Cricoarytenoideus lateralis (the lateral cricoarytenoid mus-
cricoarytenoid muscle by Maranillo and Sanudo (2016). cle) may receive a slip from the thyroid cartilage (Macalister
It originated from the superior part of the left lamina of 1875). It may be fused with thyroarytenoideus (Bergman et al.
the cricoid cartilage, just lateral to the median ridge. The 1988). The extent of its attachment on the arytenoid cartilage
muscle coursed superiorly and laterally over the transverse may vary (Mossallam et al. 1987).
and oblique arytenoid muscles. It inserted on the right side The lateral cricoarytenoid muscle is associated with
of the larynx into the apex of the arytenoid cartilage and several accessory muscles. It may be connected with cri-
the mucous membrane of the aryepiglottic fold, with some coepiglotticus, which originates from the inner surface of
fbers inserting into the transverse arytenoid (Wells and the cricoid cartilage near the lateral cricoarytenoid and
Thomas 1927; Maranillo and Sanudo 2016). attaches to the epiglottis (Knott 1883a; Maranillo and
Sanudo 2016). A similar slip termed cricomembranosus
Prevalence (cricomembranous muscle) may pass to the quadrangular
Hetherington (1934) found ceratoarytenoid in 3 out of 66 membrane, which extends between the epiglottis and the
individuals (4.5%). arytenoid cartilage (Knott 1883a; Maranillo and Sanudo
2016). The lateral cricoarytenoid muscle may send a bundle
Anomalies to the thyroarytenoid muscle and thyroid cartilage termed
Description the internal lateral cricothyroid muscle (Maranillo and
In cases of laryngeal cleft, the posterior cricoarytenoid Sanudo 2016). Syndesmoarytenoid may extend from the
muscles may be defcient or displaced (Lim et al. 1979). muscular process of the arytenoid cartilage and the lateral
cricoarytenoid to the cricothyroid ligament (Maranillo and 1866; Sonntag 1923; Starck and Schneider 1960; Gibbs
Sanudo 2016). Arythyrocricoid may originate from the 1999; Diogo et al. 2010, 2013a,b).
transverse or oblique arytenoid muscles and insert into lat-
eral cricoarytenoid (Maranillo et al. 2011).
Variations
Wells and Thomas (1927) describe an accessory muscle
of lateral cricoarytenoid that is referred to as the cricothy- Description
rohyoid muscle by Maranillo and Sanudo (2016). It was Arytenoideus transversus (the transverse arytenoid) may
situated on the lateral aspect of the larynx on the left side send fbers to thyroarytenoid (Macalister 1875; Kanthack
of the body. It originated from the superolateral angle of 1892). The direction of its fbers or the proportion of its
the cricoid cartilage, beneath the lateral cricoarytenoid fbers relative to the oblique arytenoid may vary (Macalister
muscle. The muscle coursed superiorly and anteriorly over 1875; Maranillo and Sanudo 2016). The transverse aryte-
thyroarytenoideus and thyroepiglotticus to insert into a noid may connect with the thyroarytenoid or the lateral
band of fascia that was continuous with the median thy- cricoarytenoid via a fascicle referred to as the arythyrocri-
rohyoid ligament (Wells and Thomas 1927; Maranillo and coid (Maranillo et al. 2011). It may also connect with the
Sanudo 2016). straight arycorniculate muscle (aryteno-corniculatus rec-
tus), which lies deep to the transverse arytenoid and extends
Prevalence from the base of the arytenoid cartilage to the corniculate
Krause found cricoepiglotticus in 34% of cases, and Knott cartilage (Luschka 1869; Macalister 1875; Le Double 1897;
found this muscle in 3 out of 19 subjects (15.8%) (Knott Maranillo and Sanudo 2016).
1883a).
Prevalence
Anomalies Maranillo et al. (2011) found the arythyrocricoid fasci-
N/A cle in 29 out of 30 larynges (96.7%), and due to bilateral
presence in some cases, a total of 47 arythyrocricoid fas-
Clinical Implications cicles were found. Arythyrocricoid originated from the
N/A transverse arytenoid in 16 cases (34%) and from both
the transverse and oblique arytenoid muscles in 18 cases
arytenOiDeus transversus (38.3%). Arythyrocricoid inserted into the thyroaryte-
noid muscle in 18 cases (38.3%), the lateral cricoaryte-
(transverse arytenOiD) (figure 2.8) noid muscle in 5 cases (10.6%), and into both muscles in
Synonyms 24 cases (51.1%).
Together with arytenoideus obliquus, this muscle may be
referred to as arytenoideus proprius (Macalister 1875) or Anomalies
interarytenoideus (Kanthack 1892). Description
Typical Presentation In cases of laryngeal cleft, the interarytenoid may show
partial agenesis (Lim et al. 1979). In an infant with mandib-
Description ulofacial dysostosis, the transverse arytenoid was enlarged
Arytenoideus transversus is a single muscle that extends (Herring et al. 1979).
between the muscular processes and lateral borders of the
two arytenoid cartilages (Standring 2016). It is situated Prevalence
deep into the arytenoideus obliquus muscles on the poste- N/A
rior surface of the larynx (Standring 2016).
Innervation The presence of arythyrocricoid may affect vocal fold
Arytenoideus transversus is innervated by the recurrent position following laryngeal nerve palsy or interfere with
laryngeal nerves (Standring 2016). electromyographic testing of laryngeal nerve function
(Maranillo et al. 2011).
Comparative Anatomy
In gibbons and bonobos, arytenoideus is present as a single,
unpaired muscle that extends between the two arytenoid arytenOiDeus Obliquus (Oblique
cartilages, and is not divided into an arytenoideus trans-
arytenOiD) (figure 2.8)
versus and arytenoideus obliquus (Kohlbrügge 1890–1892;
Starck and Schneider 1960; Diogo et al. 2012, 2017). In Synonyms
orangutans, gorillas, and common chimpanzees, the aryte- Together with arytenoideus transversus, this muscle may be
noideus transversus is present in all cases and the arytenoi- referred to as arytenoideus proprius (Macalister 1875) or
deus obliquus is present in some cases (Gratiolet and Alix interarytenoideus (Kanthack 1892).
Typical Presentation connect with the thyroarytenoid or the lateral cricoary-

Description tenoid via a fascicle referred to as the arythyrocricoid
(Maranillo et al. 2011). The oblique arycorniculate mus-
The arytenoideus obliquus muscles cross each other,
cle (aryteno-corniculatus obliquus) is a fascicle of aryte-
extending from the muscular process of one arytenoid car-
noideus obliquus that inserts into the corniculate cartilage
tilage to the apex of the contralateral arytenoid cartilage
(Macalister 1875; Le Double 1897; Maranillo and Sanudo
(Standring 2016). Fibers of oblique arytenoid that extend
2016). The arymembranous muscle may extend from
from the apex of the arytenoid cartilage to the aryepiglot-
the arytenoid cartilage to the quadrangular membrane
tic fold are referred to as aryepiglotticus (Standring 2016)
(Maranillo and Sanudo 2016).
or arytenoepiglottic muscles (Maranillo and Sanudo 2016).
Innervation
Prevalence
Arytenoideus obliquus is innervated by the recurrent laryn-
Maranillo et al. (2011) found the arythyrocricoid fascicle
geal nerve (Standring 2016).
in 29 out of 30 larynges (96.7%), and due to bilateral
Comparative Anatomy presence in some cases, a total of 47 arythyrocricoid
In gibbons and bonobos, arytenoideus is present as a single, fascicles were found. Arythyrocricoid originated from
unpaired muscle that extends between the two arytenoid the oblique arytenoid in 13 cases (27.7%) and from both
cartilages, and is not typically divided into an arytenoi- the transverse and oblique arytenoid muscles in 18 cases
deus transversus and arytenoideus obliquus (Kohlbrügge (38.3%). Arythyrocricoid inserted into the thyroaryte-
1890–1892; Starck and Schneider 1960; Diogo et al. 2012, noid muscle in 18 cases (38.3%), the lateral cricoaryte-
2017). In orangutans, gorillas, and common chimpanzees, noid muscle in 5 cases (10.6%), and both muscles in 24
the arytenoideus transversus is present in all cases, and the cases (51.1%).
arytenoideus obliquus is present in some cases (Gratiolet
and Alix 1866; Sonntag 1923; Starck and Schneider 1960; Anomalies
Gibbs 1999; Diogo et al. 2010, 2013a,b). Arytenoideus
Description
obliquus may blend with thyroarytenoideus in chimpanzees
(Jordan 1971a; Gibbs 1999). In cases of laryngeal cleft, the interarytenoid may show
partial agenesis (Lim et al. 1979).
Variations Prevalence
Description N/A
One or both of the arytenoideus obliquus muscles (the
oblique arytenoid muscles) may be absent (Macalister 1875;
Maranillo and Sanudo 2016). Its proportion relative to the The presence of arythyrocricoid may affect vocal fold
transverse arytenoid may vary (Macalister 1875). Some position following laryngeal nerve palsy or interfere with
fbers may connect with thyroarytenoid (Macalister 1875; electromyographic testing of laryngeal nerve function
Kanthack 1892). It may also connect to the cricoid cartilage (Maranillo et al. 2011).
or the corniculate cartilage (Macalister 1875; Maranillo and
Sanudo 2016).
The oblique arytenoid muscles are associated with thyrOarytenOiDeus (thyrOarytenOiD) (figure 2.9)
a few accessory muscles. The oblique arytenoids may See also: Superior thyroarytenoideus
FIGURE 2.9 Laryngeal muscles in superior view.

Synonyms (the ventricular muscle), a muscle situated in the vestibular

N/A fold with attachments to the angle of the thyroid cartilage and
the arytenoid cartilage (Kanthack 1892; Kotby et al. 1991;
Typical Presentation Maranillo and Sanudo 2016). A fascicle of the superfcial
Description fbers of thyroarytenoid referred to as thyreo-corniculatus
Thyroarytenoideus originates from the angle of the thyroid (thyrocorniculate muscle) may insert into the corniculate
cartilage and the cricothyroid ligament and inserts into the cartilage (Knott 1883a; Maranillo and Sanudo 2016). A
arytenoid cartilage (Standring 2016). The deep fbers form similar fascicle termed thyreo-cuneiformis (thyrocuneiform
vocalis, which is situated between thyroarytenoideus and the muscle) may insert into the cuneiform cartilage (Knott 1883a;
vocal ligament, and attach to the vocal process of the aryte- Maranillo and Sanudo 2016). Thyroconoid may originate
noid cartilage (Standring 2016). Some fbers of vocalis insert from the thyroarytenoid and thyroid cartilage and insert into
into the vocal ligament (aryvocalis of Ludwig) (Bergman et al. the conoid ligament (Maranillo and Sanudo 2016). The infe-
1988; Standring 2016). The thyroepiglotticus muscle is formed rior thyroid muscle (thyroideus internus or subthyroideus of
by fbers of thyroarytenoideus that continue into aryepiglottic Krause) may extend from the lower aspect of the angle of
fold and end in the margin of the epiglottis (Standring 2016). the thyroid cartilage to the root of the inferior cornu (Knott
1883a; Le Double 1897; Maranillo and Sanudo 2016).
Innervation
Thyroarytenoideus is innervated by the recurrent laryngeal Prevalence
nerve (Standring 2016). Knott (1883a) found thyreo-corniculatus in 2 out of 19 cases
(10.5%) and the inferior thyroid muscle in 2 out of 43 cases
Comparative Anatomy (4.7%). In a sample of 20 larynges, Kotby et al. (1991) found
Thyroarytenoideus has a similar typical presentation in the the superior thyroarytenoid muscle bilaterally in 16 laryn-
apes, extending from the thyroid cartilage to the arytenoid ges (80%) and ventricularis in 19 larynges (95%). In a sam-
cartilage (Gibbs 1999; Diogo et al. 2010, 2012, 2013a,b, ple of 100 hemilarynges from 50 cadavers, Lee et al. (2018)
2017). In the great apes, both a pars superior (corresponding found superior thyroarytenoideus in 36 specimens (36%).
to thyroarytenoideus) and a pars inferior (or thyroarytenoi-
deus medialis, corresponding to vocalis) seem to be pres- Anomalies
ent in most specimens, though the development of the pars N/A
inferior may vary (Gibbs 1999; Starck and Schneider 1960;
Saban 1968; Diogo et al. 2010, 2013a,b, 2017). Due to con- Clinical Implications
ficting accounts in the literature, it is not clear whether a N/A
bundle corresponding to vocalis is typically present in gib-
bons (Diogo et al. 2012). Thyroepiglotticus may be present
in gorillas and common chimpanzees (Gratiolet and Alix superiOr thyrOarytenOiDeus (superiOr
1866; Saban 1968; Gibbs 1999; Diogo et al. 2010, 2013a). thyrOarytenOiD) (figure 2.9)
See also: Thyroarytenoideus

Variations
The extent to which thyroarytenoid and vocalis are distinct This muscle may also be referred to as oblique thyroaryte-
may vary (Macalister 1875). The extent of the origin from the noid (Lee et al. 2018).
thyroid cartilage may also vary (Maranillo and Sanudo 2016).
Thyroarytenoideus may be connected with cricoaryte- Typical Presentation
noideus lateralis, arytenoideus transversus, or arytenoideus This muscle is present only as a variation.
obliquus (Macalister 1875; Kanthack 1892; Bergman et al.
1988; Maranillo et al. 2011; Maranillo and Sanudo 2016). Comparative Anatomy
Thyroepiglotticus may be present as thyroepiglotticus N/A
major (inferior) and thyroepiglotticus minor (superior)
(Macalister 1875; Knott 1883a; Le Double 1897; Maranillo Variations
and Sanudo 2016). Both insert into the lateral border of the Description
epiglottis (Knott 1883a; Maranillo and Sanudo 2016). The superior thyroarytenoideus is situated lateral to thyro-
Thyroarytenoideus is associated with several acces- arytenoid, extending from the upper portion of the angle of
sory muscles. The superior thyroarytenoideus may be pres- the thyroid cartilage to the muscular process of the arytenoid
ent along its lateral surface (see the entry for this muscle). cartilage (Zemlin et al. 1984; Kotby et al. 1991; Bergman
Thyromembranosus is similar to thyroepiglotticus but inserts et al. 1988; Maranillo and Sanudo 2016; Standring 2016;
into the quadrangular membrane (Maranillo and Sanudo Lee et al. 2018). It may insert into the lateral cricoarytenoid
2016). Fibers of thyroarytenoideus may form ventricularis muscle or its superfcial fascia (Lee et al. 2018).
Innervation INFRAHYOID MUSCLES

Superior thyroarytenoideus is likely innervated by the
recurrent laryngeal nerve.
sternOhyOiDeus (sternOhyOiD) (figure 2.10)
See also: Cleidohyoid
Prevalence
Zemlin et al. (1984) found the superior thyroarytenoid mus- Synonyms
cle in 12 out of 15 larynges (80%). Kotby et al. (1991) found N/A
this muscle bilaterally in 16 out of 20 larynges (80%). In a
sample of 100 hemilarynges from 50 cadavers, Lee et al. Typical Presentation
(2018) found this muscle in 36 cases (36%), and it had an
Description
insertion into the lateral cricoarytenoid muscle or its fascia
in 8 cases (8%). Sternohyoideus originates from the manubrium, medial end
of the clavicle, and posterior sternoclavicular ligament and
Anomalies inserts into the inferior margin of the body of the hyoid
(Standring 2016).
N/A
Innervation
Clinical Implications Sternohyoideus is innervated by branches of the ansa cervi-
N/A calis (Standring 2016).
FIGURE 2.10 Suprahyoid and infrahyoid muscles in anterior view.

Comparative Anatomy males and present in 29 out of 230 sides (12.61%) in

Sternohyoideus has a similar typical presentation in the females. In males, one intersection was present in 21 out of
apes, extending from the sternum and adjacent regions to the the 23 sides (91.3%), and two intersections were present in
hyoid (Gratiolet and Alix 1866; Sonntag 1923, 1924; Raven two sides (8.7%). In females, one intersection was present in
1950; Miller 1952; Starck and Schneider 1960; Gibbs 1999; 19 out of the 29 sides (65.52%) and two intersections were
Diogo et al. 2010, 2012, 2013a,b, 2017). Tendinous intersec- present in ten sides (34.48%). Right and left sternohyoid
tions are found in gibbons, common chimpanzees, and bono- muscles were fused in 23 out of 169 males (13.61%) and in
bos but are not found in gorillas (Gratiolet and Alix 1866; 5 out of 115 females (4.35%).
Champneys 1872; Deniker 1885; Kohlbrügge 1890–1892;
Starck and Schneider 1960; Diogo et al. 2010, 2012, 2013a, Anomalies
2017). Sternohyoid may have an additional origin from the Description
frst and second ribs in gibbons (Kohlbrügge 1890–1892). In both trisomy 13 and trisomy 18 neonates, Aziz (1979,
1981) found sternohyoideus azygos, a supernumerary
slip that arose from the manubrium and inserted near
Variations
the intermediate tendon of the digastricus. Bersu and
Description Ramirez-Castro (1977) also found sternohyoideus azy-
One or two tendinous intersections may be present in sterno- gos that inserted into the hyoid bone in an infant with
hyoid (Hallett 1848; Macalister 1867b, 1875; Mori 1964; Sato trisomy 18. This same infant also had a doubled left
1968b; Standring 2016). The right and left sternohyoid mus- sternohyoid. The left sternohyoid was also doubled in a
cles may fuse (Macalister 1875; Sato 1968b; Lee and Yang second infant. In the third infant with trisomy 18, two
2016). Sternohyoid may join with omohyoid or sternothyroid accessory slips were present that originated from the
(Hallett 1848; Macalister 1867b, 1875; Wood 1867b, 1868; medial ends of the clavicles and inserted into the dis-
Kim et al. 2009a; Lee and Yang 2016; Standring 2016; Çetkin tal end of the right greater horn of the hyoid (Bersu and
et al. 2017). It may also send a slip to mylohyoid (Macalister Ramirez-Castro 1977).
1875; Lee and Yang 2016). Sternohyoid may be doubled or In a neonate with trisomy 18, the left sternohyoid orig-
absent (Macalister 1875; Lee and Yang 2016; Standring 2016). inated partially behind its counterpart and sent a slip to
The origin of sternohyoid may be confned to just the the deep cervical fascia (Aziz 1979). In addition to this
sternum (Macalister 1875; Mori 1964; Bergman et al. slip and bilateral sternohyoideus azygos muscles, this
1988). It may have an additional origin from the frst and/ individual also had two other bilateral supernumerary
or second ribs (Mori 1964; Kim et al. 2009a; Lee and Yang muscles that ascended from the manubrium to the head
2016). Macalister (1875) reports a case in which sterno- (Aziz 1979). In a boy with trisomy 13q, the left sterno-
hyoid originated from only the posterior sternoclavicular hyoid was doubled (Pettersen 1979). In a neonate with
ligament and the cartilage of the frst rib. Its clavicular ori- trisomy 13, sternohyoid and omohyoid fused near their
gin may be shifted laterally to the middle of the clavicle insertions (Aziz 1980). In an individual with craniora-
(Macalister 1875; Kim et al. 2015b). Sternohyoid may also chischisis, the sternohyoid muscles were fused with each
originate only from the middle of the clavicle (Hallett 1848; other and with the proximal portions of the omohyoid
Macalister 1875; Bergman et al. 1988; Lee and Yang 2016). muscles (Alghamdi et al. 2017).
This variant may be mistaken for cleidohyoid, an accessory
slip that extends from the clavicle to the hyoid bone (Lee
and Yang 2016; see the entry for this muscle). An accessory Prevalence
cleidohyoid may fuse with sternohyoid (Leppi 1962; Mori In their literature review, Smith et al. (2015) found that ster-
1964). Sternohyoid is also associated with sternohyoideus nohyoideus azygos was present in 1 out of 20 individuals
azygos, which extends from the manubrium to the hyoid with trisomy 13 (5%) and in 2 out of 17 individuals with
(Bergman et al. 1988; Lee and Yang 2016). trisomy 18 (11.8%). They also found that a doubled sterno-
hyoid or a sternohyoid with an accessory slip was found in
Prevalence 1 out of 20 individuals with trisomy 13 (5%) and in 4 out of
Wood (1868) states that Turner found sternohyoid blended 17 individuals with trisomy 18 (23.5%).
with the superior belly of omohyoid in 4 out of 373 cases
(1.1%). In a sample of 86 sides from 43 cadavers, Mori
(1964) found that sternohyoid originated from the sternum Clinical Implications
only on fve sides (11.6%); from both the sternum and clavi- An attachment of sternohyoid to the middle of the clavicle
cle on 64 sides (74.4%); from the sternum, clavicle, and frst may present as a lateral neck mass, a condition which Kim
rib on two sides (4.6%); and from the sternum, clavicle, frst et al. (2015b) refer to as sternohyoid syndrome.
and second ribs on two sides (4.6%). A tendinous inscrip-
tion was present on 22 out of 60 sides (36.7%).
cleiDOhyOiDeus (cleiDOhyOiD) (figure 2.10)
Sato (1968b) found at least one tendinous intersection
present in 23 out of 338 sides (6.8%) in Kyushu-Japanese See also: Sternohyoideus, Omohyoideus
Synonyms OMOhyOiDeus (OMOhyOiD) (figure 2.10)

This muscle may also be referred to as cleidohyoideus See also: Cleidohyoid
accessorius (Steinbach 1923).
Synonyms
Typical Presentation N/A
Comparative Anatomy
Description
Common chimpanzees may sometimes exhibit an omo-
Omohyoideus is comprised of two bellies joined by an inter-
hyoid with three bellies including a superior belly, infero-
mediate tendon (Standring 2016). The inferior belly originates
medial belly, and an inferolateral belly (Gratiolet and Alix
from the superior margin of the scapula near the scapular
1866; Diogo and Wood 2011, 2012a; Diogo et al. 2013a).
notch and ends in the intermediate tendon (Standring 2016).
One of the inferior bellies may correspond to cleidohyoid
The superior belly arises from the intermediate tendon and
(Gratiolet and Alix 1866; Sonntag 1923; Diogo et al. 2013a).
inserts into the body of the hyoid (Standring 2016).
Variations Innervation
Omohyoideus is innervated by the ansa cervicalis
Description
(Standring 2016).
When present, cleidohyoid originates from the middle of
the clavicle between the superior belly of omohyoid and Comparative Anatomy
the sternohyoid and inserts into the hyoid bone (Macalister Omohyoideus has a similar typical presentation in the
1875; Le Double 1897; Leppi 1962; Mori 1964; Hatipoğlu apes, extending from the scapula to the hyoid (Deniker
et al. 2006; Lee and Yang 2016). Its clavicular origin may 1885; Kohlbrügge 1890–1892; Gibbs 1999; Diogo et al.
arise from behind the cleidomastoid head of sternocleido- 2010, 2012, 2013a,b, 2017). The intermediate tendon is
mastoid (Sato et al. 1987; Bergman et al. 1988; Kim et al. poorly developed or absent in gibbons (Deniker 1885;
2009a; Stark et al. 2009; Lee and Yang 2016). Cleidohyoid Kohlbrügge 1890–1892; Diogo et al. 2012). The interme-
may fuse with either sternohyoid and/or the superior belly diate tendon is absent in most orangutans (Sonntag 1924;
of omohyoid (Leppi 1962; Mori 1964; Hatipoğlu et al. Kallner 1956; Diogo et al. 2013b). A well-developed inter-
2006). Cleidohyoid may also be used to refer to a vari- mediate tendon is present in some gorillas (Macalister
ant of sternohyoid that originates only from the middle 1873; Raven 1950; Diogo et al. 2010). Raven (1950) found
clavicle, or a variant of omohyoid with an absent inferior an omohyoid with three bellies on one side of a gorilla.
belly and a superior belly that originates from the clavi- The intermediate tendon between the superior and infe-
cle (Macalister 1875; Knott 1883a; Bergman et al. 1988; rior bellies is well-developed in common chimpanzees
Novakov et al. 2012; Lee and Yang 2016). (Gratiolet and Alix 1866; Macalister 1871; Sonntag 1923;
Diogo et al. 2013a). Differing from common chimpan-
Innervation zees, the intermediate tendon is absent in most bonobos,
Cleidohyoid is innervated by the ansa cervicalis, specif- so omohyoid is typically not divided into two bellies in
cally branches from the second and/or third cervical nerves this species (Diogo et al. 2017). In bonobos, the omohyoid
(Sato et al. 1987). may be fused with sternohyoid or the cleidomastoid head
of sternocleidomastoid (Miller 1952; Diogo et al. 2017).
Prevalence
Macalister (1875) states that Schwegl found cleidohyoid in Variations
3 out of 100 subjects (3%). Mori (1964) found cleidohyoid Description
present in 8 out of 300 sides (2.7%), fusing with sternohyoid Omohyoid may partially or entirely originate from
on three sides (1%) and with omohyoid on three sides (1%). the superior transverse scapular ligament (Macalister
Sukekawa and Itoh (2006) found cleidohyoid on 2 out of 67 1875; Mori 1964; Lee and Yang 2016; Standring 2016).
sides (3%). Omohyoid may partially or entirely originate from
the acromion process (Macalister 1875; Knott 1883a;
Anomalies Bergman et al. 1988; Lee and Yang 2016). It may also
This muscle is present only as a variation, though accessory originate entirely from the frst rib (Macalister 1875)
slips in this region occur often in individuals with trisomy or from the coracoid process (coraco-hyoid of Gruber)
18 (Bersu and Ramirez-Castro 1977; Aziz 1979). (Knott 1880, 1883a). Tubbs et al. (2004a) found an origin
of omohyoid from the scapula and an insertion into the
Clinical Implications transverse process of the sixth cervical vertebra. A partial
The presence of cleidohyoid may complicate surgical origin of omohyoid from the clavicle may take the form
approaches in the neck (Leppi 1962). of an additional head (Hallett 1848; Wood 1864, 1868;
Macalister 1875; Knott 1880; Bergman et al. 1988; Lee inserts into the middle of the clavicle (Mori 1964; Bakkum
and Yang 2016). When the additional head is present, it and Miller 1964). Cervico-costo-humeralis (Gruber) is
typically joins with the inferior belly before ending in the considered an aberrant version of omohyoid that originates
intermediate tendon (Hallett 1848; Macalister 1875). The from the lesser tubercle of the humerus and inserts into the
inferior belly may also originate entirely from the clavi- cartilage of the frst rib and transverse process of the sixth
cle (Hallett 1848; Macalister 1867b, 1875; Bergman et al. cervical vertebra (Bergman et al. 1988; Lee and Yang 2016).
1988; Standring 2016; Singh et al. 2018).
The form of the intermediate tendon is variable, and Prevalence
it may be absent (Hallett 1848; Wood 1868; Macalister Hallett (1848) found that omohyoid had an additional origin
1875; Mori 1964; Bergman et al. 1988; Lee and Yang 2016; from the clavicle in about 1 out of every 15 subjects (6.7%).
Standring 2016). The cervical fascia that surrounds the Macalister (1875) states that an accessory head from the
intermediate tendon may pull the inferior belly of the omo- clavicle was found by Wood in 5 out of 70 subjects (7.1%),
hyoid down such that it lies along the clavicle (Hallett 1848; by Turner in 17 out of 373 cases (4.6%), and by Schwegl in 2
Macalister 1875; Bergman et al. 1988). out of 100 cases (2%). Wood (1868) states that Turner found
Omohyoid may join with sternohyoid (Hallett 1848; sternohyoid blended with the superior belly of omohyoid in
Macalister 1867b, 1875; Wood 1867b, 1868; Sukekawa and 4 out of 373 cases (1.1%).
Itoh 2006; Kim et al. 2009a; Lee and Yang 2016; Standring In a sample of 94 sides, Mori (1964) found that omohyoid
2016; Çetkin et al. 2017). It may connect with mylohyoid, originated from the superior margin of the scapula only on
stylohyoid, or sternomastoid (Hallett 1848; Macalister 68 sides (72.3%), from the superior transverse ligament only
1875; Lee and Yang 2016). It may send a slip to the man- on two sides (2.1%), and from both on 24 sides (25.5%). In a
dible or the greater horn of the hyoid (Macalister 1875; Lee sample of 240 sides, Mori (1964) found that the intermedi-
and Yang 2016). It may also have a tendinous connection to ate tendon was absent on 26 sides (10.8%). The intermediate
the cartilage of the frst rib or the sternoclavicular articula- tendon was short and narrow on 12 sides (5%) and long and
tion (Hallett 1848). narrow on 76 sides (31.7%). The intermediate tendon did not
One belly or the entire muscle may be doubled (Wood extend over the entire breadth of the omohyoid and was only
1864, 1867b; Macalister 1875; Bergman et al. 1988; Rai present on the medial or lateral side on 108 sides (45%). The
et al. 2007; Kim et al. 2010; Lee and Yang 2016). In cases intermediate tendon was as wide as the muscle and moder-
of doubling, there are often attachments to sternohyoid ately long on 12 sides (5%) and wide as the muscle but short
(Macalister 1875; Rai et al. 2007; Kim et al. 2020 Lee and and wavy on four sides (1.7%). Mori (1964) also found omo-
Yang 2016). The superior belly may be split into posterior clavicularis present on 6 out of 200 sides (3%).
and anterior portions or may present as a lamellar divided In a sample of 67 sides, Sukekawa and Itoh (2006) found
structure with three to fve bellies (Sukekawa and Itoh that the superior belly of omohyoid was normal on 43 sides
2006). Wood (1868) observed a case in which the superior (64.2%), comprised of undeveloped myofbers on four sides
belly of the omohyoid was doubled. The upper belly fused (6%), divided into two bellies on seven sides (10.4%), and
with hyoglossus and the middle pharyngeal constrictor and presented as a lamellar divided belly on fve sides (7.5%).
received a slip from sternothyroid (Wood 1868). The intermediate tendon was absent on one side (1.5%). The
One belly of omohyoid or the entire muscle may be absent superior belly inserted into sternohyoid on one side (1.5%).
(Hallett 1848; Macalister 1875; Bergman et al. 1988; Zhao The superior belly connected to sternohyoid via a supernu-
et al. 2015; Lee and Yang 2016; Standring 2016). When the merary belly on two sides (3%).
superior belly is absent, the inferior belly may end in the cer-
vical fascia and is referred to as the coraco-cervicalis (Hallett Anomalies
1848; Macalister 1875; Knott 1883a; Lee and Yang 2016). Description
The superior belly may also be replaced by a tendon or unde- In a female anencephalic fetus, the left superior belly of
veloped myofbers (Knott 1880, 1883a; Sukekawa and Itoh omohyoid was replaced by a band of fbrous tissue (Windle
2006). When the inferior belly is absent the superior belly 1893). Mieden (1982) describes two male fetuses with
originates from the clavicle (Bergman et al. 1988; Lee and cyclopia and alobar holoprosencephaly. On the right side of
Yang 2016). This variant may be referred to as cleidohyoid, one specimen, fbers from the superior belly of omohyoid
an accessory slip that extends from the clavicle to the hyoid joined with fbers of geniohyoid to insert to the mandible.
bone (see the entry for this muscle). An accessory cleidohyoid In an individual with craniorachischisis, the intermediate
may fuse with omohyoid (Leppi 1962; Mori 1964; Hatipoğlu tendons were absent and the proximal portions of the omo-
et al. 2006). A similar slip, cleidothyroid, extends from the hyoids fused with the sternohyoid muscles (Alghamdi et al.
clavicle to the thyroid cartilage (Bergman et al. 1988). 2017). There was also an extra slip present deep to the right
Omohyoid is associated with several other accessory omohyoid that fused with its distal end and attached proxi-
slips. Cleidofascialis extends from the clavicle into the cer- mally to the clavicle (Alghamdi et al. 2017).
vical fascia (Macalister 1867b; Bergman et al. 1988; Lee The intermediate tendon of omohyoid was poorly
and Yang 2016). Omoclavicularis originates from the supe- developed in all of the specimens with trisomy 18 exam-
rior margin of the scapula near the origin of omohyoid and ined by Bersu and Ramirez-Castro (1977). In one of the
cases, bilateral accessory fascicles were present that usually passes anteriorly to the posterior portion of the
originated from the clavicle and joined with omohyoid main body of thyrohyoid (Gratiolet and Alix 1866; Deniker
at its midpoint. In another case, the left inferior belly of 1885; Kohlbrügge 1890–1892; Sonntag 1923; Starck and
omohyoid originated entirely from the clavicle and an Schneider 1960; Swindler and Wood 1973; Diogo et al.
extra slip from the clavicle was present on the right side. 2010, 2012, 2013a, 2017). Sternothyroid may be fused with
In a third case, the right inferior belly of omohyoideus thyrohyoid in gorillas and bonobos (Diogo et al. 2010,
received an extra slip from the transverse scapular liga- 2017). A tendinous intersection may be present in gorillas
ment and coracoid process. In the female fetus with tri- (Raven 1950) and common chimpanzees (Macalister 1871).
somy 18 dissected by Alghamdi et al. (2018), omohyoid
had an extra head on the left side that arose anterior to its
Variations
intermediate tendon and ran lateral to the normal head to
insert onto the greater horn of the hyoid. Description
Aziz (1979, 1980, 1981) found that the intermediate ten- Sternothyroid may be absent (Macalister 1875; Bale and
don of omohyoid was absent bilaterally in two neonates with Herrin 2016). One or more tendinous intersections may
trisomy 18 and one neonate with trisomy 13. In a neonate be present in the muscle (Hallett 1848; Macalister 1875;
with trisomy 13, sternohyoid and omohyoid fused near their Sato 1968b). The right and left sternothyroid muscles
insertions (Aziz 1980). In another neonate with trisomy 13, may fuse (Wood 1868; Macalister 1875; Sato 1968b; Lee
there was an extra superior belly of omohyoid on the right side and Yang 2016). The origin of sternothyroid may extend
that extended between sternocleidomastoid and sternohyoid onto the medial half of the frst rib (Kim et al. 2009a) or
(Pettersen et al. 1979). In a fetus with trisomy 13, the left supe- onto the cartilage of the second rib (Macalister 1875).
rior belly of omohyoid was hypoplastic (Pettersen et al. 1979). The costal origin may also be absent (Macalister 1875).
It may have a second head that arises from the clavicle
(Wood 1868; Macalister 1875). Kang et al. (2015) report
Prevalence
an accessory belly that originated from the left ster-
In their literature review, Smith et al. (2015) found that nothyroid and pretracheal layer of cervical fascia and
anomalies of omohyoid were present in 7 out of 24 individ- inserted into the oblique line of the thyroid cartilage on
uals with trisomy 13 (29.2%) and in 11 out of 26 individuals the right side.
with trisomy 18 (42.3%). Sternothyroid may be doubled, divided into bun-
dles, or split into two layers (Hallett 1848; Wood 1868;
Clinical Implications Macalister 1867b, 1875; Bergman et al. 1988; Lee and
A doubled omohyoid muscle may cause omohyoid syn- Yang 2016). Nayak et al. (2009a) report a case in which
drome (Guo-Hua et al. 2009) or progressive dysphagia and sternothyroid divided into a lateral belly that inserted into
dyspnea (Kshirsagar et al. 2019). the thyroid cartilage and a medial belly that became ten-
dinous and inserted into the hyoid bone and intermediate
sternOthyrOiDeus (sternOthyrOiD) (figure 2.10) tendon of the digastricus. Murugan et al. (2016) report a
case in which sternothyroid divided into a medial belly
See also: Thyrotrachealis that inserted into the thyroid cartilage and an elongated
lateral belly that sent fbers to the carotid sheath, coursed
Synonyms between the internal jugular vein and internal carotid
N/A artery, and inserted into the tympanic plate and petrous
temporal bone.
Typical Presentation Sternothyroid may be joined with sternohyoid, thyrohy-
Description oid, omohyoid, levator glandulae thyroideae, cricothyroid,
Sternothyroideus originates from the manubrium and car- hyoglossus, the middle or inferior pharyngeal constric-
tilage of the frst rib and inserts into the oblique line of the tors, or the cricoid cartilage (Wood 1868; Macalister 1875;
thyroid cartilage (Standring 2016). Bergman et al. 1988; Sakamoto 2009; Maranillo and
Sanudo 2016; Lee and Yang 2016; Çetkin et al. 2017).
Innervation Sternothyroid is associated with incisurae mediae obliquus
Sternothyroideus is innervated by branches of the ansa cer- (Gruber), an accessory muscle on the inferolateral aspect of
vicalis (Standring 2016). the thyroid cartilage (Gruber 1868a; Bergman et al. 1988;
Maranillo and Sanudo 2016). It may have a second head
Comparative Anatomy that originates from the sternothyroid, and in this case, the
Sternothyroideus has a similar typical presentation in the muscle would be referred to as incisurae mediae obliquus
apes, extending from the sternum and adjacent regions to bicaudatus (Bergman et al. 1988; Maranillo and Sanudo
the thyroid cartilage (Gratiolet and Alix 1866; Sonntag 2016). It is also associated with thyrotrachealis, which may
1923; Miller 1952; Gibbs 1999; Diogo et al. 2010, 2012, be a detached slip of sternothyroid (Macalister 1875; see the
2013a,b, 2017). The main body of sternothyroid in the apes entry for this muscle).
Prevalence Variations
In a sample of 44 sides from 22 cadavers, Sakamoto (2009) Description
found that fbers of the inferior pharyngeal constrictor con- Thyrohyoid may be continuous with sternothyroid
nected with sternothyroid on 29 sides (65.9%). Sato (1968b) (Macalister 1875; Bergman et al. 1988; Lee and Yang 2016).
found at least one tendinous intersection present in 58 out It may also connect with omohyoid, the middle or inferior
of 362 sides (16.02%) in Kyushu-Japanese males and pres- pharyngeal constrictors, hyoglossus, levator glandulae thy-
ent in 29 out of 234 sides (12.39%) in females. In males, one roideae, cricothyroid, or the cricoid cartilage (Hallett 1848;
intersection was present in 43 out of the 58 sides (74.14%), Macalister 1875; Bergman et al. 1988; Sakamoto 2009,
two intersections were present in nine sides (15.52%), three 2014; Lee and Yang 2016; Maranillo and Sanudo 2016). It
intersections were present in four sides (6.9%), and fve inter- may be divided into fascicles (Macalister 1875).
sections were present in two sides (3.45%). In females, one Thyrohyoid may connect with hyotrachealis, a slip
intersection was present in 25 out of the 29 sides (86.21%) and that originates from the hyoid, passes behind the isthmus
two intersections were present in 4 sides (13.79%). Right and of the thyroid gland, and inserts into the perichondrium
left sternothyroid muscles were fused in 13 out of 181 males of the upper rings of the trachea (Macalister 1875). An
(7.18%) and in 6 out of 117 females (5.13%) (Sato 1968b). accessory muscle referred to as thyrohyoideus superior
(minor, or azygos) may extend from the upper border of
Anomalies the thyroid cartilage to the hyoid (Macalister 1875). A
Description slip referred to as ceratohyoid or thyrohyoideus latera-
In one neonate with trisomy 13, the posterior belly of the lis may extend between the superior horn of the thyroid
digastric was doubled bilaterally, and the extra belly on cartilage to the greater horn of the hyoid (Maranillo and
the left side received a contribution from sternothyroid Sanudo 2016). Macalister (1875) suggests that variants of
(Pettersen et al. 1979). ceratohyoid include thyreo-syndesmicus (Sömmerring),
which inserts into the posterior thyrohyoid ligament,
Prevalence and a thyreo-triticeal slip, which attaches to the triticeal
In their literature review, Smith et al. (2015) found that cartilage.
anomalies of sternothyroid were found in only 1 out of
20 individuals with trisomy 13 (5%, the case described by Prevalence
Pettersen et al. 1979).
In a sample of 44 sides from 22 cadavers, Sakamoto (2009)
Clinical Implications found that fbers of the inferior constrictor connected with
thyrohyoid on 13 sides (29.5%). In a sample of 82 sides from
Variations of sternothyroid may complicate surgical proce-
41 cadavers, Sakamoto (2014) found fbers of the middle
dures in the anterior neck (Kang et al. 2015).
constrictor attached to thyrohyoid on two sides (2.4%).
thyrOhyOiDeus (thyrOhyOiD) (figure 2.10) Anomalies

Synonyms Description
This muscle may also be referred to as the hyothyroid mus- In an infant with trisomy 18, the left thyrohyoid was dou-
cle (Hallett 1848). bled (Bersu and Ramirez-Castro 1977). In a fetus with
trisomy 18, there is an accessory slip along the midline
Typical Presentation between the thyrohyoid muscles (Urban and Bersu 1987).
In a fetus with craniorachischisis, thyrohyoid fused with
Description the thyroid cartilage on the right side (Alghamdi et al.
Thyrohyoideus originates from the oblique line of the thy- 2017).
roid cartilage and inserts into the body and greater horn of
the hyoid (Standring 2016). Prevalence
Doubling of the thyrohyoid was only found in one out of
Innervation eight infants with trisomy 18 (12.5%) (Bersu and Ramirez-
Thyrohyoideus is innervated by fbers of the frst cervical Castro 1977).
spinal nerve from the hypoglossal nerve (Standring 2016).
N/A
Comparative Anatomy
Thyrohyoideus has a similar typical presentation in the
apes, extending from the thyroid cartilage to the hyoid levatOr glanDulae thyrOiDeae (figure 2.10)
bone (Gibbs 1999; Diogo et al. 2010, 2012, 2013a,b, 2017).
Thyrohyoid may be fused with sternothyroid in gorillas and Synonyms
bonobos (Diogo et al. 2010, 2017). N/A
Typical Presentation Mori (1964) found 210 cases of levator glandulae thy-
This muscle is present only as a variation or anomaly. roideae in 510 cadavers (41.2%). Mori (1964) classifed the
muscle into fve types. Hyopyramidalis, from the hyoid to
Comparative Anatomy the pyramidal lobe, was present in 53 out of the 210 cases
Kohlbrügge (1890–1892) found fbers of the inferior pha- (25.2%). Thyreopyramidalis, from the thyroid cartilage
ryngeal constrictor that attached to the thyroid gland in one to the pyramidal lobe, was present in 18 cases (8.6%).
gibbon. This structure could correspond to the levator glan- Thyreoglandularis, from the thyroid cartilage to the sheath
dulae thyroideae (Diogo et al. 2012). of the thyroid gland, was present in 115 cases (54.8%).
Hyoglandularis, from the hyoid to the sheath of the thyroid
gland, was present in 28 cases (13.3%). Tracheoglandularis,
Variations
from the upper trachea to the capsule of the isthmus, was
Description present in seven cases (3.3%).
Levator glandulae thyroideae originates from the hyoid
bone or thyroid cartilage and inserts onto the thyroid Anomalies
gland (Hallett 1848; Macalister 1875; Mori 1964; Sato
1968b; Sultana et al. 2009; Kim et al. 2010; Lee and Yang Description
2016; Standring 2016; Velasco-Nieves et al. 2020). It may In a neonate with trisomy 13, levator glandulae thyroideae
insert into the lobe, pyramidal lobe, isthmus, or sheath extended from the thyroid cartilage to the isthmus of the
of the gland (Hallett 1848; Macalister 1875; Keyes 1940; thyroid gland (Pettersen et al. 1979). In another neonate
Mori 1964; Sato 1968b; Bergman et al. 1988; Gregory and with trisomy 13, the muscle extended from the hyoid bone
Guse 2007; Sultana et al. 2009; Kim et al. 2010; Murugan to the isthmus of the thyroid gland (Aziz 1980).
et al. 2016; Standring 2016). Chaudhary et al. (2013) clas-
sify levator glandulae thyroideae as a fbromusculoglan- Prevalence
dular band. In their literature review, Smith et al. (2015) found that leva-
Levator glandulae thyroideae may be doubled or bifur- tor glandulae thyroideae was present in 2 out of 20 indi-
cated at its origin or insertion (Hallett 1848; Macalister viduals with trisomy 13 (10%).
1875). Pacífco et al. (2019) report a case in which leva-
tor glandulae thyroideae had a short bundle that inserted Clinical Implications
into the cricoid cartilage and a long bundle that inserted Levator glandulae thyroideae may be mistaken for a tumor
into the pyramidal lobe. It may also consist of three slips or infected thyroglossal duct cyst in medical imaging
(Macalister 1875; Loukas et al. 2008a). It may be con- (Loukas et al. 2008a).
nected with sternothyroid or thyrohyoid (Hallett 1848;
Macalister 1875). This muscle may present as fbers
extending from the inferior pharyngeal constrictor to the SUPRAHYOID MUSCLES
thyroid gland, in which case it may be referred to as leva-
MylOhyOiDeus (MylOhyOiD) (figure 2.10)
tor glandulae thyroideae lateralis (Krause) (Knott 1883a;
Bergman et al. 1988). Synonyms
N/A
Innervation
Levator glandulae thyroideae may be innervated by a Typical Presentation
branch from the ansa cervicalis (Loukas et al. 2008a), Description
or more specifically, a branch from the second cervical
Mylohyoid originates from the mylohyoid line of the man-
nerve (Sato et al. 1987). It may also be innervated by the
dible (Standring 2016). Its posterior fbers insert onto the
external laryngeal nerve (Keyes 1940; Chaudhary et al.
body of the hyoid bone (Standring 2016). The anterior and
2013).
middle fbers of mylohyoid on each side insert onto a fbrous
median raphe (mylohyoid raphe) (Standring 2016). The
Prevalence mylohyoid raphe extends from the mental symphysis of the
Sato (1968b) found levator glandulae thyroideae present mandible to the hyoid (Standring 2016).
in 90 out of 350 sides in males (25.7%) and present in 53
out of 220 sides (24.1%) in females. Lehr (1979) reported Innervation
one case of this muscle out of 203 cases (0.49%). Sultana Mylohyoid is innervated by the nerve to mylohyoid, a
et al. (2009) found this muscle in 26 out of 60 cadav- branch of the inferior alveolar nerve (Standring 2016).
ers (43.3%). Yadav et al. (2014) found this muscle in 9
out of 26 cadavers (34.6%). In a sample of 52 fetuses, Comparative Anatomy
Chaudhary et al. (2016) found levator glandulae thyroi- Mylohyoideus has a similar typical presentation in the
deae in 10 fetuses (19.2%). apes, extending from the mylohyoid lines of the mandible
to the hyoid bone and the ventral midline (Gibbs 1999; the left mylohyoid line and inserted into the median raphe
Diogo et al. 2010, 2012, 2013a,b, 2017). The median raphe and hyoid bone. Zdilla et al. (2018) provide a comprehen-
is absent in orangutans (Sonntag 1924; Brown and Ward sive review of the “arrowhead variation,” bilateral acces-
1988) and gorillas (Saban 1968; Göllner 1982; Gibbs 1999; sory muscles that extend between the intermediate tendons
Diogo et al. 2010). The median raphe has been observed in of the digastric and the median raphe of the mylohyoid
rare cases in gibbons (DuBrul 1958) and common chim- musculature (see the entry for digastricus anterior for more
panzees (Göllner 1982). The median raphe is typically pres- information).
ent in bonobos (Gibbs 1999; Miller 1952; Diogo et al. 2017).
Prevalence
Variations Hallett (1848) found that mylohyoideus was connected to
Description the intermediate tendon of the digastricus in one out of
five subjects (20%). Mori (1964) found that the median
Mylohyoid is absent in rare cases (Hallett 1848; Macalister
raphe was only clearly present in 45 out of 210 cases
1875; Bergman et al. 1988; Lee and Yang 2016). Hallett
(21.4%). In a sample of 19 cadavers, Zdilla et al. (2018)
(1848) and Macalister (1875) observed cases in which mylo-
found that the arrowhead variation was present in two
hyoid was completely absent and replaced by an enlarged
cadavers (10.5%).
anterior belly of the digastric muscle. If the median raphe is
In a sample of 324 half-heads, Gaughran (1963) found
absent, the two mylohyoid muscles may fuse into a continu-
that a process of the sublingual gland herniated through a
ous muscle sheet (Zdilla and Lambert 2015; Standring 2016).
hiatus in mylohyoideus in 102 cases (31.5%) and fat her-
Mylohyoid may fuse with, send fbers to, or insert onto
niations were present in 15 cases (4.6%). In a sample of
the anterior belly of the digastric muscle (Macalister 1875;
300 half-heads from 150 cadavers, Nathan and Luchansky
Malpas 1926; Bergman et al. 1988; Saadeh et al. 2001; Lee
(1985) found that a part of the sublingual gland and/or fat
and Yang 2016; Standring 2016). It may also join with the
tissue herniated through a hiatus in mylohyoideus in 82
intermediate tendon of the digastricus (Hallett 1848). The
specimens (27.3%) from 63 cadavers.
anterior belly of the digastric may originate from mylo-
hyoid (Lee and Yang 2016). Accessory anterior bellies of
the digastric may have attachments to mylohyoid (Sevinç Anomalies
et al. 2009). Mylohyoid may receive a slip from omohyoid, Description
sternohyoid, or stylohyoid (Macalister 1875; Saadeh et al. Bersu et al. (1976) describe a male infant with Hanhart syn-
2001). The stylohyoid may insert into the outer border of drome in which the right mandible was represented by a
mylohyoid (Macalister 1875). Zdilla and Lambert (2015) small bone. On the right side, mylohyoid had a restricted
report a case in which the mylohyoid muscles did not reach origin from the distal end of the mandible and inserted along
the hyoid bone but inserted into the middle of the genio- the median raphe. An accessory muscle that was innervated
hyoid musculature via a fbrous “pseudo-hyoid” structure. by the mylohyoid nerve originated from the angle of the
The length of the insertion along the hyoid bone may right mandible and extended across the foor of the mouth
vary (Macalister 1875; Malpas 1926). The sublingual gland to insert onto the middle half of the inferior aspect of the
and/or fat tissue may partially or entirely herniate through left mandible. A slip from this accessory muscle extended
a hiatus in mylohyoideus (Malpas 1926; Gaughran 1963; between the geniohyoid muscles to have a diffuse insertion
Nathan and Luchansky 1985; Lee and Yang 2016; Standring onto the deep aspect of the left geniohyoid and the hyoglos-
2016; Bender-Heine and Zdilla 2018). Lobes of the subman- sus muscles.
dibular gland may also herniate through mylohyoid and Colacino and Pettersen (1978), Pettersen et al. (1979),
split it into discrete bundles (Macalister 1875; Knott 1883a; and Aziz (1980) report anomalies of mylohyoid in neonates
Bergman et al. 1988; Lee and Yang 2016). Mylohyoid with trisomy 13. In one individual, the mylohyoid muscles
may also be divided into parts by the submandibular vein were defcient anteriorly (Colacino and Pettersen 1978). In
(Saadeh et al. 2001). It can also be divided into superfcial another, the left digastric muscle sent a slip to the left mylo-
and deeper layers (Malpas 1926). hyoid muscle, which was fused with its right counterpart
Bender-Heine and Zdilla (2018) report a case in which since the median raphe was absent (Pettersen et al. 1979).
anterior and posterior bellies of mylohyoid were completely The mylohyoid raphe was also absent in another neonate
separated by fat that herniated from the sublingual space. (Aziz 1980).
The anterior bellies inserted into a single-bellied geniohy- Mylohyoid is also variable in some individuals with
oid, and the posterior bellies inserted into the intermediate trisomy 18 (Bersu and Ramirez-Castro 1977; Aziz 1979;
tendons and anterior bellies of the digastric. Smith et al. 2015). In one infant, there was an accessory
Accessory muscles associated with mylohyoid may be muscle sheet between the bellies of anterior digastric and
present (Sehirli and Çavdar 1996; Lee and Yang 2016; the mylohyoid musculature (Bersu and Ramirez-Castro
Zdilla et al. 2018). Sehirli and Çavdar (1996) report an 1977). All specimens examined by Bersu and Ramirez-
accessory mylohyoid situated between the left anterior belly Castro (1977) lacked a median raphe and the left and
of the digastric and the left mylohyoid. It originated from right mylohyoid muscles were therefore fused in all cases.
The neonate described by Aziz (1979) also lacked a median Innervation

raphe. In the fetus with trisomy 18 and cyclopia described Geniohyoid is innervated by a branch of the frst cervi-
by Smith et al. (2015), mylohyoid exchanged fbers with cal spinal nerve that travels with the hypoglossal nerve
the digastric muscle bilaterally. The mylohyoid raphe was (Standring 2016).
absent, so the mylohyoid muscles presented as a continu-
ous muscle sheet. Intermandibularis anterior, a narrow Comparative Anatomy
accessory muscle deep to mylohyoideus with transverse Geniohyoid has a similar typical presentation in the apes,
fbers that spanned the two halves of the anterior mandible, extending from the inner surface of the mandible to the
was present. hyoid bone (Gratiolet and Alix 1866; Sonntag 1923; Raven
Mieden (1982) describes an infant with median cleft lip, 1950; Miller 1952; Gibbs 1999; Diogo et al. 2010, 2012,
hypotelorism, and alobar holoprosencephaly (case I) and 2013a,b, 2017). In gibbons and gorillas, the insertion into
two male fetuses with cyclopia and alobar holoprosenceph- the hyoid bone bifurcates the posterior portion of hyoglos-
aly (cases II and III). In cases I and III, mylohyoideus was sus (Diogo et al. 2010, 2012). In common chimpanzees and
absent bilaterally (Mieden 1982). In a male neonate with bonobos, the muscle often merges with its counterpart at
Meckel syndrome, Pettersen (1984) found that the anterior the midline (Sonntag 1923; Diogo et al. 2013a, 2017).
belly of the right digastricus sent an accessory slip to the
mylohyoideus. In an infant with mandibulofacial dysostosis, Variations
mylohyoid was deep but short anteroposteriorly (Herring
et al. 1979). Description
In the fetus with craniorachischisis described by Geniohyoid may fuse with its counterpart across the mid-
Alghamdi et al. (2017), the right mylohyoid was diminu- line and present as a continuous muscle sheet (Hallett 1848;
tive and fused with the digastric. The fascia over the left Macalister 1875; Mori 1964; Zdilla and Lambert 2015; Lee
mylohyoid received a tendinous slip from the intermediate and Yang 2016; Standring 2016; Bender-Heine and Zdilla
tendon of the digastric. The two mylohyoid muscles were 2018). Zdilla and Lambert (2015) report a case in which
fused together. the mylohyoid muscles did not reach the hyoid bone but
inserted into the middle of the geniohyoid musculature
Prevalence via a fbrous “pseudo-hyoid” structure. Bender-Heine and
The median raphe was absent in all nine individuals (100%) Zdilla (2018) report a case in which anterior and posterior
with trisomy 18 described by Bersu and Ramirez-Castro bellies of mylohyoid were completely separated by fat that
(1977) and Aziz (1979). In their literature review, Smith herniated from the sublingual space. The anterior bellies
et al. (2015) found that anomalous mylohyoid muscles were inserted into a single-bellied geniohyoid, and the posterior
present in 2 out of 20 individuals (10%) with trisomy 13. bellies inserted into the intermediate tendons and anterior
bellies of the digastric.
Clinical Implications Geniohyoid may also fuse with genioglossus or origi-
Surgeons and clinicians should be aware of the poten- nate from hyoglossus (Macalister 1875; Knott 1883a;
tial herniation of the sublingual or submandibular glands Bergman et al. 1988; Lee and Yang 2016; Patel and
through mylohyoideus, as well as other variations of this Loukas 2016; Standring 2016). It may receive an acces-
muscle (Gaughran 1963; Nathan and Luchansky 1985; sory slip from the greater horn of the hyoid (Macalister
Bender-Heine and Zdilla 2018). Herniation of glandu- 1875). In a case where the mylohyoid was defcient,
lar tissue through mylohyoid (Bender-Heine and Zdilla Macalister (1875) observed mentohyoid inserting into
2018) or the presence of arrowhead musculature (Zdilla geniohyoideus. Geniohyoid may be doubled bilaterally,
et al. 2018) may affect the spread of infection through thus presenting with four geniohyoid muscles (Hallett
the neck. 1848; Macalister 1875; Bergman et al. 1988). Geniohyoid
may also be split into superfcial and deep layers (Mori
1964; Mehta et al. 2011b).
geniOhyOiDeus (geniOhyOiD) (figure 2.10)
Prevalence
Synonyms N/A
N/A
Typical Presentation Anomalies

Geniohyoid originates from the inferior mental spine on the Bersu et al. (1976) describe a male infant with Hanhart syn-
posterior surface of the mental symphysis (Standring 2016). drome in which the right mandible was represented by a
It inserts onto the body of the hyoid (Standring 2016). It is small bone. In this infant, a small portion of the right genio-
situated above (deep to) the medial portion of mylohyoid hyoid originated in association with the left geniohyoid on
(Standring 2016). the left mandible and attached to the larger, anomalous part
of the muscle that originated from hyoglossus. An accessory Diogo et al. 2010, 2012, 2013a,b, 2017). Digastricus anterior
muscle was present that sent a slip between the geniohyoid is typically absent in orangutans (Chapman 1880; Sonntag
muscles to have a diffuse insertion onto the deep aspect of 1924; Cachel 1984; Wall et al. 1994; Gibbs 1999; Diogo
the left geniohyoid and the hyoglossus muscles. et al. 2013b), having only been reported in one specimen
Mieden (1982) describes an infant with median cleft lip, by Parsons (1898a). The two digastricus anterior muscles
hypotelorism, alobar holoprosencephaly (case I) and two may make contact at the midline in some gibbons (Wall
male fetuses with cyclopia and alobar holoprosencephaly et al. 1994; Gibbs et al. 2002) and gorillas (Bischoff 1880;
(cases II and III). In case I, geniohyoideus was doubled bilat- Hosokawa and Kamiya 1961–1962; Diogo et al. 2010). The
erally. In case III, both geniohyoideus muscles were divided anterior belly on each side typically contacts its counter-
into superfcial and deep heads while the right muscle had part in common chimpanzees and bonobos (Wilder 1862;
a third head that was laterally placed (Mieden 1982). In an Gratiolet and Alix 1866; Sonntag 1923; Miller 1952;
infant with mandibulofacial dysostosis, geniohyoid was DuBrul 1958; Starck and Schneider 1960; Göllner K. 1982;
reduced (Herring et al. 1979). On the left side of a fetus with Diogo et al. 2013a, 2017).
craniorachischisis, the left geniohyoid was a broad muscle
that extended more laterally than is typical (Alghamdi et al.
Variations
2017). It originated from the hyoid region and fused with
genioglossus. The right geniohyoid was absent. Description
The anterior and posterior bellies may fail to connect
Prevalence (Bergman et al. 1988). The digastric muscle may have a
tendinous inscription (Bergman et al. 1988; Lee and Yang
In their literature review, Smith et al. (2015) found that
2016). The muscle may present as trigastric, often with the
geniohyoideus was doubled bilaterally in 1 out of 17 indi-
third head extending between the intermediate tendon and
viduals with trisomy 18 (5.9%).
the mandible or midline of the neck (Hallett 1848; Bergman
Clinical Implications et al. 1988; Lee and Yang 2016). The muscle may also be
Understanding variation in the geniohyoid muscle is impor- quadrigastric (Wood 1868; Bergman et al. 1988; Lee and
tant for planning and performing operational procedures in Yang 2016) or even have more than four bellies due to vari-
the suprahyoid region (Mehta et al. 2011b). ations in the anterior belly (see below).
Digastricus anterior may cross the midline onto the
other side of the body (Macalister 1875; Standring 2016).
Digastricus anteriOr (anteriOr belly The anterior bellies on each side of the body may also
be connected by feshy fbers (Macalister 1875; Bergman
Of theDigastric) (figure 2.10)
et al. 1988; Lee and Yang 2016) or via an aponeurotic slip
See also: Digastricus posterior (Venugopal and Mallula 2010). The anterior belly of the
digastric may be enlarged as a variation (Çelik et al. 1992;
Synonyms Holibková and Machálek 1999). It may also be absent in
This muscle is often referred to as the anterior belly of the rare cases (Sato 1968b; Larsson and Lufkin 1987; Bergman
digastric (Standring 2016). et al. 1988; Sargon et al. 1999; De-Ary-Pires et al. 2003).
The anterior belly of the digastric may fuse with, send
Typical Presentation fbers to, or originate from mylohyoid (Macalister 1875;
Description Malpas 1926; Bergman et al. 1988; Saadeh et al. 2001; Lee
and Yang 2016; Standring 2016). Mylohyoid may join with
Digastricus anterior originates from the digastric fossa, a
the intermediate tendon (Hallett 1848). Hallett (1848) and
fossa on the inner surface of the inferior margin of the man-
Macalister (1875) observed cases in which mylohyoid was
dible near the midline (Standring 2016). It ends in a tendon
completely absent and replaced by an enlarged digastri-
(the intermediate tendon) that connects it with digastricus
cus anterior. The anterior belly may send a slip onto the
posterior (Standring 2016). The intermediate tendon perfo-
hyoid bone (Macalister 1875) or insert entirely onto this
rates stylohyoid (Standring 2016).
bone when the intermediate tendon is absent (Bergman
et al. 1988; Lee and Yang 2016). Stylohyoid may insert
Innervation onto the intermediate tendon (Macalister 1875; Bergman
Digastricus anterior is innervated by the nerve to mylohy- et al. 1988). The intermediate tendon may pass in front of
oid, a branch of the inferior alveolar nerve (Standring 2016). or behind stylohyoid instead of perforating it (Macalister
1875; Mori 1964; De-Ary-Pires et al. 2003; Harvey et al.
Comparative Anatomy 2015). The intermediate tendon may send slips to the lesser
Digastricus anterior has a similar typical presentation in horn of the hyoid (Macalister 1875).
most apes, extending from the mandible to the intermediate Accessory muscles are frequently associated with the
tendon with occasional attachments to the hyoid (Gratiolet anterior belly of the digastric (Macalister 1875; Bergman
and Alix 1866; Raven 1950; Miller 1952; Gibbs 1999; et al. 1988; Lee and Yang 2016). Mentohyoid refers to a
supernumerary muscle situated along the medial margin presentation of the anterior belly that is comprised of either
of digastricus anterior that extends between the hyoid and the coexistence of both the anterior and posterior types or a
the mandibular symphysis (Macalister 1867b, 1875; Knott more intricate form that demonstrates some combination of
1883a; Bergman et al. 1988; De-Ary-Pires et al. 2003; Lee the six aforementioned types.
and Yang 2016). It may present as a single slip or as two De-Ary-Pires et al. (2003) classified the anterior belly
parallel bands (Macalister 1875). of the digastric into five types, the intermediate tendon into
The anterior belly of the digastric may be doubled and three types, and the posterior belly in two types (see the
may sometimes cross the midline to decussate with its com- entry for digastricus posterior for more details). A type I
panion and/or find its insertion (Wood 1864, 1867b, 1868; anterior belly consists of one belly that originates near the
Macalister 1875; Knott 1883a; Bergman et al. 1988; Sargon mandibular symphysis. A type II anterior belly is comprised
et al. 1999; Aktekin et al. 2003; Turan-Özdemir et al. 2004; of two bellies with extra slips to the mandible or mylohyoid
Liquidato et al. 2007; Lee and Yang 2016; Khona et al. muscle on either side of the body. A type III anterior belly is
2017; Hsiao and Chang 2019). Unilateral accessory slips comprised of three bellies with extra slips to the mandible
of digastricus anterior may also cross the midline (Larsson or mylohyoid muscle on either side of the body. A type IV
and Lufkin 1987; Zdilla et al. 2014a; Ortug et al. 2020). anterior belly is comprised of four bellies with extra slips
Çelik et al. (1993) report a case in which the right ante- to the mandible or mylohyoid muscle on either side of the
rior belly is tripled, with all three bellies inserting into the body. A type V anterior belly is a normal anterior belly that
intermediate tendon to join with the right posterior belly. is present with the mentohyoid muscle. The intermediate
Quadrification of the anterior belly has been found uni- tendon was categorized as piercing the stylohyoid muscle
laterally with insertion of all bellies into the intermediate (type I), passing superficial to the stylohyoid (type II), or
tendon (Çelik et al. 2002), and bilaterally with insertion passing deep to the stylohyoid (type III).
of all bellies into a common tendon that attached to the Multiple accessory muscles in the submental triangle
hyoid (Ozgur et al. 2007). In the latter case, two addi- are common (Wood 1868; Sargon et al. 1999; Peker et al.
tional accessory muscles were present on each side near 2000; Aktekin et al. 2003; Fujimura et al. 2003; Liquidato
the midline and also inserted into the hyoid, giving each et al. 2007; Kyung et al. 2011; Yamazaki et al. 2011; Raju
digastricus anterior the appearance of six heads (Ozgur et al. 2014; Harvey et al. 2015; Lee and Yang 2016; Hsiao
et al. 2007). and Chang 2019). These accessory muscles may have attach-
Mori (1964) classified the anterior belly of the digastric ments to the main anterior belly, intermediate tendon of the
into nine forms. The normal form refers to an anterior belly digastric, mylohyoid, the mylohyoid raphe, body or digastric
that arises from the digastric fossa and becomes continu- fossa of the mandible, and/or the hyoid (Traini 1983; Michna
ous with the intermediate tendon. The ape form refers to 1989; Sargon and Çelik 1994; Holibková and Machálek 1999;
an anterior belly that extends between the mandible and the Sargon et al. 1999; Guelfguat et al. 2001; Peker et al. 2000;
intermediate tendon, but also has a connection to the apo- Aktekin et al. 2003; Fujimura et al. 2003; Turan-Özdemir et al.
neurosis intertendines (aponeurosis interdigastricque). The 2004; Liquidato et al. 2007; Sevinç et al. 2009; Kyung et al.
anterior type refers to an anterior belly comprised of fibers 2011; Yamazaki et al. 2011; Harvey et al. 2015; Raju
arising from the aponeurosis intertendines that fuse with et al. 2014; Lee and Yang 2016; Khona et al. 2017; Zdilla et al.
the fibers arising from the intermediate tendon to form one 2018; Hsiao and Chang 2019; Ortug et al. 2020).
broad plate. This type is further divided into the continu- Harvey et al. (2015) report a case in which digastricus
ous type, which demonstrates complete fusion between the anterior on both sides of a cadaver had superficial and deep
two parts, and the discontinuous type, in which there is a bellies. In this case, four additional accessory muscle bel-
boundary between the fibers arising from the aponeurosis lies extended across the submental region in a “weave pat-
intertendines and the normal anterior belly of the digas- tern.” Zdilla et al. (2014b) report the presence of accessory
tric. The posterior type refers to when the anterior belly is anterior digastric muscles in the submental region that pres-
associated with but distinctly separated from a thin muscle ent in a “fractal” pattern, where two accessory bellies gave
plate that originates from the aponeurosis intertendines and origin to two smaller accessory bellies.
inserts on the mandible. This type is further divided into Zdilla et al. (2018) provide a comprehensive review of a
the continuous form, in which the thin muscle plate passes common presentation of the accessory anterior bellies des-
anteriorly to the inner surface of the mandible and sends ignated as “arrowhead variations.” The arrowhead variation
fibers to mylohyoideus, and the myloid form, in which the refers to bilateral accessory muscles that extend between
thin muscle plate passes medially to fuse with the mylohy- the intermediate tendons of the digastric and the median
oideus of the opposite side. The biceps form refers to a two- raphe of the mylohyoid musculature (Aktekin et al. 2003;
headed anterior belly with a medial head that inserts into Turan-Özdemir et al. 2004; Zdilla et al. 2018). These acces-
the digastric fossa of the opposite side. The sixth form is sory arrowhead muscles often send one or two additional
not named but refers to the presence of an accessory bundle accessory bundles to the mandible (Traini 1983; Aktekin
between the mandible and the lateral margin of the ante- et al. 2003; Turan-Özdemir et al. 2004; Zdilla et al. 2018;
rior belly. The “combinated” form refers to a complicated Hsiao and Chang 2019).
Prevalence in another cadaver. Therefore, 146 digastric muscles were

studied. These authors classified the anterior belly of the
Hallett (1848) found that mylohyoideus was connected to digastric into five types, the intermediate tendon into three
the intermediate tendon of the digastricus in one out of five types, and the posterior belly in two types (see above for
subjects (20%). Hallett (1848) also found a third belly of descriptions of the first two structures and the entry for
the digastric muscle in about 1 in 15 subjects (6.7%). Wood digastricus posterior for description of the latter). They
(1868) found that the anterior belly of the digastric was then identified ten patterns exhibited by the 146 muscles.
doubled in 5 out of 68 males (7.4%) and 1 out of 34 females Pattern A—a type I anterior belly, type I intermediate ten-
(2.9%). Sato (1968b) found that the digastricus muscle was don, and type I posterior belly—was present in 96 muscles
completely absent in 7 out of 324 sides (2.16%) in Kyushu- (65.8%). Pattern B—a type II anterior belly, type I inter-
Japanese males and in 2 out of 232 sides (0.86%) in females. mediate tendon, and type I posterior belly—was present in
Mori (1964) classified the anterior belly of the digastric eight muscles (5.5%). Pattern C—a type III anterior belly,
into nine forms (see above for descriptions). Out of a sample type I intermediate tendon, and type I posterior belly—was
of 262 cadavers: the normal form was found in 121 cadavers present in one muscle (0.7%). Pattern D—a type IV ante-
(46.2%), the ape form was found in five cadavers (1.9%), the rior belly, type I intermediate tendon, and type I posterior
continuous form of the anterior type was found bilaterally in belly—was present in one muscle (0.7%). Pattern E—a type
27 cadavers (10.3%) and unilaterally in 18 cadavers (6.9%), the V anterior belly, type I intermediate tendon, and type I pos-
discontinuous form of the anterior type was found bilaterally terior belly—was present in two muscles (1.4%). Pattern
in one cadaver (0.4%) and unilaterally in five cadavers (1.9%), F—a type I anterior belly, type II intermediate tendon, and
the continuous form of the posterior type was found in four type I posterior belly—was present in 14 muscles (9.6%).
cadavers (1.5%), the myloid form of the posterior type was Pattern G—a type I anterior belly, type II intermediate ten-
found in bilaterally in 12 cadavers (4.6%) and unilaterally in don, and type II posterior belly—was present in six muscles
30 cadavers (11.5%), the biceps form was found in eight cadav- (4.1%). Pattern H—a type I anterior belly, type III inter-
ers (3.1%), an accessory bundle between the mandible and the mediate tendon, and type I posterior belly—was present in
anterior belly was found bilaterally in one cadaver (0.4%) and 12 muscles (8.2%). Pattern I—a type II anterior belly, type
unilaterally in three cadavers (1.1%), the “combinated” form II intermediate tendon, and type II posterior belly—was
was found in a total of 29 cadavers (11.1%), with eight cadav- present in four muscles (2.7%). Pattern J—a type II anterior
ers (3.1%) combining the anterior and posterior types and 21 belly, type III intermediate tendon, and type II posterior
cadavers (8%) having a more intricate form. belly—was present in two muscles (1.4%).
Larsson and Lufkin (1987) examined the anterior belly Ortug et al. (2020) applied the classifications used by
of the digastric in 40 patients that underwent CT imaging of De-Ary-Pires et al. (2003) to examine the digastricus ante-
the oropharynx and 35 patients that underwent MR imag- rior muscles on 80 sides of 40 cadavers. A type I anterior
ing of the oropharynx. In the CT group, unilateral absence belly was found in 68 sides (85%), a type II anterior belly
of the anterior belly was observed in one patient (2.5%). In was found on four sides (5%), a type III anterior belly was
this case, the anterior belly was replaced by a small muscle found on three sides (3.8%), and a type IV belly was found
extending between the mylohyoid raphe and the hyoid. In on two sides (2.5%). A unilateral accessory belly was found
the MR group, an accessory muscle crossed the midline in in one cadaver (2.5%), and bilateral accessory bellies were
one patient (2.9%) with otherwise normal digastric muscles. found in another cadaver (2.5%).
Sargon et al. (1999) found variations of the anterior
digastric muscle in 5 out of 99 cadavers (5.1%). Accessory
bellies of the anterior digastric were found in four cadavers. Anomalies
The absence of the anterior belly on one side and an atypi- Description
cal origin of the muscle on the other side were observed Bersu et al. (1976) describe a male infant with Hanhart syn-
in the fifth cadaver. In a sample of 54 cadavers, Fujimura drome in which the right mandible was represented by a
et al. (2003) found variations of the anterior belly of the small bone. On the right side, digastricus posterior had a nor-
digastric in 13 cases (24.1%). In a sample of ten cadavers, mal origin and traveled laterally to hyoglossus and inserted
Liquidato et al. (2007) found variations of digastricus ante- by its intermediate tendon onto the body of the hyoid bone at
rior in four cases (40%). In a sample of 30 cadavers, Khona the base of the greater cornu. The anterior digastricus origi-
et al. (2017) found accessory bellies of the anterior digastric nated from this bony attachment and traveled anterolaterally
in three cadavers (10%). In a sample of 19 cadavers, Zdilla to attach along the inferior margin of the right mandible.
et al. (2018) found that the arrowhead variation was present Colacino and Pettersen (1978), Pettersen (1979),
in two cadavers (10.5%). In a sample of 15 cadavers, Hsiao Pettersen et al. (1979), and Aziz (1980) report anomalies
and Chang (2019) found accessory anterior bellies of the of the anterior belly of the digastric in individuals with tri-
digastric in three cases (20%). somy 13. In one neonate, the anterior belly of the digastric
De-Ary-Pires et al. (2003) studied the digastric muscle was doubled bilaterally (Colacino and Pettersen 1978). In
in 74 cadavers. An anterior belly was absent unilaterally in another neonate, the left anterior belly had an extra bun-
one cadaver and a posterior belly was absent unilaterally dle that fused with mylohyoid and decussated across the
midline (Pettersen et al. 1979). In a fetus, the anterior bel- Prevalence

lies of the digastric were doubled bilaterally (Pettersen et al. In their literature review, Smith et al. (2015) found that
1979). The medial anterior belly on the left side originated digastricus anterior was variable in 5 out of 20 individu-
from the hyoid bone. All anterior bellies inserted into the als with trisomy 13 (25%) and 5 out of 17 individuals with
mandible and had no connections to mylohyoid. In a boy trisomy 18 (29.4%). Mentohyoideus was reported in 4 out of
with trisomy 13q, there was an extra belly of the anterior 17 individuals with trisomy 18 (23.5%). The intermediate
digastric on the left side (Pettersen 1979). In one neonate, tendon was absent in all eight infants (100%) with trisomy
the left anterior belly was doubled while the right anterior 18 described by Bersu and Ramirez-Castro (1977).
belly had three bellies that inserted onto the mandible and
a fourth belly that inserted onto the masseter (Aziz 1980). Clinical Implications
Digastricus anterior is also variable in some individu- Accessory anterior bellies of the digastric may be mistaken
als with trisomy 18 (Bersu and Ramirez-Castro 1977; Aziz for masses, tumors, or metastatic lymph nodes (Aktekin
1979; Smith et al. 2015). On the right side of one infant, et al. 2003; Guelfguat et al 2001; Turan-Özdemir et al. 2004;
the anterior belly was narrow, and an accessory belly Ozgur et al. 2007; Bonala et al. 2015; Hsiao and Chang 2019).
extended between the left anterior digastric and the digas-
tric fossa (Bersu and Ramirez-Castro 1977). In another
infant, there was an accessory muscle sheet between the Digastricus posterior (Posterior Belly
bellies of anterior digastric and the mylohyoid musculature,
of the Digastric) (Figure 2.10)
and the anterior belly was doubled on the right side. In two
infants, mentohyoideus was present bilaterally. The inter- See also: Digastricus anterior
mediate tendon was absent bilaterally in all cases examined
by Bersu and Ramirez-Castro (1977), and all anterior bel- Synonyms
lies of the digastric had an insertion into the hyoid. In the This muscle is referred to as the posterior belly of the digas-
neonate described by Aziz (1979), both the left and right tric (Standring 2016).
anterior bellies of digastric were enlarged. In the fetus with
trisomy 18 and cyclopia described by Smith et al. (2015), Typical Presentation
digastricus anterior had two heads bilaterally. Its medial
head decussated with its counterpart across the midline and Description
the lateral head fused with mylohyoid. In a fetus with tri- Digastricus posterior originates from the mastoid notch of
somy 21, there was an accessory anterior digastric belly on the temporal bone (Standring 2016). It courses inferiorly and
the left side (Bersu 1980). anteriorly to end in a tendon (the intermediate tendon) that
Mieden (1982) describes an infant with median cleft connects it with digastricus anterior (Standring 2016). The
lip, hypotelorism, and alobar holoprosencephaly. This intermediate tendon perforates stylohyoid (Standring 2016).
infant had a bilateral accessory slip extending between
the intermediate tendon of digastricus and the superior Innervation
pharyngeal constrictor (Mieden 1982). This author also Digastricus posterior is innervated by the facial nerve
describes two male fetuses with cyclopia and alobar holo- (Standring 2016).
prosencephaly. In one of these cases, the right anterior
belly of digastricus was absent, and the left muscle was Comparative Anatomy
doubled in size and attached to both sides of the mandibu- Digastricus posterior has a similar typical presentation in
lar symphysis. most apes, extending from the mastoid region of the tem-
Moen et al. (1984) found that the anterior bellies of digas- poral bone to the intermediate tendon (Raven 1950; Miller
tricus were doubled on both sides of a male infant with trip- 1952; Gibbs 1999; Diogo et al. 2010, 2012, 2013a,b, 2017).
loidy, and there were extra slips associated with the anterior Digastricus posterior inserts into the angle of the mandible
digastric bellies bilaterally in a female fetus with triploidy. in orangutans since digastricus anterior is typically absent
In a male neonate with Meckel syndrome, Pettersen (1984) (Chapman 1880; Sonntag 1924; Cachel 1984; Wall et al.
found that the anterior belly of the right digastricus sent an 1994; Gibbs 1999; Diogo et al. 2013b). The origin of the
accessory slip to the mylohyoideus. In an infant with man- posterior belly of digastricus may extend onto the occipital
dibulofacial dysostosis, the anterior belly was fasciculated bone in orangutans (Sonntag 1924), gorillas (Raven 1950;
and these fascicles originated from the body of the hyoid Diogo et al. 2010), and common chimpanzees (Tyson 1699).
(Herring et al. 1979).
In the fetus with craniorachischisis described by
Variations
Alghamdi et al. (2017), the right anterior belly of the digas-
tric was fused with mylohyoid. On the left side, the anterior Description
belly was attached to the inferior border of the mandible. The anterior and posterior bellies may fail to connect
The intermediate tendon of the digastric sent a tendinous (Bergman et al. 1988). The digastric muscle may have a
slip to the fascia over the left mylohyoid. tendinous inscription (Bergman et al. 1988; Lee and Yang
2016). The muscle may present as trigastric, often with small bone. On the right side, digastricus posterior had a nor-
the third head extending between the intermediate tendon mal origin and traveled laterally to hyoglossus and inserted
and the mandible or midline of the neck (Macalister 1875; by its intermediate tendon onto the body of the hyoid bone at
Hallett 1848; Bergman et al. 1988; Lee and Yang 2016). The the base of the greater cornu. The anterior digastricus origi-
muscle may also be quadrigastric (Wood 1868; Bergman nated from this bony attachment and traveled anterolaterally
et al. 1988; Lee and Yang 2016) or have more than four bel- to attach along the inferior margin of the right mandible.
lies due to variations in the anterior belly (see the entry for Colacino and Pettersen (1978), Pettersen (1979), and
digastricus anterior). Pettersen et al. (1979) report anomalies of the posterior belly
The extent of the origin of the posterior belly may vary, of the digastric in individuals with trisomy 13. In one neonate,
with possible attachments to the mastoid process or the the left posterior belly of the digastricus originated via two dis-
lateral part of the superior nuchal line (Macalister 1875; tinct bellies from the mastoid notch and from the styloid pro-
Bergman et al. 1988; Lee and Yang 2016). It may receive a cess and inserted with stylohyoid into the hyoid bone (Colacino
slip or originate entirely from the styloid process (Macalister and Pettersen 1978). The right posterior belly was split and
1875; De-Ary-Pires et al. 2003; Standring 2016). It may also sent a deep bundle to stylopharyngeus. In another neonate, the
receive a slip from the angle of the mandible (Macalister posterior belly of the digastric was doubled bilaterally, with
1875). Splenius may send fibers to its origin (Macalister the extra bellies inserting onto the hyoid bone (Pettersen et al.
1875). An accessory posterior belly of the digastric may 1979). On the left side of the specimen, the extra posterior
originate from the sternohyoid (Lee and Yang 2016) or from belly received a contribution from sternothyroid.
the mastoid notch with the normal posterior belly (Ozgursoy In a fetus with trisomy 13, the posterior bellies of the
and Kucuk 2006; Zhao et al. 2015). If the intermediate ten- digastric were doubled bilaterally (Pettersen et al. 1979).
don is absent, the posterior belly may attach onto the man- On the left side, each posterior belly had a separate origin
dible or the styloid process (Macalister 1875; Bergman et al. from the mastoid process and a separate insertion into the
1988; Lee and Yang 2016; Standring 2016). The posterior hyoid. On the right side, the superomedial belly inserted
belly may be absent (De-Ary-Pires et al. 2003). into the hyoid bone and the inferolateral belly inserted into
Stylohyoid may fuse with digastricus posterior or insert the infrahyoid muscles. In another fetus, the left posterior
onto the intermediate tendon (Macalister 1875; Bergman belly of the digastric inserted into the hyoid bone without
et al. 1988; Lee and Yang 2016). The intermediate tendon connecting to the anterior belly (Pettersen et al. 1979). The
may pass in front of or behind stylohyoid instead of perfo- same condition was found on the right side, with an exten-
rating it (Macalister 1875; Mori 1964; De-Ary-Pires et al. sion of the hyoid attachment of the posterior belly into the
2003; Harvey et al. 2015). The intermediate tendon may posterior aspect of hyoglossus. In a boy with trisomy 13q,
send slips to the lesser horn of the hyoid (Macalister 1875). the posterior belly on the left side of the body was small and
De-Ary-Pires et al. (2003) classified the posterior belly blended with the stylohyoid (Pettersen 1979).
of the digastric into two types, the intermediate tendon Digastricus posterior is also variable in some individuals
into three types, and the anterior belly of the digastric into with trisomy 18 (Bersu and Ramirez-Castro 1977; Aziz 1979;
five types (see the entry for digastricus anterior for more Smith et al. 2015). The intermediate tendon was absent bilat-
details). A type I posterior belly originated from the mas- erally in all cases examined by Bersu and Ramirez-Castro
toid notch. A type II posterior belly originated entirely or (1977), and all posterior bellies of the digastric had an inser-
partially from the styloid process and may have sent a slip tion into the hyoid when it was present. A tendinous inscrip-
to the middle or inferior constrictors of the pharynx. The tion into the posterior belly was found on the left side of one
intermediate tendon was categorized as piercing the stylo- infant (Bersu and Ramirez-Castro 1977). In another infant, the
hyoid muscle (type I), passing superficial to the stylohyoid right posterior belly passed deep to the external and internal
(type II), or passing deep to the stylohyoid (type III). carotid arteries. In a third, the muscle was absent on the right
side. Multiple infants exhibited slips extending from digastri-
Prevalence cus posterior to the styloid process and/or to the superior or
See digastricus anterior entry for prevalence information middle pharyngeal constrictors (Bersu and Ramirez-Castro
on the patterns described by De-Ary-Pires et al. (2003). 1977). In the fetus with trisomy 18 and cyclopia dissected by
Hallett (1848) found a third belly of the digastric muscle in Smith et al. (2015), the digastricus posterior bellies were broad
about 1 in 15 subjects (6.7%). Macalister (1875) notes that on both sides of the body but had normal attachments.
Wood found a third belly in 1 out of 17 subjects (5.9%). Sato Mieden (1982) describes an infant with median cleft
(1968b) found that the digastricus muscle was completely lip, hypotelorism, and alobar holoprosencephaly (case I)
absent in 7 out of 324 sides (2.16%) in Kyushu-Japanese and two male fetuses with cyclopia and alobar holoprosen-
males and in 2 out of 232 sides (0.86%) in females. cephaly (cases II and III). In case I, there was an accessory
muscle slip extending between the intermediate tendon of
Anomalies digastricus and the superior pharyngeal constrictor. In case
Description III, an accessory muscle belly was present along the hyoid
Bersu et al. (1976) describe a male infant with Hanhart syn- bone and extended between the right stylohyoid and left pos-
drome in which the right mandible was represented by a terior digastric muscles (Mieden 1982). On both sides of the
fetus with craniorachischisis, the posterior belly attached external carotid artery (Standring 2016). Its fibers may
to the “temporo-occipital” bone (Alghamdi et al. 2017). On occasionally cover the anterior aspect of the hyoid bone
the left side, the intermediate tendon of the digastric sent a (Ozgur et al. 2010) or end in either the suprahyoid or infra-
tendinous slip to the fascia over the left mylohyoid. hyoid musculature (Standring 2016). The intermediate
tendon of the digastric may pass in front of or behind stylo-
Prevalence hyoid instead of perforating it (Macalister 1875; Mori 1964;
De-Ary-Pires et al. 2003; Harvey et al. 2015). In this case,
the stylohyoid would have a single insertion onto the hyoid
digastricus posterior was variable in 5 out of 20 individu-
(Macalister 1875; Bergman et al. 1988; Lee and Yang 2016).
als with trisomy 13 (25%). Digastricus posterior sent slips
Stylohyoid may also fuse with digastricus posterior or insert
to either the styloid process and/or the superior or middle
onto the intermediate tendon (Macalister 1875; Knott 1883a;
pharyngeal constrictors in 4 out of 17 individuals with tri-
Bergman et al. 1988; Lee and Yang 2016). Hallett (1848)
somy 18 (23.5%). The intermediate tendon was absent in all
reports a case in which stylohyoid split upon its insertion,
eight infants (100%) with trisomy 18 described by Bersu
with the lower part inserting onto the body of the hyoid and
and Ramirez-Castro (1977).
the upper part inserting onto the intermediate tendon.
Stylohyoid is associated with named accessory muscles
and other accessory slips. Stylohyoid may send a slip to omo-
McMurtry and Yahr (1966) found that an accessory poste- hyoid, mylohyoid, or to the tongue (Macalister 1875). The
rior belly of the digastric coursed diagonally across the ori- stylohyoid may also insert into the outer border of mylohyoid
gin of the internal and external carotid arteries and caused (Macalister 1875). Stylohyoid may be doubled (Macalister
extracranial occlusion of the right internal carotid artery. 1875; Knott 1880; Standring 2016) with the second bundle
sometimes referred to as stylohyoideus profundus (Bergman
Stylohyoideus (Stylohyoid) (Figure 2.10) et al. 1988) or stylohyoideus nanus (Santorini) (Lee and Yang
2016). Stylohyoideus profundus may course parallel to the
Synonyms main bundle of stylohyoideus or replace the stylohyoid liga-
N/A ment (Macalister 1875; Bergman et al. 1988; Lee and Yang
2016). A second stylohyoid may have attachments to the
Typical Presentation base of the styloid process, the tip of the styloid process, and/
Description or to the lesser horn of the hyoid (Macalister 1875; Knott
Stylohyoid originates via a tendon from the posterior aspect 1880; Bergman et al. 1988; Lee and Yang 2016). Macalister
of the styloid process (Standring 2016). It inserts onto (1875) notes that Petsche found a second slip of stylohyoid
the hyoid at the junction of the body and the greater horn that inserted onto the cartilago triticea. A doubled stylo-
(Standring 2016). The intermediate tendon of the digastric hyoid may also send one slip over the intermediate tendon
perforates stylohyoid near its insertion (Standring 2016). and another slip under the intermediate tendon (Macalister
1875). Stylohyoid may also be tripled (Macalister 1875).
Innervation Stylochondrohyoideus refers to a supernumerary muscle
Stylohyoid is innervated by the stylohyoid branch of the that extends from the styloid process to the lesser horn of
facial nerve (Standring 2016). the hyoid (Lambert et al. 2010; Lee and Yang 2016) It may
replace, envelop, or have an attachment into, the stylohy-
Comparative Anatomy oid ligament (Joshi et al. 2007; Lambert et al. 2010; Lee
Stylohyoideus has a similar typical presentation in most and Yang 2016). Stylochondrohyoideus may be used inter-
apes, extending from the styloid process to the hyoid with changeably or confused with stylohyoideus profundus in
perforation at its insertion by the intermediate tendon of the the literature (Lambert et al. 2010). A second head for an
digastric (Sonntag 1923; Raven 1950; Miller 1952; Gibbs otherwise normal stylohyoid muscle may originate from the
1999; Diogo et al. 2010, 2013a,b, 2017). As the styloid pro- angle of the mandible (Macalister 1875). When this head is
cess is typically reduced or absent in gibbons, the stylo- completely distinct from the stylohyoid, it forms the hyoan-
hyoid muscle originates from the tympanic region (Diogo gularis muscle (Macalister 1875). Stylomandibularis refers
et al. 2012). It may originate from the temporal bone in to a bundle that extends from the tip of the styloid process
gorillas (Dean 1984). Stylohyoideus is not perforated by the to the angle of the mandible (Bergman et al. 1988; Lee and
intermediate tendon in orangutans (Sonntag 1924). Yang 2016).
Description Macalister (1875) notes that Hallett claims that stylohyoid
Stylohyoid may be absent (Macalister 1875; Knott 1880, is absent in 1 out of 200 cases (0.5%). From a sample of
1883a; Bergman et al. 1988; Lee and Yang 2016; Standring Japanese cadavers, Mori (1964) found that the stylohyoid
2016). In some cases, the muscle may pass medial to the passed medial to the intermediate tendon of the digastric
in 178 sides out of 254 sides (70.1%), passed lateral to the Prevalence
intermediate tendon on five sides (2%), and was perfo- In their literature review, Smith et al. (2015) found that sty-
rated by the intermediate tendon on 71 sides (27.9%). In lohyoideus was absent in 5 out of 20 individuals with tri-
a sample of 56 sides from 28 Anatolian cadavers, Ozgur somy 13 (25%) and 7 out of 17 individuals with trisomy 18
et al. (2010) found that the stylohyoid muscle inserted (41.2%). Stylohyoideus blended with the digastric in 2 out of
into the body of the hyoid at its junction with the greater 20 individuals with trisomy 13 (10%) and was anomalous in
horn on 34 sides (60.7%). In the remaining 22 sides 5 out of 17 individuals with trisomy 18 (29.4%).
(39.3%), fibers from stylohyoid elongated in front of the
hyoid and formed an arch or circle in front of it, having Clinical Implications
the appearance of covering the hyoid bone as a collar. A variant stylohyoid muscle can be confused with some
pathological conditions and may compress surrounding
Anomalies neurovascular structures, inducing stylohyoid syndrome
(Ozgur et al. 2010).
Description
On the right side of one neonate with trisomy 13, stylo-
hyoid did not bifurcate around the intermediate tendon of EXTRAOCULAR MUSCLES
the digastric (Colacino and Pettersen 1978). On the left
side, stylohyoid inserted with some fibers of the posterior
Levator palpebrae superioris (Figure 2.11)
belly of the digastric into the hyoid. On the left side of See also: Tensor trochleae
another neonate, stylohyoid was hypoplastic and inserted
into the intermediate tendon of the digastric (Pettersen Synonyms
et al. 1979). This morphology was present on the right, N/A
but a slip was present from the stylohyoid muscle to the
hyoid. Stylohyoid was absent bilaterally in a neonate with Typical Presentation
trisomy 13 (Aziz 1980). In one fetus with trisomy 13, sty- Description
lohyoid was absent on the left side (Pettersen et al. 1979). Levator palpebrae superioris originates via a tendon from
On the left side of another fetus, the stylohyoid inserted the lesser wing of the sphenoid (Standring 2016). It ends
into the hyoid and continued into the anterior belly of the in an aponeurosis that attaches to the superior tarsus,
digastric (Pettersen et al. 1979). In a boy with trisomy with some fibers inserting into the skin of the upper eye-
13q, the posterior belly of the digastric on the left side lid (Standring 2016). Anteriorly, the fascia between levator
of the body was small and blended with the stylohyoid palpebrae superioris and the superior rectus attaches to the
(Pettersen 1979). superior conjunctival fornix, which may be considered a
Stylohyoid was absent either bilaterally or unilaterally in third attachment of this muscle (Standring 2016).
nearly all the infants with trisomy 18 studied by Bersu and
Ramirez-Castro (1977). In two cases, stylohyoid muscles Innervation
were doubled unilaterally. When present, the stylohyoid Levator palpebrae superioris is innervated by a branch from
was not perforated by the digastric tendon, except for in the the superior division of the oculomotor nerve (Standring 2016).
following case. On the right side of one infant, an acces-
sory stylohyoid muscle originated from the temporal bone Comparative Anatomy
(Bersu and Ramirez-Castro 1977). This muscle gave origin In common chimpanzees, levator palpebrae superioris has
to a tendon that split around digastricus and then attached attachments only into the superior tarsus and superior con-
to the body of the hyoid near its junction with the greater junctival fornix, lacking an insertion into the skin of the eyelid
horn. The accessory stylohyoid muscle blended with the (Sonntag 1923). All three attachments are present in orang-
normal stylohyoid muscle. On the left side of this specimen, utans and the muscle also gives off a lateral band and medial
stylohyoid was absent. In the female fetus with trisomy 18 band (Sonntag 1924). The lateral band splits the lacrimal gland
dissected by Alghamdi et al. (2018), stylohyoid was absent and attaches to the deep surface of the malar bone. The medial
bilaterally. band divides into two slips that surround the lacrimal sac and
Mieden (1982) describes an infant with median cleft insert into the lacrimal bone. Some fibers of levator palpebrae
lip, hypotelorism, and alobar holoprosencephaly (case I) superioris fused with superior rectus (Sonntag 1924).
and two male fetuses with cyclopia and alobar holoprosen-
cephaly (cases II and III). Stylohyoideus was absent bilater-
ally in case I and absent on the left side in case III. In case Variations
III, an accessory belly was present along the hyoid bone Description
and extended between the right stylohyoid and left pos- Levator palpebrae superioris may be fused with rectus supe-
terior digastric muscles (Mieden 1982). In an infant with rior at its origin (Macalister 1875). It may originate from the
mandibulofacial dysostosis, stylohyoid received a slip from posterior margin of the frontal part of the roof of the orbit
digastricus posterior (Herring et al. 1979). (Macalister 1875). Macalister (1875) observed a case in which
FIGURE 2.11 Superior view (above) and inferior view (below) of the extraocular muscles.
levator palpebrae superioris only exhibited an insertion into Lüdinghausen et al. 1999; Haładaj et al. 2020a). An acces-
the superior conjunctival fornix. Levator palpebrae superi- sory muscle of levator palpebrae superioris that attaches
oris may be absent (Macalister 1875; Bergman et al. 1988). to the superior oblique may also be referred to as obliquus
Levator palpebrae superioris may also be split, doubled, or accessorius, obliqui superioris (Albinus), gracillimus orbitis
associated with accessory slips (Macalister 1875; Knott (Bochdalek), or gracillimus oculi (Macalister 1875; Knott
1883a; Whitnall 1921; Bergman et al. 1988; Amonoo-Kuof 1883a; Bergman et al. 1988). A variant of this slip, tensor
and Darwish 1998; von Lüdinghausen et al. 1999; Plock trochleae, may insert into the trochlea of superior oblique
et al. 2005; Yalçin et al. 2009; Kocabiyik 2016; Haładaj et al. (see the entry for this muscle). Levator palpebrae superi-
2020a). It may have a bipartite origin (Haładaj et al. 2020a). oris may also be connected with transversus orbitis, which
Amonoo-Kuof and Darwish (1998) describe a bilat- extends from the os planum of the ethmoid bone to the lateral
eral supernumerary muscle situated between the superior wall of the orbit, coursing across the superior surface of the
oblique and levator palpebrae superioris. These ‘levator pal- eyeball (Bochdalek 1868; Macalister 1875; Whitnall 1921;
pebrae superioris accessorius’ muscles originated from the Bergman et al. 1988; Kocabiyik 2016). Haładaj et al. (2020a)
lesser wing of the sphenoid and inserted into the medial por- describe a slip attaching to the lateral wall of the orbit that
tion of the skin of the eyelid (Amonoo-Kuof and Darwish resembles a partially developed transversus orbitis muscle.
1998). The three accessory slips described by Yalçin et al.
(2009) include a lateral slip divided anteriorly into superior Prevalence
and inferior parts, a medial fbromuscular slip, and slip situ- In a sample of 20 sides of 10 fetuses, Plock et al. (2005)
ated between the levator and superior rectus that ended in a found medial accessory bellies of levator palpebrae superi-
wide aponeurosis. This aponeurosis attached to the medial oris on seven sides (35%), and singular muscle fbers within
orbital septum, palpebral part of orbicularis oculi, and the medial fbrous bands on fve sides (25%). Out of 35 cadav-
medial skin of the upper eyelid (Yalçin et al. 2009). ers, Amonoo-Kuof and Darwish (1998) found bilateral
A slip of levator palpebrae superioris may extend to the levator palpebrae superioris accessorius muscles in one
lacrimal gland and may be referred to as retractor glandu- cadaver (2.9%). Yalçin et al. (2009) found accessory slips
lae lacrimalis (Whitnall 1921; Bergman et al. 1988; von of levator palpebrae superioris in 3 out of 60 orbits (5%).
In a sample of 70 orbits, Haładaj et al. (2020a) found oblique (Budge 1859; Macalister 1875; Knott 1883a; Haładaj
a typical presentation of levator palpebrae superioris in 58 et al. 2020a). It may also have attachments into the sclera
orbits (82.8%). Lateral slips of levator palpebrae superioris and fascia near the trochlea, the supratrochlear artery, or the
that inserted into the lacrimal gland were present in seven medial rectus muscle (Whitnall 1921; Isomura 1977; Sacks
specimens (10%). In one case (1.4%), the muscle sent a slip 1985; Bergman et al. 1988; von Lüdinghausen 1998; von
to the lateral wall of the orbit. In another case (1.4%), leva- Lüdinghausen et al. 1999; Kocabiyik 2016). Tensor troch-
tor palpebrae superioris had a bipartite origin. leae may be distinct from levator palpebrae superioris only
at its insertion, or for most of its length (Macalister 1875).
Anomalies Tensor trochleae may be split into two bundles (Whitnall
1921; von Lüdinghausen 1998). In a case observed by
Description
Whitnall (1921), tensor trochleae consisted of two bundles,
In a fetus with 18p-, levator palpebrae superioris was one that inserted into the fascia bulbi near the superior
absent on the left side and reduced in size on the right side oblique and one that inserted beneath the trochlea. In the
(Urban and Bersu 1987). In a fetus with prosencephaly, von case described by von Lüdinghausen (1998), the muscle
Lüdinghausen et al. (1999) found a bipartite levator pal- split at its midpoint, with a medial portion that became lig-
pebrae superioris that formed a retrobulbar arch. Levator amentous and inserted into the tissue between the trochlea
palpebrae superioris was absent bilaterally in a fetus with and the eyeball, and a lateral portion that rejoined levator
occipital meningocele (Plock et al. 2007). In anenceph- palpebrae superioris.
alic fetuses, levator palpebrae superioris may be reduced,
absent, or have insertions into the lacrimal gland and peri-
orbita (Plock et al. 2007). Innervation
Tensor trochleae is innervated by a branch from the supe-
Prevalence
rior division of the oculomotor nerve (Isomura 1977; Sacks
In a sample of 15 anencephalic fetuses, levator palpebrae 1985).
superioris was absent bilaterally in six fetuses (40%) and
absent on one side in one fetus (6.7%) (Plock et al. 2007).
Prevalence
Clinical Implications Isomura (1977) found tensor trochleae in fve orbits from
Variations of levator palpebrae superioris (including 85 subjects (5.9%). Sacks (1985) found tensor trochleae in
absence, defciency, or accessory slips) may cause ptosis or 7 out of 98 cadaver orbits (7.1%). Haładaj et al. (2020a)
congenital eyelid retraction (Urban and Bersu 1987; Wylen found tensor trochleae in 3 out of 70 orbits (4.3%), and
et al. 2001; Plock et al. 2005; Yalçin et al. 2009). Haładaj et al. (2020c) found tensor trochleae in 2 out of 78
orbits (2.6%).
tensOr trOchleae (figure 2.11) Anomalies

Description
See also: Levator palpebrae superioris
Macalister (1875) notes that in 1724, Kulmus found a mus-
Synonyms cle corresponding to tensor trochleae in an abnormal fetus.
This muscle may also be referred to as rectus quintus Prevalence
(Molinetti) (Knott 1883a) or the levator-trochlear mus-
N/A
cle (Sacks 1985). Tensor trochleae is a variant of the
accessory muscle referred to as comes obliquus acces- Clinical Implications
sorius, obliqui superioris (Albinus), gracillimus orbitis
Sacks (1985) argued that it may be possible for tensor troch-
(Bochdalek), or gracillimus oculi (Macalister 1875; Knott
leae to be a cause of some hypertropias or unexplained ocu-
1883a; Whitnall 1921).
lar deviations.
This muscle is only present as a variation or anomaly.
rectus superiOr (superiOr rectus) (figure 2.11)
von Lüdinghausen (1998) states that tensor trochleae and N/A
similar muscles are remnants of the membrana nictitans
(third eyelid) of amniotes. Typical Presentation
Description
Variations
Superior rectus originates from the common tendinous
Description ring and the dural sheath surrounding the optic nerve
Tensor trochleae presents as a medial bundle of levator pal- and inserts into the upper surface of the sclera (Standring
pebrae superioris that inserts into the trochlea of the superior 2016).
Innervation Prevalence
Superior rectus is innervated by the superior division of the Diamond et al. (1980) found superior rectus absent in three
oculomotor nerve (Standring 2016). out of fve patients (60%) with craniofacial dysostosis. In
a sample of 15 anencephalic fetuses, superior rectus was
Comparative Anatomy absent on the left side in one fetus (6.7%) (Plock et al. 2007).
In common chimpanzees, orangutans, and gibbons, superior
rectus has a similar typical presentation to that of humans Clinical Implications
(Sonntag 1923, 1924; Diogo et al. 2012). In orangutans, some The absence of superior rectus may cause ptosis (Posey 1923).
fbers fuse with levator palpebrae superioris (Sonntag 1924).
rectus inferiOr (inferiOr rectus) (figure 2.11)
Variations
Superior rectus may be fused with levator palpebrae superi- N/A
oris at its origin (Macalister 1875). Drummond and Keech
(1989) describe a case in which superior rectus and superior Typical Presentation
oblique were absent in the right eye and variant in the left Description
eye. Nayak et al. (2019) describe a double-bellied superior Inferior rectus originates from the common tendinous ring
rectus. The muscle had two bellies with separate origins and inserts into the sclera below the cornea (Standring 2016).
from the common tendinous ring that joined prior to inser-
tion (Nayak et al. 2019). Innervation
Accessory slips that originate from the tendinous Inferior rectus is innervated by a branch from the inferior
ring may connect the superior rectus with inferior rectus division of the oculomotor nerve (Standring 2016).
(Whitnall 1921; Bergman et al. 1988; von Lüdinghausen
1998; von Lüdinghausen et al. 1999; Kakizaki et al. 2006; Comparative Anatomy
Kocabiyik 2016; Haładaj et al. 2018; 2020b). These muscles In common chimpanzees and gibbons, inferior rectus has
may divide into two heads, one that inserts into superior a similar typical presentation to that of humans (Sonntag
rectus and one that inserts into inferior rectus (Kakizaki 1923; Diogo et al. 2012). In orangutans, inferior rectus is
et al. 2006; Haładaj et al. 2018). cylindrical and has a more intimate attachment to the globe
Superior rectus is associated with retractor bulbi, than the other extraocular muscles (Sonntag 1924).
which has numerous presentations (Bergman et al. 1988;
Kocabiyik 2016). It may present as a slip from the common Variations
tendinous ring to the posterior globe, a slip between the Description
superior rectus and lateral rectus, four slips that attach to Inferior rectus may be absent (Ingham et al. 1986; Muñoz
the back of the globe, or four slips that insert into the deep 1996; Taylor and Kraft 1997; Astle et al. 2003; Matsuo et al.
surfaces of the recti (Whitnall 1911; Bergman et al. 1988; 2009). Accessory slips that originate from the tendinous
Valmaggia et al. 1996; Krasny et al. 2011; Kocabiyik 2016). ring may connect the inferior rectus with superior rectus
(Whitnall 1921; Bergman et al. 1988; von Lüdinghausen
Prevalence 1998; von Lüdinghausen et al. 1999; Kakizaki et al. 2006;
In a sample of 70 hemiheads, Haładaj et al. (2020b) found Kocabiyik 2016; Haładaj et al. 2018; 2020b). These muscles
slips that connected superior rectus and inferior rectus in may divide into two heads: one that inserts into superior
two cases (2.9%). rectus and one that inserts into inferior rectus (Kakizaki
et al. 2006; Haładaj et al. 2018). Inferior rectus may also
Anomalies receive a fascicle from lateral rectus (Whitnall 1921;
Description Bergman et al. 1988; Kocabiyik 2016). Macalister (1875)
observed the partial fusion of inferior rectus and medial
Superior rectus may be absent in individuals with Apert syn-
rectus in the posterior third of the orbit. There may be a
drome, craniofacial dysostosis, or anencephaly (Weinstock
muscular bridge between the inferior rectus and inferior
and Hardesty 1965; Cuttone et al. 1979; Diamond et al.
oblique (Yalçin et al. 2004). It may also receive a slip from
1980; Plock et al. 2007). In a patient with oculocutaneous
an accessory inferior oblique (Whitnall 1921). Inferior rec-
albinism, superior rectus had a bifd insertion bilaterally
tus may be attached to retractor bulbi, which has numerous
(Verma and Hertle 2014).
presentations (Bergman et al. 1988; Kocabiyik 2016; see the
In two fetuses with triploidy, the attachments of the
entry for superior rectus for more information).
recti onto the sclera were shifted posteriorly (Moen et al.
1984). In a fetus with occipital meningocele, the left supe-
rior rectus was very thin (Plock et al. 2007). In fetuses with Prevalence
anencephaly, there was hypoplasia of superior rectus (Plock In a sample of 60 orbits, Yalçin et al. (2004) found a mus-
et al. 2007). In a fetus with trisomy 18 and cyclopia, supe- cular bridge between inferior rectus and inferior oblique in
rior rectus was doubled bilaterally (Smith et al. 2015). four orbits (6.7%). In a sample of 70 hemiheads, Haładaj
et al. (2020b) found slips that connect superior rectus and be absent (Bergman et al. 1988; Lee et al. 2013; Kocabiyik
inferior rectus in two cases (2.9%). 2016). Macalister (1875) observed the partial fusion of infe-
rior rectus and medial rectus in the posterior third of the
Anomalies orbit. A fascicle of lateral rectus may join with medial rec-
Description tus (Whitnall 1921; Bergman et al. 1988; Kocabiyik 2016).
Inferior rectus muscles were thinner than normal in fetuses An accessory medial rectus may be present superior and
with anencephaly (Plock et al. 2007) and in a fetus with trisomy posterior to the insertion of medial rectus (Lee and Kim
18 and cyclopia (Smith et al. 2015). In a fetus with prosenceph- 2009). Tensor trochleae may send a slip to medial rectus
aly, there was a tripartite inferior rectus (von Lüdinghausen (Sacks 1985).
et al. 1999). Its lateral belly blended with the posterior tendon
of inferior oblique and its medial belly attached medial to the
main belly of inferior rectus (von Lüdinghausen et al. 1999). Prevalence
In two fetuses with triploidy, the attachments of the recti In a sample of 16 patients with aplasia of the inferior rectus
onto the sclera were shifted posteriorly (Moen et al. 1984). muscle, the insertion of medial rectus was shifted inferiorly
Diamond et al. (1980) noted the absence of the right inferior in five patients (31.3%) (Matsuo et al. 2009).
rectus in a child with craniofacial dysostosis. In a child with
Axenfeld-Rieger syndrome, there was hypoplasia of inferior Anomalies
rectus on the right side (Bhate and Martin 2012). Description
Prevalence In a child with craniosynostosis, the left medial rectus was
bifurcated at its insertion (Coats and Ou 2001). In fetuses
Diamond et al. (1980) found inferior rectus absent in one
with anencephaly, there was hypoplasia of medial rectus
out of five patients (20%) with craniofacial dysostosis.
(Plock et al. 2007). Medial rectus was absent bilaterally in
Clinical Implications a fetus with trisomy 18 and cyclopia (Smith et al. 2015). In
an individual with trisomy 21, medial rectus was doubled
The absence of inferior rectus may mimic inferior rectus paresis
on the right side (Fernández-de-Luna et al. 2020). In two
and cause strabismus and compensatory head posture (Muñoz
fetuses with triploidy, the attachments of the recti onto the
1996; Taylor and Kraft 1997; Astle et al. 2003). The absence of
sclera were shifted posteriorly (Moen et al. 1984).
inferior rectus may also be associated with microphthalmos,
microcornea, or coloboma (Matsuo et al. 2009).
Prevalence
N/A
Rectus medialis (Medial rectus) (Figure 2.11)
Synonyms
The absence of medial rectus may be associated with diver-
This muscle may also be referred to as rectus internus
gent strabismus (Macalister 1875; Le Double 1897; Lee et al.
(Macalister 1875).
2013). A bifid insertion of medial rectus may contribute to
Typical Presentation intermittent distance exotropia (Sundaram et al. 2005). An
accessory medial rectus muscle may be present in cases of
Description strabismus fixus convergens (Lee and Kim 2009). An infe-
Medial rectus originates from the common tendinous ring riorly shifted insertion of medial rectus may be associated
and the dural sheath surrounding the optic nerve and inserts with microcornea and iris coloboma (Matsuo et al. 2009).
into the medial surface of the sclera (Standring 2016).
Innervation Rectus lateralis (Lateral rectus) (Figure 2.11)

Medial rectus is innervated by a branch from the inferior Synonyms
division of the oculomotor nerve (Standring 2016).
This muscle may also be referred to as rectus externus
Comparative Anatomy (Macalister 1875).
In common chimpanzees, orangutans, and gibbons, medial Typical Presentation
rectus has a similar typical presentation to that of humans
Description
(Sonntag 1923, 1924; Diogo et al. 2012).
Lateral rectus originates from the common tendinous ring
and greater wing of the sphenoid and inserts into the lateral
Variations surface of the sclera (Standring 2016).
Description
Medial rectus may be bifurcated at its insertion (Bergman Innervation
et al. 1988; Sundaram et al. 2005). Its insertion may also be Lateral rectus is innervated by the abducens nerve (Standring
shifted inferiorly (Matsuo et al. 2009). Medial rectus may 2016).
Comparative Anatomy Obliquus oculi superior

In common chimpanzees, orangutans, and gibbons, lateral (Superior oblique) (Figure 2.11)
rectus has a similar typical presentation to that of humans
See also: Tensor trochleae
(Sonntag 1923, 1924; Diogo et al. 2012). An accessory
lateral rectus is often present in macaque monkeys and Synonyms
considered a remnant of retractor bulbi (Schnyder 1984;
N/A
Boothe et al. 1990; Liao and Hwang 2014).
Variations
Description
Description
Superior oblique originates from the body of the sphenoid
Some authors may describe lateral rectus as having two (Standring 2016). The muscle ends in a tendon that courses
heads, as it arises via two tendinous bands (Haładaj through a trochlea (a cartilaginous loop or pulley) that is
2019). It may be bifurcated at its insertion (Duranoğlu attached to the frontal bone (Standring 2016). The tendon
and Gözkaya 2005). Its insertion may also be shifted inserts into the sclera deep to superior rectus (Standring 2016).
superiorly (Matsuo et al. 2009). Lateral rectus may be
absent (Bergman et al. 1988; Zöller et al. 2001; Kocabiyik Innervation
2016). Superior oblique is innervated by the trochlear nerve
Lateral rectus may send a fascicle to inferior rectus, (Standring 2016).
medial rectus, or the lateral wall of the orbit (Whitnall
1921; Bergman et al. 1988; Kocabiyik 2016). An accessory Comparative Anatomy
lateral rectus may be present (Macalister 1875; Bergman In common chimpanzees and gibbons, superior oblique
et al. 1988; Park and Oh 2003; Liao and Hwang 2014). has a similar typical presentation to that found in humans
Lateral rectus is associated with retractor bulbi, which may (Sonntag 1923; Diogo et al. 2012). In orangutans, the supe-
present as a slip between the superior rectus and lateral rior oblique runs closely along the medial wall of the orbit
rectus (Bergman et al. 1988; Kocabiyik 2016). Nussbaum and its tendon is connected via fascia to the superior rec-
(1893) described an accessory muscle that originated with tus (Sonntag 1924). The tendon does not expand toward the
lateral rectus and split into three heads that united with the globe (Sonntag 1924).
superior, inferior, and lateral recti.
Variations
Prevalence
Description
In a sample of 16 patients with aplasia of the inferior rectus
The position and length of the tendinous insertion of
muscle, the insertion of lateral rectus was shifted superiorly
superior oblique are variable (Whitnall 1921; Howe 1923;
in one patient (6.3%) (Matsuo et al. 2009).
Kocabiyik 2016; Haładaj et al. 2020c). The insertion may
Anomalies also be bifurcated (Park et al. 2009a). Superior oblique or
its tendon may be absent (Helveston et al. 1981; Matsuo
Description
et al. 1988; Drummond and Keech 1989; Wallace and
In fetuses with anencephaly, there was hypoplasia of lat- von Noorden 1994; Chan and Demer 1999). There may
eral rectus (Plock et al. 2007). In a fetus with trisomy 18 be a muscular band that connects the insertion of superior
and cyclopia, lateral rectus received one slip from inferior oblique with that of inferior oblique (Macalister 1875).
oblique on the left side, and two slips from inferior oblique Summarizing Le Double, Whitnall (1921) states that
on the right side (Smith et al. 2015). In two fetuses with there may be accessory fasciculi associated with the ten-
triploidy, the attachments of the recti onto the sclera were don of the superior oblique. In another case, the belly of
shifted posteriorly (Moen et al. 1984). In a child with superior oblique was absent, and the tendon was replaced
Axenfeld-Rieger syndrome, the left lateral rectus was by muscular fibers that originated from the trochlea
absent (Sandall and Morrison 1979). (Whitnall 1921). An accessory muscle of levator palpe-
brae superioris that attaches to the superior oblique may
Prevalence
also be referred to as comes obliquus accessorius, obliqui
N/A superioris (Albinus), gracillimus orbitis (Bochdalek),
or gracillimus oculi (Macalister 1875; Knott 1883a;
Bergman et al. 1988). A variant of this slip, tensor troch-
The absence of lateral rectus may be associated with conver- leae, may insert into the trochlea of superior oblique (see
gent strabismus (Macalister 1875; Whitnall 1921; Sandall the entry for this muscle).
and Morrison 1979; Zöller et al. 2001). Park and Oh (2003)
found that an accessory lateral rectus was associated with
exotropia, ptosis, pupil dilation, and limited eye movement. Prevalence
Symptoms associated with accessory lateral rectus muscles In a sample of 34 patients diagnosed with congenital supe-
may be diagnosed as exotropic Duane syndrome (Pineles rior oblique palsy, Helveston et al. (1981) found the superior
and Velez 2015; Neves and Curi 2019). oblique tendon absent on at least one side of the body in six
patients (17.6%). In a sample of 127 patients with strabis- Comparative Anatomy

mus, Chan and Demer (1999) found superior oblique absent Inferior oblique has a tendinous insertion in common
in six patients (4.7%). chimpanzees, but the muscle is entirely fleshy in orang-
utans (Sonntag 1923, 1924). In common chimpanzees and
Anomalies orangutans, it inserts more posteriorly onto the sclera than
Description in humans (Sonntag 1923, 1924). Diogo et al. (2012) found
The tendon of superior oblique may be absent in individu- that inferior oblique in gibbons is similar to that of humans.
als with anencephaly, trisomy 21, craniofacial dysostosis,
or neurofibromatosis (Barnes and Boniuk 1972; Diamond Variations
et al. 1980; Pinchoff and Sandall 1985; Pollard 1988; Description
Newman and Cogen 1997). In a child with Axenfeld-Rieger The origin of inferior oblique may be laterally displaced
syndrome, Park et al. (2009b) found that superior oblique (Whitnall 1921). Some fibers of inferior oblique may
inserted more posteriorly than normal. originate from the periosteal covering of the lacrimal sac
In two fetuses with triploidy, the attachments of the supe- (Whitnall 1921). The size and position of the insertion
rior oblique and inferior oblique were shifted anterior to are variable (Whitnall 1921; Howe 1923; Prakash et al.
those of the recti (Moen et al. 1984). Superior oblique was 1983; Paik and Shin 2009). Inferior oblique may have two
absent bilaterally in a fetus with trisomy 18 and cyclopia bellies (Wilson and Landers 1982; Bergman et al. 1988;
(Smith et al. 2015). In fetuses with anencephaly, the belly of DeAngelis et al. 1999; DeAngelis and Kraft 2001; Yalçin
superior oblique was markedly reduced, and the insertion and Ozan 2005a). Inferior oblique may also be divided
of the tendon was variable (Plock et al. 2007). In a fetus into three or more slips at its insertion (DeAngelis et al.
with occipital meningocele, the superior oblique muscles 1999; Yalçin and Ozan 2005a; Paik and Shin 2009).
were very thin (Plock et al. 2007). There may be a muscular band that connects the inser-
tion of superior oblique with that of inferior oblique
Prevalence
(Macalister 1875). There may also be a muscular bridge
Diamond et al. (1980) found the tendon of superior oblique between the inferior rectus and inferior oblique (Yalçin
absent bilaterally in one out of five patients (20%) with cra- et al. 2004). An accessory inferior oblique, or obliquus
niofacial dysostosis. Out of seven patients with Apert syn- accessorius inferior, extends between the apex of the orbit
drome and bilateral superior oblique palsy, Pollard (1988) and inferior oblique and may send a slip to inferior rectus
found that the tendon of superior oblique was absent bilater- (Rex 1887; Whitnall 1921; Bergman et al. 1988; Kocabiyik
ally in five patients (71.4%) and superior oblique was replaced 2016; Richardson et al. 2017). The case described by
by a small fibrous band bilaterally in two patients (28.6). Richardson et al. (2017) inserted into the inferior oblique
Clinical Implications and the lower eyelid.
The absence of superior oblique or its tendon may mimic Prevalence
superior oblique palsy and be associated with vertical In a sample of 100 orbits, Whitnall (1921) found that the
deviations, horizontal strabismus, or amblyopia (Helveston origin of inferior oblique was adjacent (<1 mm away) to
et al. 1981; Matsuo et al. 1988; Pollard 1988; Wallace and the nasolacrimal fossa in 45 cases (45%), 2 mm away from
von Noorden 1994; Chan and Demer 1999). A bifid inser- the edge of the nasolacrimal fossa in 14 cases (14%), 3 mm
tion of the superior oblique may cause congenital Brown away in 19 cases (19%), 4 mm away in eight cases (8%),
syndrome (Park et al. 2009a). 5 mm away in six cases (6%), 6 mm away in four cases (4%),
and 7 mm away in four cases (4%).
Obliquus oculi inferior (Inferior In a sample of 100 orbits, DeAngelis et al. (1999) found
that 17 inferior oblique muscles (17%) had multiple divi-
oblique) (Figure 2.11)
sions upon insertion. Eight inferior oblique muscles (8%)
had two bellies. In four of these cases, the two bellies shared
Synonyms an insertion and in the other four cases, the two bellies had
N/A separate insertions (DeAngelis et al. 1999). DeAngelis and
Kraft (2001) found 27 doubled-bellied inferior oblique
Typical Presentation muscles out of a sample of 247 muscles (10.9%).
In a sample of 60 orbits, Yalçin et al. (2004) found a mus-
Description
cular bridge between inferior rectus and inferior oblique
Inferior oblique originates from the orbital surface of the in four orbits (6.7%). In a sample of 60 orbits, Yalçin and
maxilla and inserts into the inferolateral surface of the Ozan (2005a) studied how many bellies of inferior oblique
sclera (Standring 2016). were present at insertion. There was a single belly in five
cases (8.3%), two bellies in 30 cases (50%), three bellies
Innervation in 16 cases (26.7%), and more than three bellies in nine
Inferior oblique is innervated by a branch from the inferior cases (15%). Paik and Shin (2009) studied how many bel-
division of the oculomotor nerve (Standring 2016). lies of inferior oblique were present at insertion in 62 orbits.
Inferior oblique had a single belly in 44 cases (71%), two Innervation

bellies in 14 cases (22.5%), three bellies in two cases (3.2%), Scalenus anterior is innervated by the ventral rami of the
and four bellies in two cases (3.2%). fourth through sixth cervical spinal nerves (Standring 2016).
Anomalies Comparative Anatomy

Description Scalenus anterior has a similar typical presentation in the
Inferior oblique may be absent in individuals with anen- apes, extending from the transverse processes of various
cephaly or craniofacial dysostosis (Diamond et al. 1980; cervical vertebrae to the first rib (Stewart 1936; Sonntag
Plock et al. 2007). In fetuses with anencephaly that had infe- 1923, 1924; Miller 1952; Gibbs 1999; Diogo et al. 2010,
rior oblique muscles, Diamond et al. (1980) found they were 2012, 2013a,b, 2017). The origin is typically from the third
reduced in size. In two fetuses with triploidy, the attach- through fifth or fourth through sixth cervical vertebrae
ments of the superior oblique and inferior oblique were (Stewart 1936; Sonntag 1923, 1924; Miller 1952; Gibbs
shifted anterior to those of the recti (Moen et al. 1984). In 1999). It may originate from only the fifth and sixth cer-
a fetus with trisomy 18 and cyclopia, both inferior oblique vical vertebrae in gorillas (Raven 1950). There may be an
muscles sent a slip to lateral rectus and had another slip that attachment to the seventh cervical vertebra in common
fused with its counterpart on the other side (Smith et al. chimpanzees (Champneys 1872).
2015). On the right side, an additional slip inserted into lat-
eral rectus, and on the left side, a slip passed to an accessory Variations
band of muscle parallel to the limbus of the eye (Smith et al. Description
2015). In an infant with mandibulofacial dysostosis, infe-
The origin of scalenus anterior may vary in terms of which
rior oblique originated from an enlarged and poorly ossified
vertebrae contribute fascicles to the muscle (Macalister 1875;
fossa for the lacrimal sac (Herring et al. 1979).
Knott 1883a; Mori 1964; Rusnak-Smith et al. 2001; Bergman
Prevalence et al. 1988; Sakamoto 2012, 2016b). Its costal attachment
Diamond et al. (1980) found the inferior oblique absent may also vary (Bergman et al. 1988; Wayman et al. 1993;
bilaterally in one out of five patients (20%) with craniofacial Sakamoto 2016b). Width of the anterior scalene varies
dysostosis. Plock et al. (2007) found inferior oblique absent among individuals (Rusnak-Smith et al. 2001). The lower
unilaterally in 2 out of 15 anencephalic fetuses (13.3%). third of the muscle may be tendinous (Macalister 1875). The
muscle may be divided into two bellies (Macalister 1875;
Clinical Implications Goubran et al. 2010; Sakamoto 2016b; Perumal et al. 2018).
The absence of inferior oblique may cause ptosis (Posey Scalenus anterior may exchange fibers with scalenus
1923). DeAngelis and Kraft (2001) found that eyes with medius or scalenus minimus (Wood 1867b; Macalister 1875;
double-bellied inferior oblique muscles had a higher Rajanigandha et al. 2008; Sakamoto 2012, 2016b). Its fibers
incidence of fundus excyclotropia. A band from the apex may be continuous with longus capitis or the intertransverse
of the orbit to the inferior oblique and lower eyelid may muscles (Macalister 1875; Rusnak-Smith et al. 2001; Sakamoto
cause congenital paradoxical lower eyelid retraction 2016b). Scalenus anterior may be absent (Macalister 1875;
(Richardson et al. 2017). A variant insertion of inferior Murakami et al. 2003; Collins et al. 2014). In the case reported
oblique may be associated with divergent strabismus by Murakami et al. (2003), scalenus anterior was replaced by
(Prakash et al. 1983). aberrant muscle slips that originated from the lower cervical
vertebrae and joined with scalenus medius.
SCALENE, PREVERTEBRAL, AND An accessory anterior scalene muscle may be present
SUBOCCIPITAL MUSCLES (Bergman et al. 1988; Sakamoto 2016b). Scalenus ante-
rior may be associated with transversalis cervicis medius
Scalenus anterior (Anterior scalene) (Figure 2.12) (Krause), which originates from the second and fourth cer-
vical vertebrae and inserts into the seventh and eighth cer-
See also: Roos band, scalenus minimus
vical vertebrae (Knott 1883a; Bergman et al. 1988). A Roos
Synonyms band may extend from anterior scalene to subclavius and
This muscle may also be referred to as scalenus anticus the costal cartilage (see the entry for this muscle).
(Macalister 1875). Prevalence
In a sample of 240 sides (120 cadavers), Mori (1964) found
that scalenus anterior arose from the second through fifth
Description cervical vertebrae on eight sides (3.3%), from the third
Scalenus anterior originates from the anterior tubercles through fifth cervical vertebrae on 108 sides (45%), and
of the transverse processes of the third through sixth from the fourth through sixth cervical vertebrae on 124
cervical vertebrae (Standring 2016). It inserts onto the sides (51.7%). In a sample of ten cadavers, Rusnak-Smith
scalene tubercle and superior margin of the first rib et al. (2001) found that scalenus anterior arose from the
(Standring 2016). third through sixth cervical vertebrae in two cases (20%),
from the third through seventh cervical vertebrae in three Comparative Anatomy
cases (30%), from the fourth and fifth cervical vertebrae in In the apes, scalenus medius and scalenus posterior are
two cases (20%), and from the fourth through sixth cervical fused, and the combined muscle complex typically origi-
vertebrae in three cases (30%). Tendinous fibers of scalenus nates from the second through sixth or seventh cervi-
anterior blended with longus capitis in one case (10%). cal vertebrae and inserts onto the first rib (Sonntag 1923,
In a sample of 52 cadaveric sides, Sakamoto (2012) found 1924; Stewart 1936; Miller 1952; Jouffroy 1971; Gibbs
that scalenus anterior originated from the fifth and sixth 1999; Diogo et al. 2010, 2012, 2013a,b, 2017). In common
cervical vertebrae on three sides (5.8%), from the fourth chimpanzees, there may be an insertion onto the second rib
through sixth cervical vertebrae on 26 sides (50%), and (Macalister 1871). In one common chimpanzee, Sonntag
from the third through sixth cervical vertebrae on 23 sides (1923) found that the scalenus medius-posterior muscle
(44.2%). Scalenus anterior sent fibers to scalenus minimus sheet attached to the first five ribs.
on one side (1.9%). Scalenus anterior and the ventral part of
scalenus medius exchanged fibers on seven sides (13.5%). Variations
Description
Anomalies
The origin of scalenus medius may vary in terms of which
Description vertebrae contribute fascicles to the muscle (Macalister
The right anterior scalene muscle was absent in a fetus with 1875; Knott 1883a; Cave 1933; Bergman et al. 1988; Rusnak-
craniorachischisis (Alghamdi et al. 2017). Smith et al. 2001; Sakamoto 2012, 2016b; Standring 2016).
It often has an origin from the transverse process of the first
Prevalence cervical vertebra (Wood 1867b; Macalister 1875; Cave 1933;
N/A Bergman et al. 1988; Rusnak-Smith et al. 2001; Sakamoto
2016b; Standring 2016). Fibers of scalenus medius may
Clinical Implications originate from a cervical rib when this structure is pres-
An anteriorly shifted insertion of scalenus anterior may ent (Tokat et al. 2011). The insertion of scalenus medius
cause venous compression syndromes in the root of the onto the first rib may be shifted anteriorly to just behind the
neck (Wayman et al. 1993). Blending of some anterior sca- anterior scalene (Thomas et al. 1983). It may also insert par-
lene fibers with longus capitis may cause tension and head- tially or entirely into the second rib (Macalister 1875; Knott
aches in the occipital region (Rusnak-Smith et al. 2001). A 1883a; Sakamoto 2012, 2016b). Width of the middle sca-
relatively wide scalenus anterior muscle may impinge the lene varies among individuals (Rusnak-Smith et al. 2001).
surrounding structures, leading to vascular compromise Scalenus medius may be absent (Macalister 1875).
causing symptoms of coldness and pallor (Rusnak-Smith Scalenus medius may exchange fibers with scalenus ante-
et al. 2001). Compression of the subclavian artery between rior (Wood 1867b; Macalister 1875; Rajanigandha et al. 2008;
the two bellies of a divided scalenus anterior may lead Sakamoto 2012, 2016b). Paraskevas et al. (2007) found an
to the development of vascular thoracic outlet syndrome accessory middle scalene that arose from the middle portion
(Goubran et al. 2010). The absence of scalenus anterior may of scalenus medius and inserted with scalenus anterior onto
present with the symptoms of thoracic outlet syndrome the first rib. Scalenus medius may be fused with scalenus
(Collins et al. 2014). A Roos band that arises from scalenus posterior (Macalister 1875; Knott 1883a; Cave 1933; Rusnak-
anterior and attaches to subclavius and the costal cartilage Smith et al. 2001; Sakamoto 2016b; Standring 2016). It may
may cause subclavian vein compression (Spears et al. 2011). also join with multifidus or semispinalis cervicis (Macalister
1875). It may send a slip to levator scapulae (Macalister 1875;
Wood 1867b, 1868). A Roos band may originate from the
Scalenus medius (Middle scalene) (Figure 2.12) middle scalene and attach to the first rib or its cartilage (see
See also: Roos band the entry for this muscle).
Synonyms Prevalence
N/A Out of 60 subjects, Cave (1933) found that scalenus medius
originated from all cervical vertebrae in 37 subjects
Typical Presentation (61.7%), from the second through seventh cervical verte-
Description brae in nine cases (15%), from the third through seventh
cervical vertebrae in six cases (10%), and from the third
Scalenus medius originates from the transverse processes
through sixth cervical vertebrae in six cases (10%). The
of the second through seventh cervical vertebrae (Standring
origins could not be determined in two cases (3.3%). In a
2016). It inserts into the superior margin of the first rib
sample of 29 arms, Thomas et al. (1983) found that scalenus
(Standring 2016).
medius inserted anteriorly and behind anterior scalene in
Innervation 14 arms (483%). In a sample of ten cadavers, Rusnak-Smith
Scalenus medius is innervated by the ventral rami of the et al. (2001) found that scalenus medius originated from
third through eighth cervical spinal nerves (Standring 2016). all cervical vertebrae in five cases (50%), from the second
through seventh cervical vertebrae in one case (10%), and Comparative Anatomy
from the second through sixth cervical vertebrae in four In the apes, scalenus medius and scalenus posterior are
cases (40%). Scalenus medius was fused with scalenus pos- fused, and the combined muscle complex typically origi-
terior in three cases (30%). nates from the second through sixth or seventh cervi-
In a sample of 52 cadaveric sides, Sakamoto (2012) cal vertebrae and inserts onto the first rib (Sonntag 1923,
found that the ventral part of scalenus medius originated 1924; Stewart 1936; Miller 1952; Jouffroy 1971; Gibbs
from the fourth through seventh cervical vertebrae on 42 1999; Diogo et al. 2010, 2012, 2013a,b, 2017). In common
sides (80.8%) and from the third through seventh cervical chimpanzees, there may be an insertion onto the second rib
vertebrae on ten sides (19.2%). The ventral part of scale- (Macalister 1871). In one common chimpanzee, Sonntag
nus medius inserted onto the first rib on 42 sides (80.8%) (1923) found that the scalenus medius-posterior muscle
and onto both the first and second ribs on ten sides sheet attached to the first five ribs.
(19.2%). Scalenus anterior and the ventral part of scale-
nus medius exchanged fibers on seven sides (13.5%). The
Variations
dorsal part of scalenus medius originated from the third
through seventh cervical vertebrae on two sides (3.8%), Description
from the second through seventh cervical vertebrae on The origin of scalenus posterior may vary in terms of which
seven sides (13.5%), and from all cervical vertebrae on 43 vertebrae contribute fascicles to the muscle (Mori 1964;
sides (82.7%). Bergman et al. 1988; Rusnak-Smith et al. 2001; Sakamoto
2012, 2016b). It inserts variously into the first three ribs
(Macalister 1875; Knott 1883a; Mori 1964; Bergman et al.
Anomalies 1988; Rusnak-Smith et al. 2001; Sakamoto 2012, 2016b).
Description The muscle may be divided into three separate portions, one
In a boy with trisomy 13q, scalenus medius inserted onto portion inserting onto each of the first three ribs (Macalister
the first and second ribs on the right side and to the first rib 1875). Width of the posterior scalene varies among individ-
only on the left side (Pettersen 1979). In a fetus with cranio- uals (Rusnak-Smith et al. 2001). Scalenus posterior may be
rachischisis, levator scapulae fused with the distal portion absent (Macalister 1875; Knott 1883a; Rusnak-Smith et al.
of scalenus medius on the left side (Alghamdi et al. 2017). 2001; Sakamoto 2012, 2016b).
Scalenus posterior may be fused with scalenus medius
(Macalister 1875; Knott 1883a; Cave 1933; Rusnak-Smith
Prevalence
et al. 2001; Standring 2016). It may also be fused with the
N/A first external intercostal muscle (Bergman et al. 1988).
Clinical Implications Scalenus posterior may also connect with serratus anterior,
levator scapulae, or the first levator costae (Wood 1868;
An anterior insertion of scalenus medius (Thomas et al. 1983)
Macalister 1875; Rickenbacher et al. 1985; Smith et al.
or the presence of an accessory middle scalene (Paraskevas
2003; Bakkum and Miller 2016).
et al. 2007) may cause thoracic outlet syndrome. Fusion of
scalenus medius and scalenus posterior may compromise the Prevalence
long thoracic and dorsal scapular nerves, leading to shoulder In a sample of 102 cadaveric sides, Mori (1964) found that
pain and weakness of levator scapulae, serratus anterior, and scalenus posterior inserted into the first rib on 21 sides
the rhomboid muscles (Rusnak-Smith et al. 2001). (20.6%), the first and second ribs on 71 sides (69.6%), and
the first three ribs on nine sides (8.8%). In a sample of ten
Scalenus posterior (Posterior scalene) (Figure 2.12) cadavers, Rusnak-Smith et al. (2001) found that scalenus
posterior originated from the fourth through sixth cervical
Synonyms vertebrae in two cases (20%), from the fourth and fifth cer-
This muscle may also be referred to as scalenus posticus vical vertebrae in two cases (20%), and from the fifth and
(Macalister 1875). sixth cervical vertebrae in four cases (40%). Scalenus poste-
rior inserted into the third rib in one case (10%). The muscle
Typical Presentation was absent in two cases (20%). Scalenus medius was fused
Description with scalenus posterior in three cases (30%).
Scalenus posterior originates from the posterior tubercles In a sample of 52 cadaveric sides, Sakamoto (2012) found
of the transverse processes of the fourth through sixth cer- that scalenus posterior originated from the fifth cervical
vical vertebrae (Standring 2016). It inserts onto the outer vertebra only on 25 sides (61%), from the fifth and sixth
aspect of the second rib (Standring 2016). cervical vertebrae on nine sides (21.9%), from the fourth
through sixth cervical vertebrae on four sides (9.8%), from
Innervation the fourth and fifth cervical vertebrae on two sides (4.9%),
Scalenus posterior is innervated by the ventral rami of and from the third and fourth cervical vertebrae on one side
the sixth through eighth cervical spinal nerves (Standring (2.4%). Scalenus posterior inserted onto the first rib on two
2016). sides (4.9%), onto the second rib on 28 sides (68.3%), onto
FIGURE 2.12 Lateral vertebral muscles in anterior view.
the first and second ribs on eight sides (19.5%), and onto the Scalenus minimus (Figure 2.12)
second and third ribs on three sides (7.3%). Scalenus poste-
rior was absent on 11 sides (21.2%). See also: Roos band
Synonyms
Anomalies This muscle may also be referred to as scalenus pleuralis
Description (Sibson) or scalenus intermedius (Testut) (Bergman et al.
1988; Standring 2016).
The origin of scalenus posterior may blend with leva-
tor scapulae in individuals with trisomy 18 (Smith et al. Typical Presentation
2015).
Prevalence Comparative Anatomy

In their literature review, Smith et al. (2015) found that the In a gorilla, Stewart (1936) reported a slip that extended
origin of scalenus posterior was blended with levator scapu- from the transverse processes of the sixth and seventh cer-
lae in 1 out of 26 individuals with trisomy 18 (3.8%). vical vertebrae to the first rib, which may correspond to sca-
lenus minimus. Diogo et al. (2017) found scalenus minimus
in one bonobo extending between the transverse process of
the seventh cervical vertebra and the first rib.
A relatively wide scalenus posterior muscle may affect
the long thoracic nerve and dorsal scapular nerve, causing
pain and tension in serratus anterior and levator scapulae Variations
(Rusnak-Smith et al. 2001). Fusion of scalenus medius Description
and scalenus posterior may also compromise the long Scalenus minimus originates from the transverse process
thoracic and dorsal scapular nerves, leading to shoulder of the seventh cervical vertebra and inserts onto the first rib
pain and weakness of levator scapulae, serratus anterior, just behind the groove for the subclavian artery, and into
and the rhomboid muscles (Rusnak-Smith et al. 2001). the cervical pleura (Fawcett 1896; Shore 1926; Stott 1928;
Boyd 1934; Rickenbacher et al. 1985; Bergman et al. 1988; Longus colli (Figure 2.13)
Rusnak-Smith et al. 2001; Natsis et al. 2013; Sakamoto
2012, 2016b; Standring 2016). Synonyms
It may sometimes have an origin from the sixth cervical This muscle may also be referred to as rectus colli
vertebra (Bergman et al. 1988; Rusnak-Smith et al. 2001; (Luschka) or longus cervicis (Macalister 1875; Rusnak-
Sakamoto 2012, 2016b). Some early accounts report that Smith et al. 2001).
scalenus minimus extends from the fifth through seventh
cervical vertebrae to the second rib (Macalister 1875; Knott Typical Presentation
1883a). Under the Roos band classification scheme, a type 5 Description
band is the scalenus minimus muscle attached to the first rib Longus colli is divided into inferior oblique, vertical
between the subclavian artery and the brachial plexus, and a (intermediate), and superior oblique parts (Standring
type 6 band is the scalenus minimus muscle attached only to 2016). The inferior oblique part originates from the bod-
the cervical pleura (Roos 1976; entry for this muscle). ies of the first three thoracic vertebrae and inserts onto
Scalenus minimus may exchange fibers with scalenus the transverse processes of the fifth and sixth cervical
anterior (Sakamoto 2012). If scalenus minimus is not pres- vertebrae (Standring 2016). The vertical part originates
ent, a fibrous band often referred to as the scalenus medius from the bodies of the fifth cervical through third tho-
band may take its place (Shore 1926; Bonney 1965; Thomas racic vertebrae and inserts onto the bodies of the second
et al. 1983; Bergman et al. 1988; Sakamoto 2016b). through fourth cervical vertebrae (Standring 2016). The
Innervation superior oblique part originates from the transverse pro-
Scalenus minimus is innervated by the seventh and eighth cesses of the third through fifth cervical vertebrae and
cervical spinal nerves (Sakamoto 2012). inserts onto the anterior tubercle of the first cervical ver-
tebra (Standring 2016).
Prevalence
Knott (1883a) found scalenus minimus in 5 out of 23 bodies Innervation
(21.7%). Stott (1928) found scalenus minimus on 39 out of 100 Longus colli is innervated by the ventral rami of the second
cadaveric sides (39%). In a sample of 30 cadaveric sides, Boyd through sixth cervical spinal nerves (Standring 2016).
(1934) found scalenus minimus on ten sides (33.3%). In a sam-
ple of 29 arms, Thomas et al. (1983) found scalenus minimus Comparative Anatomy
in one case (3%). In a sample of 102 cadaveric sides, Harry There is no information available for gibbons. In orang-
et al. (1997) found scalenus minimus in 46% of sides. In a utans, the superior oblique part is typically absent, while
sample of 10 cadavers, Rusnak-Smith et al. (2001) found sca- the inferior oblique and vertical parts originate from the
lenus minimus definitively present in one cadaver (10%) and seventh cervical through the fifth thoracic vertebrae and
potentially present in two other cadavers (20%). Natsis et al. insert onto the first cervical vertebra (Sonntag 1924). In
(2013) found scalenus minimus in 3 out of 73 cadavers (4.1%). gorillas, there is no origin from the thoracic vertebrae
Out of 241 patients that underwent surgery for thoracic (Gibbs 1999; Raven 1950; Diogo et al. 2010). The pre-
outlet syndrome, Roos (1976) found type 5 bands in 40 cases sentation in common chimpanzees and bonobos is most
(16.6%) and type 6 bands in five cases (2.1%). Out of 58 tho- like that of humans, with similar origins and insertions
racic outlet dissections in 29 cadavers, Roos (1976) found (Gibbs 1999; Sonntag 1923; Miller 1952; Diogo et al.
type 5 bands present in eight cases (13.8%). Out of 100 cases, 2013a, 2017).
Spears et al. (2011) found type 5 bands in 11 cases (11%).
In a sample of 52 cadaveric sides, Sakamoto (2012) found Variations
that scalenus minimus was present on 25 sides (48.1%). Out Description
of these 25 cases, it originated from the sixth cervical verte- Longus colli may vary in terms of which structures it arises
bra on two sides (8%), from the seventh cervical vertebra on from and inserts into (Macalister 1875; Rickenbacher et al.
14 sides (56%), and from both the sixth and seventh cervical 1985; Bergman et al. 1988; Sakamoto 2012, 2016b). It may
vertebrae on nine sides (36%). Scalenus anterior sent fibers have an origin from the body of the fourth thoracic vertebra
to scalenus minimus on one side (1.9%). (Eisler 1912; Rickenbacher et al. 1985; Sakamoto 2016b).
It may also originate from the heads of the first, second, or
Anomalies third rib (Eisler 1912; Rickenbacher et al. 1985; Sakamoto
N/A 2016b). The origins from the bodies of the thoracic vertebrae
or lower cervical vertebrae may be absent (Rickenbacher
Clinical Implications et al. 1985). The most superior bundles of the muscle may
Scalenus minimus may compress the brachial plexus and/or insert into the base of the skull (Rickenbacher et al. 1985).
subclavian artery and cause thoracic outlet syndrome (Levi Longus colli may be thin or absent (Macalister 1875).
1948; Harry et al. 1997; Rusnak-Smith et al. 2001; Natsis Longus colli may fuse with or send bundles to longus capi-
et al. 2013). tis (Macalister 1875; Rickenbacher et al. 1985; Bergman
et al. 1988; Sakamoto 2012, 2016b). It may be associated Comparative Anatomy

with an accessory intertransverse muscle referred to as There is no information available for gibbons. The mus-
transversalis cervicis anticus (Retzius), which extends from cle has a similar typical presentation to that of humans in
the fourth through sixth cervical vertebrae to the first three the other apes, extending from the transverses processes
cervical vertebrae (Macalister 1875; Knott 1883a; Bakkum of the cervical vertebrae to the basiocciput (Sonntag
and Miller 2016; Sakamoto 2016b). 1923, 1924; Raven 1950; Miller 1952; Dean 1985; Gibbs
1999; Diogo et al. 2010, 2013a,b, 2017). It may origi-
Prevalence nate from the second through sixth cervical vertebrae
In a sample of 52 cadaveric sides, Sakamoto (2012) in gorillas (Raven 1950) and from the second through
found that the vertical and inferior oblique parts were fifth (Diogo et al. 2017) or fourth through seventh cervi-
fused at their origins, with their lowermost origin from cal vertebrae (Miller 1952) in bonobos. It may receive a
the third thoracic vertebra on 15 sides (28.9%), from the slip from the anterior scalene in common chimpanzees
second thoracic vertebra on 36 sides (69.2%), and from (Sonntag 1923).
the first thoracic vertebra on one side (1.9%). The inferior
oblique part inserted onto the fifth and sixth cervical ver-
tebrae on 44 sides (84.7%), the sixth cervical vertebra on Variations
two sides (3.8%), the fourth and fifth cervical vertebrae Description
on two sides (3.8%), and the fourth through sixth cervi- Longus capitis may vary in terms of which structures it arises
cal vertebrae on four sides (7.7%). The vertical part origi- from and inserts into (Macalister 1875; Rickenbacher et al.
nated from the fifth cervical vertebra on one side (1.9%), 1985; Bergman et al. 1988; Sakamoto 2012, 2016b). It may have
from the fourth and fifth cervical vertebrae on three sides additional origins from the first and/or second cervical verte-
(5.8%), from the fifth and sixth cervical vertebrae on 18 brae (Macalister 1875; Rickenbacher et al. 1985; Sakamoto
sides (34.6%), from the fourth through sixth cervical ver- 2012, 2016b). The origin from the fifth or sixth cervical verte-
tebrae on 24 sides (46.2%), and from the third through brae may be absent (Macalister 1875; Rickenbacher et al. 1985;
sixth cervical vertebrae on six sides (11.5%). The superior Sakamoto 2016b). At its insertion, fibers may decussate with
oblique part originated from the third through fifth cervi- those of the contralateral muscle (Rickenbacher et al. 1985).
cal vertebrae on 26 sides (50%), the third and fourth cervi- Longus capitis may split into two muscles or multiple fascicles
cal vertebrae on 17 sides (32.7%), from the second through (Macalister 1875; Knott 1883a; Rickenbacher et al. 1985). It
fourth cervical vertebrae on one side (1.9%), from the sec- may also be bilaminar (Macalister 1875). Longus capitis may
ond through fifth cervical vertebrae on four sides (7.7%), fuse with longus colli (Macalister 1875; Rickenbacher et al.
and from the third through sixth cervical vertebrae on 1985; Bergman et al. 1988; Sakamoto 2012, 2016b). It may
four sides (7.7%). Longus capitis and the inferior oblique also connect with the anterior scalene, rectus capitis anterior,
part connected via tendinous fibers on 43 sides (82.7%). or intertransverse muscles (Macalister 1875; Bergman et al.
1988; Sakamoto 2016b).
Anomalies Longus capitis is associated with atlantobasilaris
N/A internus (Gruber), which originates from the anterior
tubercle of the first cervical vertebra, courses along
Clinical Implications the medial margin of longus capitis, and inserts near
N/A longus capitis into the base of the skull (Rickenbacher
et al. 1985; Bergman et al. 1988; Sakamoto 2016b). If
the insertion of this accessory muscle is shifted onto the
Longus capitis (Figure 2.13) anterior longitudinal ligament, it is referred to as axo-
Synonyms basilaris or axiobasilaris (Rickenbacher et al. 1985). If it
originates from the second cervical vertebra, it is referred
This muscle may also be referred to as rectus capitis anticus
to as epistropheobasilaris (Gruber) (Bergman et al. 1988;
major (Macalister 1875).
Sakamoto 2016b).
Prevalence
Description Likely summarizing the work of Gruber, Rickenbacher at al.
Longus capitis originates from the anterior tubercles of the (1985) states that atlantobasilaris internus is present in about
transverse processes of the third through sixth cervical ver- 4% of subjects and axobasilaris is present in about 2% of
tebrae and inserts into the basilar part of the occipital bone subjects. In a sample of 52 cadaveric sides, Sakamoto (2012)
(Standring 2016). found that longus capitis originated from the third through
sixth cervical vertebrae on 32 sides (61.5%) and from the
Innervation second through sixth cervical vertebrae on 20 sides (38.5%).
Longus capitis is innervated by the ventral rami of the first Longus capitis and the inferior oblique part of longus colli
through third cervical spinal nerves (Standring 2016). connected via tendinous fibers on 43 sides (82.7%).
Anomalies et al. 1988; Sakamoto 2016b). It may insert exclusively

N/A into the anterior atlanto-occipital ligament (Macalister
1875; Rickenbacher et al. 1985; Bergman et al. 1988;
Clinical Implications Sakamoto 2016b). It may receive a slip from the sec-
N/A ond cervical vertebra (Macalister 1875). It may connect
with longus capitis (Macalister 1875). Rectus capitis
Rectus capitis anterior (Figure 2.13) anterior may be doubled (Macalister 1875; Knott 1883a;
Rickenbacher et al. 1985). There may be a separate bundle
Synonyms along its medial border, rectus capitis minimus, that orig-
This muscle may also be referred to as rectus capitis anticus inates from the anterior arch of the first cervical vertebra
minor (Macalister 1875). (Macalister 1875; Rickenbacher et al. 1985; Bergman et
al. 1988; Sakamoto 2016b). If the origin of this bundle is
Typical Presentation shifted inferiorly over the lateral atlantoaxial joint, it may
be referred to as rectus capitis anterior medius (Eisler)
Description (Rickenbacher et al. 1985).
Rectus capitis anterior originates on the first cervical ver-
tebra from the lateral mass and base of the transverse pro-
cess (Standring 2016). It inserts into the basilar part of the Prevalence
occipital bone (Standring 2016). Rickenbacher et al. (1985) state that rectus capitis anterior
is reduced or absent in 4% of cases and rectus capitis ante-
Innervation rior medius is present in 10%–12% of cases.
Rectus capitis anterior is innervated by the ventral rami of
the first and second cervical spinal nerves (Standring 2016). Anomalies
N/A
Comparative Anatomy
Rectus capitis anterior has a similar typical presentation in Clinical Implications
apes, extending from the first cervical vertebra to the basi- N/A
lar part of the occipital bone (Sonntag 1923, 1924; Raven
1950; Dean 1985; Gibbs 1999; Diogo et al. 2010, 2012,
2013a,b, 2017). Rectus capitis lateralis (Figure 2.13)
Synonyms
Variations
N/A
Description
Rectus capitis anterior may be reduced or absent
(Macalister 1875; Rickenbacher et al. 1985; Bergman
FIGURE 2.13 Anterior vertebral muscles in anterior view.

Typical Presentation Rectus capitis posterior major (Figure 2.14)

Rectus capitis lateralis originates from the transverse pro- This muscle may also be referred to as rectus capitis posticus
cess of the first cervical vertebra and inserts into the jugular major or rectus capitis posterior superficialis (Macalister
process of the occipital bone (Standring 2016). 1875; Bakkum and Miller 2016).
Innervation Typical Presentation

Rectus capitis lateralis is innervated by the ventral rami of Description
the first and second cervical spinal nerves (Standring 2016). Rectus capitis posterior major originates from the spinous
process of the second cervical vertebra (Standring 2016). It
Comparative Anatomy inserts onto the occipital bone along the lateral half of the
Rectus capitis lateralis has a similar typical presentation inferior nuchal line and onto the bone just below the line
in apes, inserting from the transverse process of the first (Standring 2016).
cervical vertebra to the jugular process of the occipital
Innervation
bone (Sonntag 1923; Raven 1950; Dean 1984; Gibbs 1999;
Diogo et al. 2010, 2012, 2013a,b, 2017). Rectus capitis Rectus capitis posterior major is innervated by the suboc-
lateralis may be absent in some orangutans (Gibbs 1999; cipital nerve (Standring 2016).
Sonntag 1924).
Comparative Anatomy
Rectus capitis posterior major has a similar typical presen-
Variations tation in the apes, extending from the spinous process of
Description the second cervical vertebra to the occipital bone (Sonntag
Rectus capitis lateralis may be deficient (Macalister 1923, 1924; Raven 1950; Miller 1952; Gibbs 1999; Diogo
1875). The muscle may be fan-shaped (Macalister 1875; et al. 2010, 2012, 2013a,b, 2017). Sonntag (1923) notes
Rickenbacher et al. 1985). Levator scapulae may fuse that the rectus capitis posterior major muscles mostly con-
with the inferior attachment of rectus capitis lateralis ceal the rectus capitis posterior minor muscles in common
(Gonzales et al. 2017). Rectus capitis lateralis may be chimpanzees. The rectus capitis posterior minor muscles
doubled (Macalister 1875; Rickenbacher et al. 1985). are concealed to a lesser extent in orangutans (Sonntag
The accessory belly may be referred to as rectus capitis 1924). Grider-Potter (2017) found an accessory suboccipital
lateralis accessorius (Winslow) (Macalister 1875; Knott muscle in one gibbon. It had the same origin and insertion
1883a). A rectus capitis lateralis longus (Otto) may be as rectus capitis posterior major but was situated lateral and
present, extending from the transverse process of the sec- deep to that muscle and deep to obliquus capitis superior.
ond cervical vertebra to the occipital bone (Macalister This location is similar to that of the rectus capitis posterior
1875; Rickenbacher et al. 1985; Bergman et al. 1988; medius muscle found in cats.
Sakamoto 2016b).
Variations
Prevalence Description
Knott (1883a) found rectus capitis lateralis accessorius in 3 Rectus capitis posterior major may contact its counterpart
out of 33 subjects (9.1%). at the midline such that the rectus capitis posterior minor
muscles are not visible (Mori 1964). Rectus capitis pos-
Anomalies terior major may receive accessory slips from the spine
of the third cervical vertebra or the ligamentum nuchae
Description (Rickenbacher et al. 1985). It may also receive slips from
Rectus capitis lateralis may be absent in individuals with the spinalis or semispinalis (Macalister 1875), or from rec-
atlas assimilation, or in individuals with a well-developed tus capitis posterior minor or obliquus capitis inferior (Mori
paracondylar process that articulates with the transverse 1964). There may be a soft-tissue connection (myodural
process of the atlas (Rickenbacher et al. 1985). bridge) between rectus capitis posterior major and the dura
mater in the atlanto-axial interspace (Scali et al. 2011, 2013;
Prevalence Bakkum and Miller 2016).
N/A Rectus capitis posterior major may be doubled (Wood
1867b; Macalister 1875; Mori 1964; Rickenbacher et al.
Clinical Implications 1985; Tagil et al. 2005; Nayak et al. 2011; Bakkum and
Fusion of levator scapulae with the origin of rectus capi- Miller 2016). When doubled, the more lateral belly may
tis lateralis may cause greater flexion of the craniocervical extend to near rectus capitis lateralis (Rickenbacher et al.
junction (Gonzales et al. 2017). 1985). Loukas and Tubbs (2007) describe an accessory
FIGURE 2.14 Suboccipital muscles in posterior view.
suboccipital muscle that originated from the spinous pro- minor bilaterally in three cadavers (2.7%), on the right
cess of the second cervical vertebra inferior to rectus capi- side only in two cadavers (1.8%), and on the left side only
tis posterior major and inserted into the inferior nuchal in nine cadavers (8%). The left rectus capitis posterior
line lateral and posterior to the insertion of rectus capitis major muscle received a slip from obliquus capitis infe-
posterior major. rior in one cadaver (0.9%).
Prevalence Anomalies
In a sample of 13 cadavers, 11 cadavers (84.6%) had con-
Description
nections between rectus capitis posterior major and the
dura mater in the atlanto-axial interspace (Scali et al. When the spinous process of the second cervical vertebra
2011). In a sample of 112 cadavers, Mori (1964) found is absent, rectus capitis posterior major may originate from
rectus capitis posterior major with a typical presentation the spinous process of the third cervical vertebra (Asakawa
bilaterally in 84 cadavers (75%), on the right side only et al. 1999). In an individual with atlas assimilation, the
in seven cadavers (6.3%), and on the left side only in six right rectus capitis posterior major muscle was comprised
cadavers (5.4%). The muscle was longitudinally split into of two fan-shaped bellies, and the left muscle was reduced
two parts bilaterally in three cadavers (2.7%), on the right in size (Ciołkowski et al. 2014).
side only in one cadaver (0.9%), and on the left side only
in two cadavers (1.8%). Rectus capitis posterior major Prevalence
received accessory slips from rectus capitis posterior N/A
Clinical Implications et al. 2016). Yuan et al. (2016) also found that rectus capitis
The myodural communication between rectus capitis pos- posterior minor may connect to the dura via the posterior
terior major and the dura mater may monitor dural tension atlanto-axial interspace.
or be involved in some cervicogenic pathologies (Scali et al. Rectus capitis posterior minor may be doubled
2011, 2013). A doubled rectus capitis posterior major may (Macalister 1875; Mori 1964; Rickenbacher et al. 1985;
strain the spine of the second cervical vertebra or contribute Tagil et al. 2005; Bakkum and Miller 2016; Standring
to cervicogenic headaches (Nayak et al. 2011). Accessory 2016). It may also be divided into three parts (Mori 1964;
muscles like the one described by Loukas and Tubbs (2007) Bakkum and Miller 2016). When doubled or divided, the
may compress the vertebral artery. second muscle is often smaller and situated lateral to the
normal muscle and deep to rectus capitis posterior major
(Mori 1964; Rickenbacher et al. 1985).
rectus capitis pOsteriOr MinOr (figure 2.14)
Synonyms Prevalence
This muscle may also be referred to as rectus capitis posti- In a sample of 112 cadavers, Mori (1964) found rectus
cus minor, rectus posticus minor, or rectus capitis posterior capitis posterior minor with a typical presentation bilat-
profundus (Macalister 1875; Bakkum and Miller 2016). erally in 54 cadavers (48.2%), on the right side only in 16
cadavers (14.3%), and on the left side only in ten cadavers
(8.9%). Rectus capitis posterior minor was absent bilater-
ally in two cadavers (1.8%), on the right side only in one
Description cadaver (0.9%), and on the left side only in three cadav-
Rectus capitis posterior minor originates from the posterior ers (2.7%). The muscle was longitudinally split into two
tubercle of the frst cervical vertebra and inserts onto the parts bilaterally in 17 cadavers (15.2%), on the right side
occipital bone along the medial half of the inferior nuchal only in 11 cadavers (9.8%), and on the left side only in 12
line and onto the bone just below the line (Standring 2016). cadavers (10.7%). Rectus capitis posterior minor received
an accessory slip that originated from the spine of the
Innervation second cervical vertebra or the ligamentum nuchae bilat-
Rectus capitis posterior minor is innervated by the suboc- erally in seven cadavers (6.3%) and on the left side only
cipital nerve (Standring 2016). in one cadaver (0.9%). Rectus capitis posterior minor sent
accessory slips to rectus capitis posterior major bilater-
Comparative Anatomy ally in three cadavers (2.7%), on the right side only in
Rectus capitis posterior minor has a similar typical pre- two cadavers (1.8%), and on the left side only in nine
sentation in the apes, extending from the posterior tubercle cadavers (8%).
of the frst cervical vertebra to the occipital bone (Sonntag In a sample of 75 cadavers, Zumpano et al. (2006) found
1923, 1924; Raven 1950; Miller 1952; Gibbs 1999; Diogo that rectus capitis posterior minor was absent bilaterally in
et al. 2010, 2012, 2013a,b, 2017). Sonntag (1923) notes that fve cadavers (6.7%). A soft tissue bridge between rectus
the rectus capitis posterior major muscles mostly conceal capitis posterior minor and the posterior atlanto-occipi-
the rectus capitis posterior minor muscles in common chim- tal membrane was found on the right side in 53 cadavers
panzees. The rectus capitis posterior minor muscles are (70.7%) and on the left side in 55 cadavers (73.3%). Out of
concealed to a lesser extent in orangutans (Sonntag 1924). 11 cases, Hack et al. (1995) found a connection between
rectus capitis posterior minor and the dorsal spinal dura
Variations at the atlanto-occipital junction in all cases (100%). In a
Description sample of 13 cadavers, all of them (100%) had connection
Rectus capitis posterior minor may be reduced or absent between rectus capitis posterior minor and the dura mater
(Mori 1964; Rickenbacher et al. 1985; Zumpano et al. 2006; (Scali et al. 2011).
Nayak et al. 2011). It may be separated from its counterpart Yuan et al. (2016) studied the origin of rectus capitis
at the midline (Mori 1964). Rectus capitis posterior minor posterior minor in 14 specimens using P45 sheet plastina-
may receive accessory slips from the second cervical ver- tion and in fve cadavers via dissection. Out of the sample
tebra or the ligamentum nuchae (Mori 1964; Rickenbacher of 14 specimens, the tendon of rectus capitis posterior
et al. 1985). It may send a slip to rectus capitis posterior minor attached only to the posterior arch of the atlas in
major (Mori 1964). It may fuse with spinalis capitis at its one case (type I, 7.1%). The tendon terminated in both
insertion (Martin 1994; Bakkum and Miller 2016). Rectus the posterior arch of the atlas and the posterior atlanto-
capitis posterior minor often has a soft-tissue connection occipital interspace in six cases (type II, 42.9%) and
to the posterior atlanto-occipital membrane and the dura in both the posterior arch of the atlas and the posterior
mater in the atlanto-occipital interspace (Hack et al. 1995; atlanto-axial interspace in one case (type III, 7.1%). The
Hack and Hallgren 2004; Zumpano et al. 2006; Scali tendon terminated in all three locations in six cases (type
et al. 2011; Bakkum and Miller 2016; Standring 2016; Yuan IV, 42.9%). Among the fve cadavers, a type I attachment
was found in two cases (40%), a type II attachment was Variations

found in one case (20%), and a type III attachment was Description
found in two cases (40%).
Obliquus capitis superior may be bilaminar (Flower and
Murie 1867; Macalister 1875; Rickenbacher et al. 1985). Its
Anomalies origin may receive bundles from obliquus capitis inferior
Description (Rickenbacher et al. 1985). It is associated with atlantomas-
When the posterior tubercle of the frst cervical vertebra is toideus, which extends from the transverse process of the
absent, rectus capitis posterior minor may originate from an frst cervical vertebra to the mastoid process (see the entry
unusual prominence on the spinous process of the second for this muscle).
cervical vertebra (Brown 1941). In an individual with atlas
assimilation, the right rectus capitis posterior minor muscle Prevalence
was vestigial, and the left muscle was replaced by a band of
N/A
connective tissue (Ciołkowski et al. 2014).
N/A Description
In a fetus with trisomy 13, the left obliquus capitis superior
had an extra belly (Pettersen et al. 1979). In an individual
The absence of rectus capitis posterior minor may lead with atlas assimilation, the left obliquus capitis superior
to a lack of coordination among the suboccipital muscles muscle was replaced by a bundle of connective tissue
while balancing the head or contribute to cervicogenic (Ciołkowski et al. 2014).
headaches (Nayak et al. 2011). The myodural commu-
nication between rectus capitis posterior minor and the
dorsal spinal dura may help resist dural infolding (Hack Prevalence
et al. 1995). Surgical separation of this myodural connec- In their literature review, Smith et al. (2015) found that an
tion can provide relief from chronic headache (Hack and extra belly of obliquus capitis superior was present in only
Hallgren 2004). The rectus capitis posterior minor myo- 1 out of 20 individuals with trisomy 13 (5%).
dural bridge may infuence the circulation of cerebrospi-
nal fuid (Yuan et al. 2016). Clinical Implications
N/A
Obliquus capitis superiOr (figure 2.14)
See also: Atlantomastoideus
Obliquus capitis inferiOr (figure 2.14)
Synonyms
This muscle may be referred to as obliquus superior or Synonyms
obliquus capitis major (Macalister 1875; Bakkum and This muscle may be referred to as obliquus inferior or
Miller 2016). obliquus capitis minimus (Macalister 1875; Bakkum and
Miller 2016).
Obliquus capitis superior originates from the transverse Obliquus capitis inferior originates from the spinous process
process of the frst cervical vertebra and inserts onto the of the second cervical vertebra and inserts onto the trans-
occipital bone between the inferior and superior nuchal verse process of the frst cervical vertebra (Standring 2016).
lines (Standring 2016).
Innervation
Innervation Obliquus capitis inferior is innervated by the suboccipital
Obliquus capitis superior is innervated by the suboccipital nerve (Standring 2016).
nerve (Standring 2016).
Comparative Anatomy
Comparative Anatomy Obliquus capitis inferior has a similar typical presenta-
Obliquus capitis superior has a similar typical presentation tion in the apes, extending from the spinous process of
in the apes, extending from the transverse process of the the second cervical vertebra to the transverse process of
frst cervical vertebra to the occipital bone (Sonntag 1923, the frst cervical vertebra (Sonntag 1923, 1924; Raven
1924; Raven 1950; Miller 1952; Gibbs 1999; Diogo et al. 1950; Miller 1952; Gibbs 1999; Diogo et al. 2010, 2012,
2010, 2012, 2013a,b, 2017). 2013a,b, 2017).
Variations (lateral) head originates from the medial third of the clavicle
Description (Standring 2016). The two heads join and insert into the mas-
toid process and the superior nuchal line (Standring 2016).
Obliquus capitis inferior may overlap rectus capitis posterior
major (Macalister 1875). It may send a slip to rectus capi-
tis posterior major (Mori 1964) or bundles to the origin of Innervation
obliquus capitis superior (Rickenbacher et al. 1985). It may Sternocleidomastoid is innervated by the accessory nerve
also send bundles to the transverse processes of the second (Standring 2016).
or third cervical vertebrae (Rickenbacher et al. 1985). There
may be a myodural bridge connecting obliquus capitis infe- Comparative Anatomy
rior with the dura mater (Scali et al. 2011; Pontell et al. 2013; Sternocleidomastoid has a similar typical presentation
Bakkum and Miller 2016). Obliquus capitis inferior may be in the apes, with a sternomastoid head that extends from
doubled (Macalister 1875; Rickenbacher et al. 1985). the sternum to the mastoid and nuchal crest and a cleido-
mastoid head that extends from the medial clavicle to the
Prevalence mastoid (Gratiolet and Alix 1866; Primrose 1899, 1900;
In a sample of 112 cadavers, Mori (1964) found that the left Sonntag 1923, 1924; Miller 1952; Gibbs 1999; Diogo et al.
rectus capitis posterior major muscle received a slip from 2010, 2012, 2013a,b, 2017). Accessory heads may be found
obliquus capitis inferior in one cadaver (0.9%). In a sample in gibbons or bonobos (Schück 1913; Miller 1952). Schück
of 14 obliquus capitis inferior muscles, Pontell et al. (2013) (1913) found that the muscle may fuse with trapezius in
found that all of the muscles (100%) were connected with gibbons. In bonobos, the clavicular attachment may be
the dura mater. absent (Diogo et al. 2017). An insertion of the cleidomas-
toid head into the axis has been observed in orangutans
(Owen 1868; Diogo et al. 2013b) while an insertion into the
Anomalies atlas has been observed in common chimpanzees (Sutton
Description 1883; Diogo et al. 2013a).
On the right side of one male neonate with trisomy 13,
obliquus capitis inferior sent an accessory slip to semispina- Variations
lis cervicis (Pettersen et al. 1979). On the left side, obliquus Description
capitis inferior had an extra belly. In an individual with
The sternal head (sternomastoid) and the clavicular head
atlas assimilation, the left obliquus capitis inferior muscle
(cleidomastoid) may be completely separated (Hallett 1848;
was thinner than the right muscle (Ciołkowski et al. 2014).
Macalister 1875; Knott 1880, 1883a; Mori 1964; Lee and
Prevalence Yang 2016). They may also be completely fused (Hallett
1848; Macalister 1875). The sternal head (Macalister 1875)
In their literature review, Smith et al. (2015) found that acces-
or the entire sternocleidomastoid muscle may be absent
sory muscles associated with obliquus capitis inferior were
(McKinley and Hamilton 1976; Vajramani et al. 2010). The
found only in 1 out of 20 individuals with trisomy 13 (5%).
sternal head may be doubled (Macalister 1875; Nayak et al.
Clinical Implications 2006; Raikos et al. 2012). The clavicular origin may be
doubled or tripled (Wood 1868; Macalister 1875; Ramesh
The obliquus capitis inferior myodural bridge may be
Rao et al. 2007; Raikos et al. 2012; Heo et al. 2020). The
involved in monitoring dural tension or resisting dural
clavicular head may also be divided into multiple fascicles,
infolding (Pontell et al. 2013).
or into superfcial and deep sheets (Macalister 1875; Knott
1880, 1883a; Mori 1964). There are reports of sternoclei-
POSTEROLATERAL NECK MUSCLES domastoid with six heads, comprised of a doubled sternal
head and quadrupled clavicular head (Natsis et al. 2009;
sternOcleiDOMastOiDeus (sternOcleiDOMastOiD) Kaur et al. 2013). Surendran et al. (2016) report a sterno-
(figures 2.3 anD 3.3) cleidomastoid with 13 tendons of origin and fve feshy
bellies (four clavicular bellies, one sternal belly). The inser-
See also: Cleido-occipitalis cervicalis, Platysma cervicale, tion of sternocleidomastoid may also be doubled or divided
Levator claviculae, Supraclavicularis proprius, Sternalis (Macalister 1875; Mori 1964). The entire sternocleidomas-
toid muscle may be doubled (Lee and Yang 2016).
Synonyms
The extent of the origin along the clavicle may vary
N/A (Hallett 1848; Macalister 1875; Lee and Yang 2016). The
clavicular head may be narrow and tendinous (Macalister
1875). Sato (1968b) found that sternocleidomastoid
Description formed a thick band anteriorly and a thin band posteri-
Sternocleidomastoid has two heads. The sternal (medial) orly. Sternocleidomastoid may blend with the clavicular
head originates from the manubrium, and the clavicular attachment of trapezius (Hallett 1848; Macalister 1875;
Rickenbacher et al. 1985; Bakkum and Miller 2016; to platysma. In a neonate with trisomy 18, the left sternoclei-
Standring 2016). It may send accessory slips to the mas- domastoid was divided into two distinct parts (Aziz 1979).
toid process, tympanic ring, parotid fascia, stylomandibular In a fetus with trisomy 18 and cyclopia, sternocleidomastoid
ligament, thyroid cartilage, hyoid, angle of the mandible, or was split into fascicles; had a broad attachment to the skull, a
clavicle on the opposite side of the body (Macalister 1875; strong fascial connection to platysma; and formed a complex
Lee and Yang 2016). with rhomboideus occipitalis and trapezius (Smith et al. 2015).
Sternocleidomastoid is associated with multiple named In a fetus with craniorachischisis, the left sternocleidomastoid
accessory muscles. Coskun et al. (2002) report a case in had a small clavicular head and a large sternal head, attached
which sternomastoid and cleidomastoid were accompa- to the “temporo-occipital bone,” and fused with trapezius
nied by a “sternocleidooccipital” muscle, which had a (Alghamdi et al. 2017). The right sternocleidomastoid had
single origin from the occiput and a split attachment onto three heads: a lateral head attached to the “temporo-occipital
the clavicle and the manubrium. This accessory muscle bone,” a middle head attached to the skull inferior to the ear,
sent an additional bundle to sternomastoid. Platysma cer- and a small head attached to the mandibular ramus.
vicale (transversus nuchae, occipital platysma) runs from
the occipital region to the mouth or posterior ear region Prevalence
and may insert into the anterior border of sternocleido-
In their literature review, Smith et al. (2015) found that ster-
mastoid (see the entry for this muscle). Levator claviculae
nocleidomastoid was divided into three bellies in 1 out of 20
originates from the transverse processes of some cervical
individuals with trisomy 13 (5%). The muscle was divided
vertebrae and inserts onto the clavicle, sometimes insert-
into sternomastoid and cleidomastoid portions in 2 out of 17
ing into sternocleidomastoid (see the entry for this muscle).
individuals with trisomy 18 (11.8%).
Supraclavicularis proprius originates from the cervical fas-
cia overlying the clavicular head of sternocleidomastoid and Clinical Implications
the sternoclavicular joint and inserts onto the distal end of
The absence of sternocleidomastoid may be associated with
the clavicle or the fascial sheath of trapezius (see the entry
torticollis (McKinley and Hamilton 1976; Vajramani et al.
for this muscle). Sternalis, a variable accessory muscle that
2010). Accessory heads may lead to stenosis of the supra-
extends from the sternal/infraclavicular area to the upper
clavicular fossa (Natsis et al. 2009; Raikos et al. 2012; Lee
abdominal wall or costal cartilages, may originate from
and Yang 2016).
or blend with sternocleidomastoid (see the entry for this
muscle). Cleido-occipitalis cervicalis is situated near the
posterior border of sternocleidomastoid, extending between
the occiput and the clavicle (see the entry for this muscle). cleiDO-Occipitalis cervicalis (figure 2.3)
See also: Sternocleidomastoideus, Trapezius
Prevalence
Hallett (1848) found a lengthened attachment of the cla- Synonyms
vicular head toward the middle of the clavicle in one out of This muscle may be referred to as the cleido-occipital
eight cases (12.5%). Wood (1868) found that sternocleido- muscle (Wood 1866, 1867b, 1870; Macalister 1875),
mastoid had a doubled or divided clavicular origin in 7 out cleido-occipitalis (Rickenbacher et al. 1985; Rahman and
of 36 subjects (19.4%). Mori (1964) found complete separa- Yamadori 1994), or accessory cleido-occipital muscle of
tion of the sternal and clavicular heads in 450 out of 510 trapezius (Paraskevas et al. 2013; note: these authors con-
cases (88.2%). Mori (1964) also found sternocleidomastoid sider the cleido-occipital muscle of Wood to be separate
separated into superfcial and deep layers in 10 out of 1020 muscle that is a variant of sternocleidomastoid).
sides (1%). The occipital insertion was separated from the
rest of the muscle in 102 out of 1020 sides (10%). The vari- Typical Presentation
ant described by Sato (1968b, see above) was present on 12 This muscle is present only as a variation or anomaly.
out of 354 sides (3.39%) in Kyushu-Japanese males and on
10 out of 216 sides (4.63%) in females. Comparative Anatomy
A cleido-occipital head of sternocleidomastoid (but not
trapezius) has been observed in a few orangutans and com-
Anomalies
mon chimpanzees (Sonntag 1923, 1924; Miller 1952; Diogo
Description et al. 2013a,b).
Sternocleidomastoid was absent on the right side in a
male infant with triploidy studied by Moen et al. (1984).
Variations
Sternocleidomastoid had three bellies on both sides of an
infant with trisomy 13 (Pettersen et al. 1979). The lateral Description
belly attached to the clavicle, the medial belly attached to the When present, cleido-occipitalis cervicalis (Kwak et al.
manubrium, and the middle belly attached to the sternocla- 2003) presents as a separate slip that courses along the ante-
vicular junction. On the left side, the medial belly sent slips rior border of trapezius, originating from the occiput and
inserting into the clavicle (Hallett 1848; Wood 1866, 1867b, Trapezius (Figures 2.3 and 3.3)
1870; Macalister 1867b, 1875; Mori 1964; Rickenbacher
et al. 1985; Rahman and Yamadori 1994; Kwak et al. 2003; See also: Cleido-occipitalis cervicalis, Platysma cervicale,
Ravindra et al. 2012; Paraskevas et al. 2013; Bakkum and Levator claviculae, Supraclavicularis proprius
Miller 2016). It likely forms as an isolated muscle during Synonyms
the separation of trapezius and sternocleidomastoid from
This muscle may also be referred to as cucullaris (Henle)
the common muscle anlage (Rahman and Yamadori 1994;
(Bakkum and Miller 2016).
Kwak et al. 2003; Paraskevas et al. 2013; Bakkum and
Miller 2016). Typical Presentation
It may join with trapezius or sternocleidomastoid at its
origin or insertion (Wood 1867b, 1870; Macalister 1875; Description
Rahman and Yamadori 1994; Kwak et al. 2003). It may be Trapezius originates from the superior nuchal line, external
split into separate fascicles (Wood 1870; Macalister 1875). occipital protuberance, nuchal ligament, and the spinous
It may be doubled (Wood 1870; Macalister 1875). It may processes and supraspinous ligaments of the seventh cervical
be comprised of tendinous or muscular fibers (Mori 1964). through the twelfth thoracic vertebrae (Standring 2016). It
The external jugular vein may pass through the space cre- inserts onto the lateral third of the clavicle, the acromion, and
ated between cleido-occipitalis cervicalis and the clavicle the scapular spine (Standring 2016). The muscle can there-
(Mori 1964; Ravindra et al. 2012). Paraskevas et al. (2013) fore be divided into three parts: the superior part (descend-
observed a fibrous arch at the clavicular insertion of cleido- ing fibers with occipital and clavicular attachments), middle
occipitalis, through which the main trunk of the supracla- part (transverse fibers with vertebral, acromial, and scapular
vicular nerves passed. Rickenbacher et al. (1985) report spine attachments), and inferior part (ascending fibers with
that cleido-occipitalis cervicalis may take the form of vertebral and scapular spine attachments) (Rickenbacher
cleido-occipitalis totalis, which connects trapezius to ster- et al. 1985; Bakkum and Miller 2016; Standring 2016).
nocleidomastoid, filling the posterior triangle of the neck.
Innervation
Trapezius is innervated by the accessory nerve (Standring
Innervation 2016).
The cleido-occipitalis cervicalis is likely innervated by the
accessory nerve. Comparative Anatomy
Trapezius has a similar typical presentation in the great apes,
Prevalence differentiated into three parts with origins from the cranium
and vertebral column and insertions into the acromion, scap-
Wood (1867b) found the cleido-occipital muscle in 12
ular spine, and lateral third of the clavicle (Diogo et. 2010,
out of 34 subjects (35.3%). Wood (1870) states that the
2013a,b, 2017). As in humans, the extent of the attachment
muscle was found in 37 out of 102 subjects (36.3%). Mori
along the clavicle may vary (Hepburn 1892; Diogo et. 2010,
(1964) found a bundle along the anterior margin of tra-
2013a,b). The clavicular attachment was reported absent in
pezius that inserted into the clavicle bilaterally in 16 out
one orangutan (Schück 1913). The lowest attachment of tra-
of 130 cadavers (12.3%) and unilaterally in 11 cadavers
pezius in common chimpanzees varies between the eighth
(8.5%).
and thirteenth thoracic vertebrae (Gratiolet and Alix 1866;
Macalister 1871; Sonntag 1923; Stewart 1936; Swindler and
Anomalies Wood 1973). The lowest attachment of trapezius in bonobos
Description is either onto the ninth or tenth thoracic vertebrae (Miller
In an infant with median cleft lip, hypotelorism, and alobar 1952; Diogo et al. 2017). The occipital origin is absent in
holoprosencephaly, Mieden (1982) found cleido-occipitalis most gibbons (Deniker 1885; Kohlbrügge 1890–1892; Gibbs
present on the right side, extending between the superior 1999; Michilsens et al. 2009; Diogo et al. 2012).
nuchal line and the clavicle.
Variations
Mieden (1982) found cleido-occipitalis in one out of three Trapezius may be partially or entirely absent (Hallett 1848;
specimens with alobar holoprosencephaly (33%). Wood 1864; Macalister 1875; Testut 1884; Le Double 1897;
Bing 1902; Rickenbacher et al. 1985; Emsley and Davis 2001;
Clinical Implications Garbelotti et al. 2001; Allouh et al. 2004; Vajramani et al. 2010;
The external jugular vein may become entrapped between Bakkum and Miller 2016; Standring 2016). The occipital attach-
cleido-occipitalis cervicalis and trapezius (Ravindra et al. ment may be missing (Hallett 1848; Macalister 1875; Testut
2012). The supraclavicular nerve trunk may be entrapped 1884; Le Double 1897; Rickenbacher et al. 1985; Standring
by a fibrous arch at the insertion of the cleido-occipital 2016). The inferior part may be reduced or absent (Hallett
muscle (Paraskevas et al. 2013). 1848; Wood 1867b; Macalister 1875; Mori 1964; Rickenbacher
et al. 1985; Bakkum and Miller 2016; Standring 2016). Isolated trapezius, extending between the occiput and the clavicle
absence of the middle part can also occur (Macalister 1875; (see the entry for this muscle).
Rickenbacher et al. 1985). The clavicular insertion may also be
reduced or absent (Hallett 1848; Macalister 1875; Rickenbacher Prevalence
et al. 1985). Reduction or absence of the scapular insertion is In a review of 186 cases of congenital muscle absences,
rare (Macalister 1875; Rickenbacher et al. 1985). Due to these Bing (1902) reported that trapezius was absent in 18 cases.
variations, there may be asymmetry in development of the left Macalister (1875) states that Wood found that the inferior
and right muscles (Macalister 1875; Hallett 1848; Wood 1867b; part of trapezius extended only to the eighth and ninth,
Mori 1964; Rickenbacher et al. 1985; Emsley and Davis 2001; or tenth, thoracic vertebrae in about 4 out of 70 subjects
Bakkum and Miller 2016). (5.7%). In a sample of 216 sides, Mori (1964) found that the
Hallett (1848) reports a case in which trapezius was par- lower limit of trapezius was the eighth thoracic vertebra on
tially absent on the left side, with muscle fbers ending at the two sides (0.9%), the ninth thoracic vertebra on seven sides
level of the third thoracic vertebra and a thin fascial layer (3.2%), the tenth thoracic vertebra on 49 sides (22.7%), the
replacing the inferior part of the muscle. Similarly, Garbelotti eleventh thoracic vertebra on 79 sides (36.6%), the twelfth
et al. (2001) and Valtanen et al. (2017) report cases in which thoracic vertebra on 65 sides (30%), the frst lumbar verte-
an aponeurosis completely replaced the ascending fbers and bra on 12 sides (5.6%), and the second lumbar vertebra on
partially replaced transverse fbers of left trapezius muscles. two sides (0.9%). Mori (1964) also found the cervical and
Its origin from the occiput or its insertion into the clav- occipital parts separated on 15 sides (6.9%).
icle may occur via a tendinous arch that also connects to
sternocleidomastoid (Macalister 1875; Rickenbacher et al. Anomalies
1985). The origin of the inferior part may extend to the Description
lumbar vertebrae (Mori 1964). When the occipital origin is On the right side of a fetus with cyclopia and alobar holo-
missing, it may originate from the posterior border of ster- prosencephaly, an “occipito-trapezius” muscle was present,
nocleidomastoid (Macalister 1875). The extent of the inser- extending from the superior nuchal line to the deep surface
tion along the clavicle may vary (Hallett 1848; Macalister of trapezius (Mieden 1982). In cases of triploidy, the middle
1875; Rickenbacher et al. 1985; Bakkum and Miller 2016; part of trapezius may be absent and replaced with connec-
Standring 2016). The occipital and cervical parts may be tive tissue (Moen et al. 1984). In a trisomy 18 cyclopic fetus,
divided into multiple fasciculi (Macalister 1875). The dor- trapezius had a broad origin from the skull and was part of a
sal and cervical parts of the trapezius muscles can be par- complex with rhomboideus occipitalis and sternocleidomas-
tially separated (Macalister 1875; Mori 1964; Rickenbacher toid (Smith et al. 2015). Trapezius was also closely associ-
et al. 1985; Standring 2016; Ferreli et al. 2019). A cleft may ated with splenius capitis at the midline origin on both sides
also be found separating the inferior part of the muscle (Smith et al. 2015). In a fetus with craniorachischisis, the
(Macalister 1875; Rickenbacher et al. 1985). The entire left trapezius presented as a small muscle on top of supraspi-
muscle may be partially or entirely split into superfcial and natus, with attachments to the clavicle, acromion, scapular
deep layers (Rickenbacher et al. 1985). spine, and temporo-occipital bone (Alghamdi et al. 2017).
Trapezius may blend with sternocleidomastoid at its On the right side, trapezius presented as undifferentiated tis-
clavicular attachment (Hallett 1848; Macalister 1875; sue between the right eye and scapular spine and was fused
Rickenbacher et al. 1985; Bakkum and Miller 2016; with sternocleidomastoid (Alghamdi et al. 2017). Poland
Standring 2016). It may send slips to latissimus dorsi, ster- syndrome, a rare congenital anomaly, can cause the partial
nocleidomastoid, levator claviculae, deltoideus, the fascia of or complete absence of trapezius (Debeer et al. 2002; Yiyit
the neck, the lower angle or medial margin of the scapula, or et al. 2014). Bilateral absence of trapezius was observed in
the sternum (Wood 1867b; Macalister 1875; Rickenbacher male fetus with CHARGE syndrome (Beger et al. 2019).
et al. 1985; Bakkum and Miller 2016).
Trapezius is also associated with multiple named acces- Prevalence
sory muscles. Platysma cervicale (transversus nuchae, Moen et al. (1984) found that the middle fbers of trape-
occipital platysma) may extend from near the occipital zius were absent bilaterally and replaced with a sheet of
attachment of trapezius to the region of auricularis poste- connective tissue in three specimens with triploidy (100%).
rior or to the mouth near the parotid fascia (see the entry Mieden (1982) found occipito-trapezius in one out of three
for this muscle). Levator claviculae (cleidocervical muscle) specimens with alobar holoprosencephaly (33%).
originates from the transverse processes of some cervi-
cal vertebrae and inserts onto the clavicle near trapezius Clinical Implications
(see the entry for this muscle). Supraclavicularis proprius The absence of trapezius may be associated with torticol-
originates from the cervical fascia overlying the clavicular lis (Vajramani et al. 2010). Stevenson et al. (2014) describe a
head of sternocleidomastoid and the sternoclavicular joint cadaver with a severe case of congenital scoliosis and found
and inserts onto the distal end of the clavicle or the fascial signifcant asymmetry in the trapezius muscles. Valtanen et al.
sheath of trapezius (see the entry for this muscle). Cleido- (2017) report a partial unilateral absence of a left trapezius asso-
occipitalis cervicalis courses along the anterior margin of ciated with 5-degree thoracic scoliosis toward the right side.
3 Upper Limb Muscles
Eve K. Boyle
Vondel S. E. Mahon
Rui Diogo
CONTENTS
Axial Pectoral Girdle Muscles............................................................................................................................................ 121
Serratus anterior............................................................................................................................................................. 121
Synonyms.................................................................................................................................................................. 121
Typical Presentation.................................................................................................................................................. 121
Comparative Anatomy............................................................................................................................................... 121
Variations................................................................................................................................................................... 121
Anomalies.................................................................................................................................................................. 122
Clinical Implications................................................................................................................................................. 122
Rhomboideus major (Rhomboid major)........................................................................................................................ 122
Synonyms.................................................................................................................................................................. 122
Variations................................................................................................................................................................... 123
Anomalies.................................................................................................................................................................. 123
Rhomboideus minor (Rhomboid minor)........................................................................................................................ 123
Synonyms.................................................................................................................................................................. 123
Variations................................................................................................................................................................... 123
Anomalies.................................................................................................................................................................. 124
Rhomboideus occipitalis................................................................................................................................................ 124
Synonyms.................................................................................................................................................................. 124
Variations................................................................................................................................................................... 124
Anomalies.................................................................................................................................................................. 125
Rhomboideus tertius (Not Illustrated)............................................................................................................................ 125
Synonyms.................................................................................................................................................................. 125
Variations................................................................................................................................................................... 125
Anomalies.................................................................................................................................................................. 125
Levator scapulae............................................................................................................................................................. 125
Synonyms.................................................................................................................................................................. 125
DOI: 10.1201/9781003083535-3 111

Variations.................................................................................................................................................................. 126
Anomalies................................................................................................................................................................. 126
Subclavius ..................................................................................................................................................................... 126
Synonyms ................................................................................................................................................................. 126
Typical Presentation ................................................................................................................................................. 126
Variations.................................................................................................................................................................. 127
Anomalies................................................................................................................................................................. 127
Pectoralis intermedius ................................................................................................................................................... 127
Synonyms ................................................................................................................................................................. 127
Variations.................................................................................................................................................................. 128
Anomalies................................................................................................................................................................. 128
Subclavius posticus ....................................................................................................................................................... 128
Synonyms ................................................................................................................................................................. 128
Variations.................................................................................................................................................................. 128
Anomalies................................................................................................................................................................. 128
Levator claviculae ......................................................................................................................................................... 128
Synonyms ................................................................................................................................................................. 128
Variations.................................................................................................................................................................. 129
Anomalies................................................................................................................................................................. 130
Sternoscapularis ............................................................................................................................................................ 130
Synonyms ................................................................................................................................................................. 130
Variations.................................................................................................................................................................. 130
Anomalies................................................................................................................................................................. 130
Supraclavicularis proprius............................................................................................................................................. 130
Synonyms ................................................................................................................................................................. 130
Variations.................................................................................................................................................................. 130
Anomalies................................................................................................................................................................. 131
Pectoral Girdle and Arm Muscles ...................................................................................................................................... 131
Sternalis......................................................................................................................................................................... 131
Synonyms ................................................................................................................................................................. 131
Variations.................................................................................................................................................................. 131
Anomalies................................................................................................................................................................. 133
Dorsoepitrochlearis ....................................................................................................................................................... 133
Synonyms ................................................................................................................................................................. 133
Upper Limb Muscles 113
Variations.................................................................................................................................................................. 133
Anomalies................................................................................................................................................................. 134
Panniculus carnosus (Not Illustrated) ........................................................................................................................... 134
Synonyms ................................................................................................................................................................. 134
Variations.................................................................................................................................................................. 134
Anomalies................................................................................................................................................................. 134
Pectoralis major............................................................................................................................................................. 135
Synonyms ................................................................................................................................................................. 135
Variations.................................................................................................................................................................. 135
Anomalies................................................................................................................................................................. 136
Pectoralis minor ............................................................................................................................................................ 136
Synonyms ................................................................................................................................................................. 136
Variations.................................................................................................................................................................. 137
Anomalies................................................................................................................................................................. 137
Chondroepitrochlearis ................................................................................................................................................... 137
Synonyms ................................................................................................................................................................. 137
Variations.................................................................................................................................................................. 137
Anomalies................................................................................................................................................................. 138
Sternoclavicularis.......................................................................................................................................................... 138
Synonyms ................................................................................................................................................................. 138
Variations.................................................................................................................................................................. 138
Anomalies................................................................................................................................................................. 139
Pectoralis quartus .......................................................................................................................................................... 139
Synonyms ................................................................................................................................................................. 139
Variations.................................................................................................................................................................. 139
Anomalies................................................................................................................................................................. 139
Pectorodorsalis .............................................................................................................................................................. 140
Synonyms ................................................................................................................................................................. 140
Variations.................................................................................................................................................................. 140
Anomalies................................................................................................................................................................. 140
Pectoralis minimus ........................................................................................................................................................ 140
Synonyms ................................................................................................................................................................. 140
Variations.................................................................................................................................................................. 140
Anomalies................................................................................................................................................................. 141
Costocoracoideus .......................................................................................................................................................... 141
Synonyms ................................................................................................................................................................. 141
Variations.................................................................................................................................................................. 142
Anomalies................................................................................................................................................................. 142
Tensor semivaginae articulationis humeroscapularis .................................................................................................... 142
Synonyms ................................................................................................................................................................. 142
Variations.................................................................................................................................................................. 142
Anomalies................................................................................................................................................................. 142
Infraclavicularis............................................................................................................................................................. 142
Synonyms ................................................................................................................................................................. 142
Variations.................................................................................................................................................................. 143
Anomalies................................................................................................................................................................. 143
Costoclavicularis ........................................................................................................................................................... 143
Synonyms ................................................................................................................................................................. 143
Variations.................................................................................................................................................................. 143
Anomalies................................................................................................................................................................. 143
Roos Bands/Muscles ..................................................................................................................................................... 143
Synonyms ................................................................................................................................................................. 143
Variations.................................................................................................................................................................. 144
Anomalies................................................................................................................................................................. 144
Deltoideus (Deltoid)...................................................................................................................................................... 144
Synonyms ................................................................................................................................................................. 144
Variations.................................................................................................................................................................. 144
Anomalies................................................................................................................................................................. 145
Infraspinatus.................................................................................................................................................................. 145
Synonyms ................................................................................................................................................................. 145
Variations.................................................................................................................................................................. 146
Anomalies................................................................................................................................................................. 146
Supraspinatus ................................................................................................................................................................ 146
Synonyms ................................................................................................................................................................. 146
Variations.................................................................................................................................................................. 146
Anomalies................................................................................................................................................................. 146

Teres minor.................................................................................................................................................................... 147
Synonyms ................................................................................................................................................................. 147
Variations.................................................................................................................................................................. 147
Anomalies................................................................................................................................................................. 147
Teres major.................................................................................................................................................................... 147
Synonyms ................................................................................................................................................................. 147
Variations.................................................................................................................................................................. 148
Anomalies................................................................................................................................................................. 148
Subscapularis................................................................................................................................................................. 148
Synonyms ................................................................................................................................................................. 148
Variations.................................................................................................................................................................. 148
Anomalies................................................................................................................................................................. 149
Subscapularis minor ...................................................................................................................................................... 149
Synonyms ................................................................................................................................................................. 149
Variations.................................................................................................................................................................. 149
Anomalies................................................................................................................................................................. 149
Latissimus dorsi ............................................................................................................................................................ 149
Synonyms ................................................................................................................................................................. 149
Variations.................................................................................................................................................................. 150
Anomalies................................................................................................................................................................. 151
Levator tendinis musculi latissimus dorsi ..................................................................................................................... 151
Synonyms ................................................................................................................................................................. 151
Variations.................................................................................................................................................................. 151
Anomalies................................................................................................................................................................. 151
Coracobrachialis............................................................................................................................................................ 151
Synonyms ................................................................................................................................................................. 151
Variations.................................................................................................................................................................. 152
Anomalies................................................................................................................................................................. 152
Coracobrachialis profundus .......................................................................................................................................... 153
Synonyms ................................................................................................................................................................. 153
Variations.................................................................................................................................................................. 153
Anomalies................................................................................................................................................................. 153
Coracobrachialis longus ................................................................................................................................................ 153

Synonyms ................................................................................................................................................................. 153
Variations.................................................................................................................................................................. 153
Anomalies................................................................................................................................................................. 154
Brachialis....................................................................................................................................................................... 154
Synonyms ................................................................................................................................................................. 154
Variations.................................................................................................................................................................. 154
Anomalies................................................................................................................................................................. 155
Biceps brachii................................................................................................................................................................ 155
Synonyms ................................................................................................................................................................. 155
Variations.................................................................................................................................................................. 156
Anomalies................................................................................................................................................................. 156
Triceps brachii............................................................................................................................................................... 157
Synonyms ................................................................................................................................................................. 157
Variations.................................................................................................................................................................. 157
Anomalies................................................................................................................................................................. 158
Epitrochleoanconeus ..................................................................................................................................................... 158
Synonyms ................................................................................................................................................................. 158
Variations.................................................................................................................................................................. 158
Anomalies................................................................................................................................................................. 159
Anterior Forearm Muscles ................................................................................................................................................. 159
Flexor pollicis longus.................................................................................................................................................... 159
Synonyms ................................................................................................................................................................. 159
Variations.................................................................................................................................................................. 160
Anomalies................................................................................................................................................................. 161
Gantzer’s Muscle........................................................................................................................................................... 161
Synonyms ................................................................................................................................................................. 161
Variations.................................................................................................................................................................. 161
Anomalies................................................................................................................................................................. 162
Flexor digitorum profundus .......................................................................................................................................... 162
Synonyms ................................................................................................................................................................. 162
Variations.................................................................................................................................................................. 162
Anomalies................................................................................................................................................................. 163
Flexor digitorum superfcialis ....................................................................................................................................... 163

Synonyms ................................................................................................................................................................. 163
Variations.................................................................................................................................................................. 164
Anomalies................................................................................................................................................................. 165
Pronator quadratus ........................................................................................................................................................ 165
Synonyms ................................................................................................................................................................. 165
Comparative Anatomy ............................................................................................................................................. 166
Variations.................................................................................................................................................................. 166
Anomalies................................................................................................................................................................. 166
Radiocarpus................................................................................................................................................................... 166
Synonyms ................................................................................................................................................................. 166
Variations.................................................................................................................................................................. 166
Anomalies................................................................................................................................................................. 167
Palmaris longus ............................................................................................................................................................. 167
Synonyms ................................................................................................................................................................. 167
Variations.................................................................................................................................................................. 167
Anomalies................................................................................................................................................................. 169
Flexor carpi ulnaris ....................................................................................................................................................... 169
Synonyms ................................................................................................................................................................. 169
Variations.................................................................................................................................................................. 170
Anomalies................................................................................................................................................................. 170
Flexor carpi radialis....................................................................................................................................................... 170
Synonyms ................................................................................................................................................................. 170
Variations.................................................................................................................................................................. 170
Anomalies................................................................................................................................................................. 171
Flexor carpi radialis brevis............................................................................................................................................ 171
Synonyms ................................................................................................................................................................. 171
Variations.................................................................................................................................................................. 171
Anomalies................................................................................................................................................................. 171
Pronator teres ................................................................................................................................................................ 171
Synonyms ................................................................................................................................................................. 171
Variations.................................................................................................................................................................. 172
Anomalies................................................................................................................................................................. 172
Posterior Forearm Muscles ................................................................................................................................................ 173
Brachioradialis .............................................................................................................................................................. 173

Synonyms ................................................................................................................................................................. 173
Variations.................................................................................................................................................................. 173
Anomalies................................................................................................................................................................. 174
Supinator ....................................................................................................................................................................... 174
Synonyms ................................................................................................................................................................. 174
Variations.................................................................................................................................................................. 175
Anomalies................................................................................................................................................................. 175
Anconeus....................................................................................................................................................................... 176
Synonyms ................................................................................................................................................................. 176
Variations.................................................................................................................................................................. 176
Anomalies................................................................................................................................................................. 176
Extensor carpi radialis longus ....................................................................................................................................... 176
Synonyms ................................................................................................................................................................. 176
Variations.................................................................................................................................................................. 176
Anomalies................................................................................................................................................................. 177
Extensor carpi radialis brevis ........................................................................................................................................ 178
Synonyms ................................................................................................................................................................. 178
Variations.................................................................................................................................................................. 178
Anomalies................................................................................................................................................................. 179
Extensor carpi radialis accessorius................................................................................................................................ 179
Synonyms ................................................................................................................................................................. 179
Variations.................................................................................................................................................................. 179
Anomalies................................................................................................................................................................. 180
Extensor carpi ulnaris.................................................................................................................................................... 180
Synonyms ................................................................................................................................................................. 180
Variations.................................................................................................................................................................. 180
Anomalies................................................................................................................................................................. 180
Ulnaris digiti quinti ....................................................................................................................................................... 180
Synonyms ................................................................................................................................................................. 180
Variations.................................................................................................................................................................. 180
Anomalies................................................................................................................................................................. 181
Extensor digitorum........................................................................................................................................................ 181
Synonyms ................................................................................................................................................................. 181

Variations.................................................................................................................................................................. 182
Anomalies................................................................................................................................................................. 183
Extensor digiti minimi................................................................................................................................................... 183
Synonyms ................................................................................................................................................................. 183
Variations.................................................................................................................................................................. 183
Anomalies................................................................................................................................................................. 184
Extensor indicis............................................................................................................................................................. 185
Synonyms ................................................................................................................................................................. 185
Variations.................................................................................................................................................................. 185
Anomalies................................................................................................................................................................. 186
Extensor medii digiti ..................................................................................................................................................... 186
Synonyms ................................................................................................................................................................. 186
Variations.................................................................................................................................................................. 186
Anomalies................................................................................................................................................................. 187
Extensor digitorum brevis manus.................................................................................................................................. 187
Synonyms ................................................................................................................................................................. 187
Variations.................................................................................................................................................................. 187
Anomalies................................................................................................................................................................. 188
Extensor pollicis longus ................................................................................................................................................ 188
Synonyms ................................................................................................................................................................. 188
Variations.................................................................................................................................................................. 188
Anomalies................................................................................................................................................................. 189
Extensor pollicis brevis ................................................................................................................................................. 189
Synonyms ................................................................................................................................................................. 189
Variations.................................................................................................................................................................. 189
Anomalies................................................................................................................................................................. 190
Abductor pollicis longus ............................................................................................................................................... 191
Synonyms ................................................................................................................................................................. 191
Variations.................................................................................................................................................................. 191
Anomalies................................................................................................................................................................. 192
Muscles of the Hand .......................................................................................................................................................... 192
Contrahentes digitorum manus ..................................................................................................................................... 192
Synonyms ................................................................................................................................................................. 192

Variations.................................................................................................................................................................. 193
Anomalies................................................................................................................................................................. 193
Palmaris brevis .............................................................................................................................................................. 193
Synonyms ................................................................................................................................................................. 193
Variations.................................................................................................................................................................. 194
Anomalies................................................................................................................................................................. 194
Lumbricales (lumbricals)...................................................................................................................................................194
Synonyms ................................................................................................................................................................. 194
Variations.................................................................................................................................................................. 195
Anomalies................................................................................................................................................................. 195
Adductor pollicis........................................................................................................................................................... 196
Synonyms ................................................................................................................................................................. 196
Variations.................................................................................................................................................................. 197
Anomalies................................................................................................................................................................. 197
Interossei palmaris (Palmar interossei)...........................................................................................................................197
Synonyms ................................................................................................................................................................. 197
Variations.................................................................................................................................................................. 198
Anomalies................................................................................................................................................................. 198
Interossei dorsalis (Dorsal interossei).............................................................................................................................. 199
Synonyms ................................................................................................................................................................. 199
Variations.................................................................................................................................................................. 199
Anomalies................................................................................................................................................................. 199
Flexor pollicis brevis.....................................................................................................................................................200
Synonyms .................................................................................................................................................................200
Variations..................................................................................................................................................................200
Anomalies.................................................................................................................................................................200
Opponens pollicis.......................................................................................................................................................... 201
Synonyms ................................................................................................................................................................. 201
Variations.................................................................................................................................................................. 201
Anomalies................................................................................................................................................................. 201
Flexor digiti minimi brevis............................................................................................................................................ 201
Synonyms ................................................................................................................................................................. 201

Variations..................................................................................................................................................................202
Anomalies.................................................................................................................................................................202
Opponens digiti minimi.................................................................................................................................................202
Synonyms .................................................................................................................................................................202
Variations..................................................................................................................................................................202
Anomalies................................................................................................................................................................. 203
Abductor pollicis brevis ................................................................................................................................................ 203
Synonyms ................................................................................................................................................................. 203
Variations.................................................................................................................................................................. 203
Anomalies................................................................................................................................................................. 203
Abductor digiti minimi..................................................................................................................................................204
Synonyms .................................................................................................................................................................204
Variations..................................................................................................................................................................204
Anomalies.................................................................................................................................................................204
AXIAL PECTORAL GIRDLE MUSCLES In a bonobo fetus, serratus anterior was fused with the
insertion of levator scapulae (Diogo et al. 2017).
serratus anteriOr (figure 3.1)
Synonyms Variations
Serratus anterior may also be referred to as serratus anti- Description
cus, serratus magnus, or serratus anticus major (Macalister The superior part of serratus anterior may be split into two
1875; Bakkum and Miller 2016). layers or it may be absent (Macalister 1875). The middle
portion of the muscle may also be absent, or it may be
Typical Presentation reduced and replaced with connective tissue (Macalister
Description 1875; Bergman et al. 1988; Bakkum and Miller 2016;
Serratus anterior originates from ribs one through eight and Standring 2016). Slips to the frst or eighth rib may be
inserts onto the medial border of the scapula (Standring absent (Macalister 1875; Mori 1964; Bergman et al. 1988;
2016). Smith et al. 2003; Standring 2016). Serratus anterior may
have origins from ribs nine and ten (Macalister 1875; Mori
Innervation 1964; Bergman et al. 1988; Bakkum and Miller 2016;
Serratus anterior is innervated by the long thoracic nerve Standring 2016). Every slip beyond the frst two may be
(Standring 2016). absent (Macalister 1875). There may be connections to
coracobrachialis, pectoralis minor, or scalenus posterior
Comparative Anatomy (Smith et al. 2003; Bakkum and Miller 2016). Serratus
Serratus anterior has a similar typical presentation in the anterior may also be continuous with the external oblique,
apes, extending from the ribs to the medial side of the intercostal muscles, supracostalis anterior, or levator scap-
scapula (Gratiolet and Alix 1866; Macalister 1871, 1873; ulae (Macalister 1875; Bergman et al. 1988; Bakkum and
Champneys 1872; Deniker 1885; Kohlbrügge 1890–1892; Miller 2016; Standring 2016).
Hepburn 1892; Beddard 1893; Primrose 1899, 1900; Schück
1913; Sonntag 1923; Sullivan and Osgood 1927; Stewart Prevalence
1936; Raven 1950; Miller 1952; Gibbs 1999; Diogo et al. In a study cited by Bergman et al. (1988) and Bakkum and
2010, 2012, 2013a,b, 2017). In the apes, serratus anterior Miller (2016), the inferior most attachment of serratus ante-
tends to have origins more often from the inferior-most rior was to rib seven in 1% of cases, rib eight in 40% of
ribs (Gibbs 1999; Diogo et al. 2010, 2012, 2013a,b, 2017). cases, rib nine in 38% of cases, rib ten in 10% of cases, and
FIGURE 3.1 Thoracoscapular muscles in posterior view.
rib 11 in 0.5% of cases. The superior most attachment of Prevalence

the muscle was to rib one in 75% of cases, to the fascia of N/A
the rib one in 21% of cases, and to rib two in 3% of cases
(Bergman et al. 1988). Based on a study of 204 sides, Mori Clinical Implications
(1964) reported that the inferior most attachment of serratus The absence of serratus anterior can cause winged scapula
anterior was to rib seven on 14 sides (6.8%), rib eight on 26 (Levin and Trummer 1973).
sides (12.7%), rib nine on 152 sides (74.5%), and rib ten on
12 sides (5.8%). The superior most attachment was to rib
one on 182 sides (89.2%) and to rib two on 22 sides (10.7%) rhOMbOiDeus MajOr (rhOMbOiD MajOr) (figure 3.1)
of cases. In a study of 13 sides, Smith et al. (2003) observed See also: Rhomboideus minor
that the superior most attachment was to both ribs one and
two on six sides (46%), to rib one only on two sides (15%), Synonyms
to rib two only on four sides (31%), and to ribs two and three Rhomboideus major may also be referred to as rhomboi-
on one side (8%). deus inferior (Bakkum and Miller 2016).
Anomalies Typical Presentation

Poland syndrome, a rare congenital anomaly, can lead to Rhomboideus major originates from the spines of the sec-
the absence of serratus anterior (Cingel et al. 2013). In a ond through ffth thoracic vertebrae (Standring 2016). It
male fetus with triploidy, the upper half of serratus anterior attaches to the medial border of the scapula inferior to the
was absent bilaterally (Moen et al. 1984). spine (Standring 2016).
Innervation of the rhomboid muscles in this specimen was about

Rhomboideus major is innervated by the dorsal scapular one-third fbrous and two-thirds tendinous (Smith et al.
nerve (Standring 2016). 2015). In a male fetus with triploidy, rhomboid major had
an extra muscle slip on the right side (Moen et al. 1984).
Comparative Anatomy In a cadaver with a severe case of congenital scoliosis,
In the apes, the rhomboid muscles are typically fused and Stevenson et al. (2014) found signifcant asymmetry in the
extend between the cervical and thoracic vertebrae and the rhomboid muscles.
scapula (Macalister 1873; Deniker 1885; Kohlbrügge 1890–
1892; Schück 1913; Sonntag 1923, 1924; Stewart 1936; Prevalence
Raven 1950; Miller 1952; Preuschoft 1965; Swindler and In their literature review, Smith et al. (2015) found that
Wood 1973; Gibbs 1999; Diogo et al. 2010, 2012, 2013a,b, rhomboideus major and minor were fused in 2 out of 26
2017). In a bonobo fetus, rhomboideus was fused with its individuals with trisomy 18 (7.7%).
counterpart on the other side of the body and partially fused
with levator scapulae and serratus anterior (Diogo et al. Clinical Implications
2017). N/A
Variations
rhOMbOiDeus MinOr (rhOMbOiD MinOr) (figure 3.1)
Description
See also: Rhomboideus major
The rhomboid major muscle may vary in its vertebral
attachments, with the inferior most attachments varying Synonyms
anywhere between the third and sixth thoracic vertebrae
Rhomboideus minor may also be referred to as rhomboi-
(Macalister 1875; Mori 1964; Bakkum and Miller 2016).
deus superior or kleiner Rautenmuskel (Bakkum and Miller
Rhomboideus major can also be divided into many discrete
2016).
bundles (Macalister 1875; Bergman et al. 1988; Bakkum
and Miller 2016). When fasciculated, it may connect with Typical Presentation
serratus anterior (Macalister 1875). The lower part of the
Description
muscle may be bilaminar (Macalister 1875). The rhom-
boid muscles may be fused (Macalister 1875; Mori 1964; Rhomboid minor originates from the inferior end of the
Bergman et al. 1988). Together, the rhomboids can send ligamentum nuchae and spines of the seventh cervical and
slips to infraspinatus, latissimus dorsi, or teres major frst thoracic vertebrae (Standring 2016). It attaches to the
(Macalister 1875; Rickenbacher et al. 1985; Bergman et al. medial border of the scapula at the root of the scapular
1988; Bakkum and Miller 2016). Rhomboideus minimus, spine (Standring 2016).
or rhomboid minus, refers to a slip that extends between
Innervation
the upper thoracic or lower cervical vertebral spines and
the scapula and/or teres major fascia (Mori 1964; Bergman Rhomboideus minor is innervated by the dorsal scapular
et al. 1988; Bakkum and Miller 2016). nerve (Standring 2016).
Prevalence Comparative Anatomy

Based on a study of 60 sides, Mori (1964) found that the In the apes, the rhomboid muscles are typically fused and
inferior most vertebral attachment was T3 on two sides extend between the cervical and thoracic vertebrae and
(3.3%), T4 on 20 sides, (33.3%), T5 on 28 sides (46.7%), and the scapula (Macalister 1873; Deniker 1885; Kohlbrügge
T6 on 10 sides (16.7%). Mori (1964) also observed that the 1890–1892; Schück 1913; Sonntag 1923, 1924; Stewart
rhomboids were fused on 70 out of 500 sides (14%). Mori 1936; Raven 1950; Miller 1952; Preuschoft 1965; Swindler
(1964) reports the presence of rhomboideus minimus in and Wood 1973; Gibbs 1999; Diogo et al. 2010, 2012,
56 out of 505 sides (11.1%) and states that Nishi found this 2013a,b, 2017). In a bonobo fetus, rhomboideus was fused
muscle in 16% of cases. with its counterpart on the other side of the body and par-
tially fused with levator scapulae and serratus anterior
Anomalies (Diogo et al. 2017).
Description
Variations
In both a neonate with trisomy 18 (Aziz 1979) and a
fetus with craniorachischisis (Alghamdi et al. 2017), the Description
rhomboid muscles were present as a continuous sheet of The rhomboid minor muscle may vary in its vertebral attach-
muscle. Rhomboideus major and minor were also fused ments, with the superior most attachments varying anywhere
in a trisomy 18 cyclopic fetus dissected by Smith et al. between the fourth and sixth cervical vertebrae (Macalister
(2015), and these muscles attached higher onto the scapula 1875; Mori 1964; Rickenbacher et al. 1985; Bakkum and
than is normally observed. Furthermore, the composition Miller 2016). Rhomboideus minor can also be divided into
many discrete bundles (Bergman et al. 1988; Bakkum and Synonyms

Miller 2016). It may also be bilaminar (Macalister 1875). Rhomboideus occipitalis may also be referred to as occipito-
The rhomboid muscles may be fused (Macalister 1875; Mori scapularis (Wood), omo-occipitalis, rhomboideus cervicis,
1964; Bergman et al. 1988). Together, the rhomboids can rhomboideus capitis, levator scapulae minor vel posterior,
send slips to infraspinatus, latissimus dorsi, or teres major or levator anguli scapulae minor (Wood 1867a,b; Macalister
(Macalister 1875; Rickenbacher et al. 1985; Bergman et al. 1875; Knott 1883a; Patten 1935; Aziz 1981; Sullivan and
1988; Bakkum and Miller 2016). Rhomboideus minimus, Osgood 1927; Jouffroy 1971; Bakkum and Miller 2016).
or rhomboid minus, refers to a slip that extends between
the upper thoracic or lower cervical vertebral spines and Typical Presentation
the scapula and/or teres major fascia (Mori 1964; Bergman This muscle is only present as a variation or anomaly.
et al. 1988; Bakkum and Miller 2016).
Comparative Anatomy
Prevalence
Among the apes, rhomboideus occipitalis seems to only be
Based on a study of 60 sides, Mori (1964) found that
present in orangutans as a variation and extends from the
the superior most vertebral attachment was C4 on 10
medial border of the scapula to the cranium (Diogo et al.
sides (~16.7%), C5 on 36 sides (60%), and C6 on 14 sides
2013b). In orangutans, this muscle is innervated by C4 and
(~23.3%). Mori (1964) also observed that the rhomboids
C5 (Kallner 1956).
were fused on 70 out of 500 sides (14%). Mori (1964)
reports the presence of rhomboideus minimus in 56 out of
Variations
505 sides (~11.1%) and states that Nishi found this muscle
in 16% of cases. Description
Rhomboideus occipitalis is present in karyotypically nor-
Anomalies mal humans as an extremely rare variation (Rogawski
Description 1990; Smith et al. 2015). It is not known whether this
In both a neonate with trisomy 18 (Aziz 1979) and a fetus muscle appears during normal muscle formation and then
with craniorachischisis (Alghamdi et al. 2017), the rhom- disappears before birth. However, Jelev and Landzhov
boid muscles were present as a continuous sheet of muscle. (2012–2013) suggest that variation in the rhomboid mus-
Rhomboideus major and minor were also fused in a trisomy cles may happen in the embryo when the rhomboid mass
18 cyclopic fetus dissected by Smith et al. (2015), and these migrates to its usual position after its formation at CR14 mm
muscles attached higher onto the scapula than is normally (crown-rump length of 14 mm).
observed. Furthermore, the composition of the rhomboid When present, rhomboideus occipitalis passes from the
muscles in this specimen was about one-third fibrous and superior nuchal line on the occipital bone to attach to the
two-thirds tendinous (Smith et al. 2015). The bilateral pres- medial border of the scapula at the level of the scapular
ence of a bipartite rhomboid minor with superficial and spine (Wood 1867a,b; Knott 1883a; Bakkum and Miller
deep parts was observed in a 26-week-old male fetus with 2016). As noted by Patten (1935), Zağyapan et al. (2008),
CHARGE syndrome (Beger et al. 2019). Stevenson et al. and Stanchev et al. (2017), this muscle can also originate
(2014) describe a cadaver with a severe case of congenital from the superior angle of the scapula between the origins
scoliosis and found significant asymmetry in the rhomboid of rhomboideus minor and levator scapulae.
muscles. In the case described by Patten (1935), rhomboideus
occipitalis sent slips to the fascia of serratus posterior
Prevalence superior. This muscle can also send slips to levator scapu-
In the literature review completed by Smith et al. (2015), lae, splenius, and serratus anterior (Wood 1870; Bakkum
they found that rhomboideus major and minor were fused in and Miller 2016). It may also be connected to trapezius
2 out of 26 individuals with trisomy 18 (7.7%). (Macalister 1875). In the cadaver examined by Rogawski
(1990), there were three other muscular variations on the
Clinical Implications left side of the body, including a third head of biceps bra-
N/A chii, absence of plantaris, and insertion of the tertius onto
the distal phalanx of the fifth digit. Stanchev et al. (2017)
describe a bilateral presentation of rhomboideus occipitalis
Rhomboideus occipitalis (Figure 3.1) that appeared to be comprised of an inferior oblique and
Entry adapted by permission from Springer Nature Customer superior oblique part on the left side, and an inferior oblique,
Service Centre GmbH: Springer Current Molecular Biology middle straight, and superior oblique part on the right side.
Reports, Muscles Lost in Our Adult Primate Ancestors
Still Imprint in Us: on Muscle Evolution, Development, Innervation
Variations, and Pathologies. E. Boyle, V. Mahon, R. Diogo, Zağyapan et al. (2008) and Stanchev et al. (2017) observed
2020. innervation of rhomboideus occipitalis by the dorsal scapu-
See also: Rhomboideus major, Rhomboideus minor lar nerve.
N/A Description
Rhomboideus tertius is a rare variation of the rhomboid
Anomalies
muscles. Jelev and Landzhov (2012–2013) document a case
Description in which rhomboideus tertius is present bilaterally, originat-
Mieden (1982) describes two male fetuses with cyclo- ing from the spinous processes of the sixth, seventh, and (on
pia and alobar holoprosencephaly. Rhomboideus occipi- the left side) eighth thoracic vertebrae and attaching to the
talis was present on the right side of one specimen and inferior most part of the medial border of the scapula. The
on the left side of the other specimen. In both cases, the muscle was nearly 1.5 times as wide (40 mm) on the left
muscle extended between the superior nuchal line and the side than on the right side (27 mm). Lee and Jung (2015)
scapular attachment of rhomboid minor (Mieden 1982). describe a similar case in which this muscle originates from
In an otocephalic fetus examined by Lawrence and Bersu the spinous processes of the fourth and ffth thoracic verte-
(1984), rhomboideus occipitalis extended from the spinous brae on the left side and the spinous processes of the second
processes of the fourth and ffth cervical vertebrae to the through ffth thoracic vertebrae on the right side. This case
medial border of the scapula, just above the level of the also presented with asymmetry, as the right muscle was
scapular spine. about 3.5 times as wide at origin (90.50 mm) than the left
Aziz (1979) found rhomboideus occipitalis present muscle (25.13 mm).
bilaterally in a neonate with trisomy 18. It arose deep to
trapezius near the superior nuchal line and attached to the Innervation
scapula. In a fetus with trisomy 18 and cyclopia, rhomboi- Rhomboideus tertius is innervated by the dorsal scapular
deus occipitalis was part of a complex with trapezius and a nerve (Jelev and Landzhov 2012–2013).
broad sternocleidomastoid (Smith et al. 2015). Rhomboideus
occipitalis was also present in a fetus with craniorachischi- Prevalence
sis (Alghamdi et al. 2017). On the left sides of one adult and N/A
one fetus with trisomy 21, Bersu (1980) found an “occipito-
scapular” muscle that extended between the superior nuchal Anomalies
line and serratus anterior. N/A
Prevalence Clinical Implications

In the literature review conducted by Smith et al. (2015), Jelev and Landzhov (2012–2013) suggest that when present,
they found that rhomboideus occipitalis was present in 1 rhomboideus tertius may be used with rhomboid major and
out of 24 individuals with trisomy 13 (4.2%) and 8 out of rhomboid minor for intrathoracic muscle fap transfer, or for
26 individuals with trisomy 18 (30.8%). The “occipito- muscle transfer for the paralysis of trapezius.
scapular” muscle described by Bersu (1980) was found in
two out of fve individuals with trisomy 21 (40%).
levatOr scapulae (figure 3.1)
Clinical Implications Synonyms
N/A Levator scapulae may also be referred to as levator anguli
scapulae (Macalister 1875).
rhOMbOiDeus tertius (nOt illustrateD)
See also: Rhomboideus major, Rhomboideus minor Description
Synonyms Levator scapulae originates from the transverse processes
of the frst four cervical vertebrae and inserts onto the
It is possible that rhomboideus tertius may be synonymous
medial border of the scapula (Standring 2016).
with rhomboideus minimus (rhomboideus minus), as it is
in a similar location to this muscle (Jelev and Landzhov Innervation
2012–2013).
Levator scapulae is innervated C3 and C4, and by the dorsal
scapular nerve (C5) (Standring 2016).
This muscle is only present as a variation. Comparative Anatomy
Levator scapulae has a similar typical presentation in the
Comparative Anatomy apes, extending between various cervical vertebrae and the
In the apes, the rhomboid muscles are typically fused, and scapula (Gratiolet and Alix 1866; Champneys 1872; Deniker
there is no portion that has been designated as a “rhomboi- 1885; Kohlbrügge 1890–1892; Hepburn 1892; Primrose
deus tertius” (Diogo et al. 2010, 2012, 2013a,b, 2017). 1899, 1900; Pira 1913; Schück 1913; Sonntag 1923, 1924;
Sullivan and Osgood 1927; Stewart 1936; Raven 1950; Anomalies

Miller 1952; Preuschoft 1965; Swindler and Wood 1973; Description
Gibbs 1999; Michilsens et al. 2009; Diogo et al. 2010, 2012,
In a fetus with craniorachischisis, levator scapulae was
2013a,b, 2017).
absent on the right side and fused with the distal portion
Variations of the middle scalene and rhomboideus occipitalis on the
left side (Alghamdi et al. 2017). In a male fetus with trip-
Description
loidy, Moen et al. (1984) found that the muscle bellies of
Levator scapulae may be completely absent (Rickenbacher levator scapulae were replaced by tendons on both sides
et al. 1985). Levator scapulae may vary in its vertebral of the body. Levator scapulae may blend with the origin of
attachments (Mori 1964; Macalister 1875; Rickenbacher scalenus posterior in cases of trisomy 18 (Smith et al. 2015).
et al. 1985; Standring 2016; Bakkum and Miller 2016). Mieden (1982) describes two male fetuses with cyclo-
Levator scapulae can separate into multiple slips and may pia and alobar holoprosencephaly. On the right side of one
have additional attachments to the temporal bone/mastoid specimen, levator scapulae was divided into superfcial and
process, occipital bone, frst or second rib, upper thoracic deep heads, with the former attaching to the frst and sec-
vertebrae, ligamentum nuchae, scalene muscles, longissi- ond cervical vertebrae and the latter to the third and fourth
mus cervicis, subclavius, subscapularis, trapezius, splenius cervical vertebrae. A supernumerary slip of the superfcial
capitis, rhomboideus muscles, or serratus muscles (Wood head originated from the third rib just posterior to serratus
1870; Curnow 1873; Flower and Murie 1867; Macalister anterior (Mieden 1982).
1875; Humphry 1873; Mori 1964; Rickenbacher et al. 1985;
Bergman et al. 1988; Loukas et al. 2006a, Lima et al. 2012; Prevalence
Standring 2016; Bakkum and Miller 2016; Au et al. 2017).
In their literature review, Smith et al. (2015) found that the
Gonzales et al. (2017) describe a variable presentation of
origin of scalenus posterior was blended with levator scapu-
this muscle on the right side of a cadaver in which the upper
lae in 1 out of 26 individuals with trisomy 18 (3.8%).
fbers of levator scapulae fused with the inferior attachment
of rectus capitis lateralis. Au et al. (2017) suggest that the
frequent connections between levator scapulae, serratus
anterior, and serratus posterior superior may be due to the Shpizner and Holliday (1993) suggest that asymmetry of
shared somitic cell origin of these muscles. A slip of leva- levator scapulae may present as a palpable muscle mass in
tor scapulae that extends from some of the cervical verte- the posterior triangle and may be confused with disease.
brae to the lateral end of the clavicle and the acromion has Loukas et al. (2006a) suggest that variable slips and inser-
been identifed as levator claviculae (see the entry for this tions of levator scapulae may contribute to myofascial pain
muscle) (Bergman et al. 1988). syndrome.
Prevalence
subclavius (figure 3.2)
Based on observations from 60 sides, Mori (1964) reported
that levator scapulae arose from cervical vertebrae one See also: Sternoscapularis, Pectoralis intermedius,
through four on 40 sides (66.7% of cases), from the frst Subclavius posticus
three cervical vertebrae on 16 sides (26.7%), from the frst
two cervical vertebrae on two sides (3.3%), and from the Synonyms
frst fve cervical vertebrae on two sides (3.3%). This author N/A
also found two types of anomalous slips. One type ran from
the medial margin of levator scapulae to the spinous pro- Typical Presentation
cess of the second thoracic vertebra, to the dorsal surface Description
of serratus posterior superior, or to the lumbodorsalis fascia Subclavius originates from the frst rib and its costal carti-
in 9 out of 100 sides (9%) (Mori 1964). The other ran from lage and inserts onto the inferior aspect of the middle third
the medial margin of levator scapulae to the ventral surface of the clavicle (Standring 2016).
of subscapularis in 22 out of 100 sides (22%) (Mori 1964).
Au et al. (2017) studied variations of levator scapulae Innervation
using MRI and found that this muscle had caudal accessory Subclavius is innervated by the subclavian nerve (Standring
attachments in 16 out of 37 subjects (43.2%). Ten subjects 2016).
exhibited unilateral attachments to serratus anterior, ser-
ratus posterior superior, or the frst or second rib. Four Comparative Anatomy
subjects exhibited bilateral attachments to serratus ante- Subclavius has a similar typical presentation in the apes,
rior. One subject exhibited bilateral attachments to serratus extending from the frst rib to the clavicle (Primrose 1899,
posterior superior and a unilateral attachment to serratus 1900; Sullivan and Osgood 1927; Stewart 1936; Kallner
anterior. One subject had bilateral attachments to both 1956; Gibbs 1999; Diogo et al. 2010, 2012, 2013a,b,
serratus anterior and serratus posterior superior. 2017). There may be an origin from the second rib in
FIGURE 3.2 Thoracoclavicular muscles in anterior view.
orangutans, and from the second and/or third ribs in gib- Prevalence
bons (Kohlbrügge 1890–1892; Hepburn 1892; Sonntag Out of 72 cases, Mori (1964) noted that 12 out of 72 sides
1924; Gibbs 1999; Michilsens et al. 2009). (16.7%) exhibited type I variation (see above), while 6 out of
72 sides (8.3%) exhibited type II variation.
Variations
Anomalies
Description
Description
Subclavius may also arise from the second rib (Bergman
Subclavius was absent bilaterally in a neonate with trisomy
et al. 1988). Subclavius may insert onto the coracoclavicu-
18 (Aziz 1979) and a neonate with trisomy 13 (Aziz 1980).
lar ligament, transverse ligament of the scapula, acromion
process, coracoid process, the superior border of the scap- Prevalence
ula, or the humerus (Macalister 1875; Bergman et al. 1988;
In their literature review, Smith et al. (2015) found that sub-
Standring 2016). Subclavius may be doubled or divided
clavius was absent in 10 out of 20 individuals with trisomy 13
into two parts (Macalister 1875). This muscle may also be
(50%) and in 11 out of 17 individuals with trisomy 18 (64.7%).
completely absent and in some cases can be replaced by a
fbrous band (Macalister 1875; Knott 1883a; Crerar 1892; Clinical Implications
Georgiev and Jelev 2009; Snosek and Loukas 2016; Yun Georgiev and Jelev (2009) found that the fbrous band that may
et al. 2018). replace subclavius in cases of its absence can reduce the cos-
Mori (1964) observed that subclavius could have a lat- toclavicular space and possibly lead to entrapment of the bra-
eral end that divides into two portions: one inserting on chial plexus and the subclavian/axillary vessels and could thus
the inferior surface of the clavicle and the other inserting contribute to the development of thoracic outlet syndrome.
onto the base of the coracoid process (type I variation).
Alternatively, the lateral end of the muscle may insert into
the inferior surface of the clavicle, but then courses on to
pectOralis interMeDius (figure 3.2)
insert onto the base of the coracoid or the superior border of See also: Subclavius, Pectoralis major, Pectoralis minor
the scapula (type II variation).
Subclavius is associated with scapuloclavicularis, which Synonyms
passes from the base of the coracoid and transverse scapu- N/A
lar ligament to the posterior side of the clavicle (Figure 3.3)
(Macalister 1875; Bergman et al. 1988). It may be con- Typical Presentation
nected to omohyoid at its origin and subclavius at its inser- This muscle is only present as a variation.
tion (Macalister 1875). A variant of scapuloclavicularis is
scapulocostoclavicularis, which originates from the upper Comparative Anatomy
border of the scapula and inserts onto the clavicle and frst Pectoralis intermedius was not present in the common
costal cartilage (Bergman et al. 1988). Other variations of chimpanzees or bonobos dissected by Potau et al. (2018),
subclavius include sternoscapularis, pectoralis intermedius, and there is little information about this muscle for other
and subclavius posticus (see the entries for these muscles). species.
Variations laterally over the brachial plexus and subclavian vessels,

Description and ended on the fascia of serratus anterior between its
frst two digitations near the superior border of the scapula.
Pectoralis intermedius is a variant of subclavius (Bergman
et al. 1988). It originates from the third and fourth ribs, Innervation
in between pectoralis major and minor, and inserts onto Innervation to subclavius posticus is variable (Cogar et al.
the coracoid process (Bergman et al. 1988). Mori (1964) 2015). This muscle can be innervated by the subclavian
describes pectoralis intermedius as the deep layer of pec- branch of the brachial plexus, the nerve to the omohyoid
toralis major that arises from ribs three through six and is from the ansa cervicalis, or the suprascapular nerve (Cogar
fused with the lateral insertion of pectoralis major. Sawada et al. 2015).
et al. (1991) observed that pectoralis intermedius can arise
from the fourth and ffth ribs. Arican et al. (2006) found Prevalence
that this muscle originated from the third and fourth ribs Macalister (1875) states that subclavius posticus is present
and joined the tendon of the short head of biceps brachii in about 1 out of 15 subjects (6.7%). Mori (1964) found this
upon insertion, just below the coracoid process. muscle in fve out of 500 sides from 250 cadavers (1%) and
states that Nishi found this muscle in 2 out of 12 cadavers
Innervation (16.7%). Akita et al. (2000) recorded the presence of this
Pectoralis intermedius is innervated by the lateral pectoral muscle in 11 out of 124 cadavers (8.9%).
nerve (Arican et al. 2006).
Anomalies
Mori (1964) states that pectoralis intermedius was found in In individuals with trisomy 13 and trisomy 18, subclavius
112 out of 351 cases (31.9%). posticus extends from the upper margin of the scapula, and
sometimes the transverse scapular ligament, to the frst rib
Anomalies (Ramirez-Castro and Bersu 1978; Aziz 1979; Smith et al.
N/A 2015).
Prevalence
In the literature review conducted by Smith et al. (2015),
Potau et al. (2018) suggest that pectoralis intermedius could
they found that subclavius posticus was present in 3 out of
be mistaken for masses or tumors in medical imaging or
20 individuals with trisomy 13 (15%) and 5 out of 17 indi-
interfere with the surgical use of pectoralis major faps.
viduals with trisomy 18 (29.4%).
subclavius pOsticus (figure 3.2) Clinical Implications

The presence of subclavius posticus can contribute to
See also: Subclavius
thoracic outlet syndrome and its consequences, including
Synonyms Paget-von-Schroetter syndrome (Akita et al. 2000; Forcada
et al. 2001; Cogar et al. 2015). Kolpattil et al. (2009) report
This muscle is also referred to as scapulo-costalis (Knott
that the presence of the muscle could be confused with a
1883a), scapulocostalis minor (Gruber) (Macalister 1875),
pathological mass on a mammogram.
or chondroscapularis (Shetty et al. 2006).
Typical Presentation levatOr claviculae (figure 3.3)

This muscle is only present as a variation or anomaly. Entry adapted by permission from Springer Nature Customer
Service Centre GmbH: Springer Current Molecular Biology
Comparative Anatomy
Reports, Muscles Lost in Our Adult Primate Ancestors Still
N/A Imprint in Us: on Muscle Evolution, Development, Variations,
and Pathologies. E. Boyle, V. Mahon, R. Diogo, 2020.
Variations
See also: Levator scapulae
Description
This muscle is a variant of subclavius (Bergman et al. 1988). Synonyms
Subclavius posticus (Rosenmüller) extends from the frst Levator claviculae is also referred to as omocervicalis, leva-
rib or the frst costal cartilage and inserts onto the superior tor scapulae anticus, or levator scapulae ventralis (Diogo
margin of the scapula (Knott 1883a; Mori 1964; Bergman et al. 2018), trachelo-clavicularis superior (Knott 1883a),
et al. 1988; Snosek and Loukas 2016). Moyano et al. (2018) cleido-cervicalis superior (Knott 1883a) or the cleidocervi-
report the presence of subclavius posticus in an adult male cal muscle (Leon et al. 1995), cleidoatlanticus (Rodríguez-
cadaver that originated from the cranial surface of the frst Vázquez et al. 2009), cleidotrachelian (Newell et al. 1991), or
costal cartilage near the insertion of subclavius, coursed musculus trachleo-acromialis (Kuiper et al. 2014).
FIGURE 3.3 Lateral view of additional thoracoclavicular variations, shown relative to trapezius and sternocleidomastoid.
Typical Presentation considered a variant of levator scapulae (Loukas et al.

This muscle is only present as a variation or anomaly. 2008b; Odate et al. 2012), levator claviculae has been
argued to have embryological origins from scalenus ante-
Comparative Anatomy rior (Gruber 1876), sternocleidomastoid (Wood 1870), tra-
pezius (Parsons 1898a), longus colli (Tomo et al. 1994), or
Levator claviculae is normally present as a distinct muscle
the ventrolateral muscle primordia of the neck (Leon et al.
in the apes (Diogo et al. 2010, 2012, 2013a,b, 2017). In the
1995). In their review, Odate et al. (2012) arrive at the con-
apes, levator claviculae typically passes deep to trapezius,
clusion that levator scapulae shares a common embryologi-
originating from the atlas and inserting onto the lateral
cal origin with levator scapulae (i.e., arise from the same
end of the clavicle (Gratiolet and Alix 1866; Champneys
myotome), based on the work of McKenzie (1955) and
1872; Deniker 1885; Kohlbrügge 1890–1892; Chapman
shared characteristics between these two muscles, includ-
1900; Schück 1913; Sullivan and Osgood 1927; Stewart
ing similar innervation and origin/insertion. This idea is
1936; Raven 1950; Miller 1952; Kallner 1956; Preuschoft
reinforced by a detailed study of the comparative myology
1965; Jouffroy 1971; Gibbs 1999; Diogo et al. 2010, 2012,
of tetrapods (Diogo et al. 2018).
2013a,b, 2017). It may have an additional attachment onto
When present, levator claviculae seems to be found
the scapula in gorillas (Preuschoft 1965).
more often on the left side than on the right side of the
body (Rubinstein et al. 1999; Loukas et al. 2008b; Odate
Variations
et al. 2012; Ferreli et al. 2019). Levator claviculae extends
Description from the transverse processes of various cervical vertebrae
Early in development, specifcally between CR9 mm (rarely from the frst, but often from the second and third,
(crown-rump length of 9 mm) to CR 10.5 mm, the axial or third and fourth) and attaches onto the middle third or
pectoral muscles resemble those of adult humans as sub- lateral end of the clavicle, and the acromion (Wood 1864,
clavius is present but levator claviculae is absent (Diogo Macalister 1875; Macdonald Brown 1880; Knott 1883a;
and Abdala 2010). Though closely associated with and Rickenbacher et al. 1985; Bergman et al. 1988; Leon et al.
1995; Ginsberg and Eicher 1999; Loukas et al. 2008b; Odate Typical Presentation
et al. 2012; Raikos et al. 2012; Bakkum and Miller 2016). It This muscle is only present as a variation.
may also originate from the ffth or sixth cervical vertebrae
(Macalister 1875; Knott 1883a; Billings and Sherrill 2015). Comparative Anatomy
Levator claviculae may insert onto the sternocleidomastoid N/A
or serratus anterior muscles (Flower and Murie 1867; Feigl
and Pixner 2011) or blend with trapezius (Macalister 1875; Variations
Knott 1883a). Description
This muscle is a variant of subclavius (Bergman et al.
Innervation
1988). It arises from the sternum, courses between the
This muscle can be innervated by cervical spinal nerve 3, 4, clavicle and the coracoid process, and then attaches to the
5, or 6 (Leon et al. 1995; Loukas et al. 2008b; Odate et al. medial border of the acromion (Huntington 1904; Snosek
2012) and Loukas 2016).
Prevalence Innervation
Wood (1870) observed that levator claviculae is present This muscle is innervated by the lateral pectoral nerves
in only 4 out of 202 humans (2%). Loukas et al. (2008b) (Snosek and Loukas 2016).
observed levator claviculae in only 2 out of 2000 cadavers
(0.1%). Based on CT scans of 300 individuals, Rubinstein Prevalence
et al. (1999) noted seven instances of the muscle (one bilat- N/A
eral case, fve unilateral cases) in six individuals (2%).
Review articles (e.g., Leon et al. 1995; Ginsberg et al. 1999; Anomalies
Rubinstein et al. 1999; Odate et al. 2012) report a preva- N/A
lence of 2%–3%.
Anomalies N/A
Description
Levator claviculae was present bilaterally in an adult with supraclavicularis prOprius (figure 3.3)
trisomy 21, extending from the basilar portion of the occipi-
Synonyms
tal bone to the clavicle (Bersu 1980).
This muscle is also referred to as tensor fascia colli (Gruber)
(Laidlaw 1902) or anomalus claviculae (Knott 1883a).
Prevalence
In the literature review conducted by Smith et al. (2015), Typical Presentation
they found that levator claviculae was present in one out of This muscle is only present as a variation.
fve individuals with trisomy 21 (20%).
Comparative Anatomy
Levator claviculae can be misidentifed as various patholo-
gies. It can resemble a neck mass, cyst, lymphadenopathy Variations
or cervical adenopathy, metastasis, neurofbroma, arte- Description
rial aneurysm, or thrombosed vein, and can present as an Supraclavicularis proprius courses from the sternal end of
abnormal levator scapulae or sternocleidomastoid (Rüdisüli the clavicle to the acromial end and is sheathed in cervical
1995; Ginsberg et al. 1999; Rubinstein et al. 1999; Santiago fascia (Macalister 1875; Bergman et al. 1988; Snosek and
et al. 2001; Feigl and Pixner 2011; Odate et al. 2012). Loukas 2016). It may have origins from the sternoclavicu-
This muscle may also compress the supraclavicular nerve lar joint and the fascia over sternocleidomastoid (Macalister
(Billings and Sherrill 2015) or contribute to thoracic outlet 1875; Bergman et al. 1988; Ottone and Medan 2009; Snosek
syndrome (Aydoğ et al. 2007). and Loukas 2016). It inserts onto the distal portion of the
clavicle or the trapezius fascia (Macalister 1875; Laidlaw
1902; Bergman et al. 1988; Ottone and Medan 2009; Raikos
sternOscapularis (figure 3.3)
et al. 2014; Snosek and Loukas 2016). Raikos et al. (2014)
See also: Subclavius describe a case in which this muscle arose posteriorly from
the medial surface of the clavicle, formed a muscular arch
Synonyms over the supraclavicular nerve, and split into three slips that
Sternoscapularis also known as sternoacromialis (Snosek inserted onto the trapezius, acromion, and the distal end of
and Loukas 2016). the clavicle.
Innervation Typical Presentation

This muscle is innervated by the supraclavicular nerve This muscle is only present as a variation or anomaly.
(Laidlaw 1902).
Comparative Anatomy
Prevalence Sternalis is normally absent in nonhuman primates (Diogo
Ottone and Medan (2009) observed the presence of this muscle and Wood 2012a). According to the literature reviews and
on the left side of 1 out of 156 sides from 78 cadavers (0.64%). dissections done by Diogo et al. (2010, 2012, 2013a,b, 2017),
this muscle is absent in orangutans, gorillas, common chim-
Anomalies panzees, and bonobos, and its presence is variable in gibbons.
N/A
Variations
Ottone and Medan (2009) and Raikos et al. (2014) suggest that Sternalis has several presentations and varies widely in
the presence of supraclavicularis proprius could be a poten- width, length, and attachments (Macalister 1875; Snosek
tial cause of supraclavicular nerve entrapment syndrome. and Loukas 2016; Standring 2016). When present, sternalis
is a vertical slip of fbers in the anterior thoracic wall that
PECTORAL GIRDLE AND ARM MUSCLES
lies close to the sternum and runs on top of pectoralis major,
sternalis (figure 3.4) extending from the sternal/infraclavicular area to the upper
abdominal wall or costal cartilages (Mori 1964; Jouffroy
Synonyms 1971; Jelev et al. 2001; Snosek and Loukas 2016; Standring
This muscle may also be referred to as episternalis, japoni- 2016). It may originate from the manubrium, sternum, clav-
cas, parasternalis, presternalis, rectus sternalis, rectus icle, upper ribs, upper costal cartilages, pectoralis fascia,
sterni, rectus thoracis, rectus thoracicus superfcialis, ster- pectoralis major and/or sternocleidomastoid and its fas-
nalis brutorum, superfcial rectus abdominis, and thoracicus cia (Macalister 1875; Mori 1964; Jelev et al. 2001; Hung
(Macalister 1875; Jelev et al. 2001; Snosek and Loukas 2016). et al. 2012; Poveda et al. 2013; Snosek and Loukas 2016;
FIGURE 3.4 Thoracohumeral muscles in anterior view.

Standring 2016; Duque-Parra et al. 2019). It may insert into Prevalence

the lower ribs and costal cartilages, pectoral fascia, pecto- Sternalis is estimated to be present in 3%–8% of the world
ralis major, external oblique aponeurosis, and/or the rectus population (Pérez et al. 2008). Snosek and Loukas (2016)
sheath (Macalister 1875; Mori 1964; Jelev et al. 2001; Hung provide a comprehensive list of prevalence rates from
et al. 2012; Poveda et al. 2013; Snosek and Loukas 2016; cadaveric studies and clinical investigations and state that
Standring 2016; Duque-Parra et al. 2019). Duque-Parra the overall prevalence in the general population is about
et al. (2019) found that sternalis fibers extended between 7.8%. Macalister (1875) states that Gruber found sternalis
sternocleidomastoid to the fascia of the seventh intercostal in 5 out of 95 cases (5.3%) and that Turner found it in 21
muscle. In this case, the muscle had two bellies. out of 650 subjects (3.2%). Macalister (1875) himself found
Sternalis can be partially or completely absent and sternalis present in 11 out of 350 subjects (3.1%) and con-
also can have unilateral or bilateral presentations (Young nected to the sternal head of sternocleidomastoid in 2 out of
Lee et al. 2006; Duque-Parra et al. 2019), though a unilat- 600 subjects (0.3%).
eral presence is more frequent (Snosek and Loukas 2016; Prevalence can be anywhere from ~1%, as observed in
Standring 2016). When muscles are present on both sides, a Chilean population (Molina et al. 2017) and a Taiwanese
they may fuse across the sternum (Mori 1964; Snosek and population, to 23.5% as observed in a Chinese popu-
Loukas 2016). Kalpana and Usha (2010) and Dudgeon lation (Jelev et al. 2001; Raikos et al. 2011; Vishal et al.
et al. (2017) report sternalis muscles with three bellies. 2013; Snosek and Loukas 2016). Sato (1968c) reported
Macalister (1875) states that tendinous intersections in the that sternalis was present in 61 out of 426 sides (14.3%)
muscle have been observed by some researchers. in Kyushu-Japanese males and present in 42 of 266 sides
Snosek and Loukas (2016) provide a comprehensive review (15.79%) in females. Mehta et al. (2010) found sternalis in 1
of this muscle, including speculation on its homology and out of 88 cadavers of Asian origin (1.1%).
embryological origin. The authors review the four common Mori (1964) searched for sternalis in 350 Japanese male
arguments for origin of sternalis—that it is a remnant of pan- cadavers and 25 Japanese female cadavers. This author
niculus carnosus, a downward extension of sternocleidomas- found sternalis in 36 male cadavers (10.3%) and 3 female
toid, an upward extension of rectus abdominis, or a derivative cadavers (12%). Among males, 12 presented the muscle
of pectoralis major—and appear to favor this last hypothesis, bilaterally and 24 presented the muscle unilaterally, while
suggesting that sternalis should be classified as one of the two females presented the muscle bilaterally and one pre-
many muscular variations caused by abnormal pectoral mus- sented the muscle unilaterally. Therefore, sternalis was
cle development (Snosek and Loukas 2016). Standring (2016) present in 53 out of 750 sides (7.1%). Mori (1964) also
also considers sternalis a variant of pectoralis major. classified the 53 instances of sternalis into six types and
Snosek and Loukas (2016) also note that other super- noted that on seven sides (1%) both ends of the muscle
numerary muscles found on the anterior thoracic wall may attached to the pectoral fascia, on seven sides (1%) the
be described as novel muscle variants but are more likely muscle extended from the sternum superiorly to the pecto-
variants of sternalis. Examples include oblique pectoralis ral fascia inferiorly, on 12 sides (1.7%) the muscle extended
anterior, which passes from the sternum to the lower ribs from the sternum superiorly to the rectus sheath inferiorly,
and the rectus sheath (Huber et al. 2012), rectus thoracis on eight sides (1.1%) the muscle extended from the pec-
bifurcalis, which extends between the aponeurosis of exter- toral fascia superiorly to the rectus sheath inferiorly, on
nal oblique and the manubrium and sends two diverging two sides (0.3%) the muscle extended from the tendon of
aponeurotic bands to sternocleidomastoid (Mehta et al. sternocleidomastoid superiorly to the pectoral fascia infe-
2010), and sternomastalis, which passes from the pectoral riorly, and on 17 sides (2.4%) the muscle extended from
fascia and sternocostal junction to the tissue of the breast the tendon of sternocleidomastoid superiorly to the rectus
and nipple-areola complex (Kale et al. 2006). sheath inferiorly.
Duque-Parra et al. (2019) found this muscle in 2 out
of 68 Colombian cadavers (2.94%). Saeed et al. (2002)
Innervation
found a prevalence of 4% in the Kingdom of Saudi
Musculus sternalis is innervated by the medial or lateral Arabia. In Europeans, prevalence is estimated at 4.4%,
pectoral nerve in most cases (Mori 1964; O’Neill and while prevalence is estimated at 8.4% in Africans (Loukas
Folan-Curran 1998; Kida et al. 2000; Kumar et al. 2003; et al. 2004). Jelev et al. (2001) found sternalis in 3 out
Hung et al. 2012; Snosek and Loukas 2016; Duque-Parra of 102 Bulgarian cadavers (2.9%). Krishnan et al. (2017)
et al. 2019). It may also be innervated the anterior cuta- observed the presence of sternalis in 3 out of 18 cadav-
neous branches of the intercostal nerves, or both of these ers from Wales (16.7%). Sonne (2020) found a bilateral
nerves in rare cases (Mori 1964; O’Neill and Folan-Curran presentation of sternalis in 2 out of 36 cadavers from the
1998; Jelev et al. 2001; Motabagani et al. 2004; Snosek and United States (5.5%). This muscle may be more prevalent
Loukas 2016; Duque-Parra et al. 2019). It may be the case in women than in men (Snosek and Loukas 2016; Aguado-
that sternalis is not supplied by the intercostal nerves, but Henche et al. 2018).
that they may pass near to or through sternalis on their way
to innervate other structures (Snosek and Loukas 2016).

Description While typically absent in adult humans, dorsoepitrochle-
Anomalous presentations are similar to the above, extend- aris is present in nonhuman primates (Diogo and Wood
ing from the infraclavicular region near the sternum to 2011, 2012a). This muscle is present in all apes, typi-
the lower ribs or rectus abdominis (Smith et al. 2015). cally extending from latissimus dorsi to the medial epi-
Macalister (1875) states that Barkow (1828) found sternalis condyle of the humerus (Tyson 1699; Gratiolet and Alix
in an abnormal individual extending between the second 1866; Barnard 1875; Chapman 1880, 1900; Kohlbrügge
and seventh ribs. 1890–1892; Beddard 1893; Dwight 1895; Primrose 1899,
1900; Michaelis 1903; MacDowell 1910; Schück 1913;
Prevalence Sonntag 1923, 1924; Sullivan and Osgood 1927; Raven
1950; Miller 1952; Swindler and Wood 1973; Michilsens
Windle (1893) conducted dissections on ten anencephalic
et al. 2009; Gibbs 1999; Diogo et al. 2010, 2012, 2013a,b,
fetuses and noted the presence of sternalis on the left side in
2017). Dorsoepitrochlearis is innervated by the radial
two females and one male, and on both sides in one female,
nerve in the apes (Gibbs 1999; Diogo et al. 2010, 2012,
yielding a prevalence of 5 out of 20 sides (25%). Abraham
2013a,b, 2017).
(1883) found sternalis in 6 out of 11 anencephalic fetuses
In gibbons, it is often blended with the short head of the
(54.5%). In their literature review, Smith et al. (2015) found
biceps (Howell and Straus 1932; Michilsens et al. 2009;
that sternalis was reported in 1 out of 17 (5.8%) trisomy 18
Diogo et al. 2012). MacDowell (1910) observed a slip from
individuals.
coracobrachialis to dorsoepitrochlearis in one common
chimpanzee. There may be an additional origin from the
Clinical Implications coracoid process in common chimpanzees (Gratiolet and
Sternalis can be mistaken for pathology, especially mam- Alix 1866; Diogo et al. 2013a). It may attach to the inter-
mary cancer, when evaluating mammograms, CT scans, or muscular septum in orangutans (Barnard 1875; Hepburn
MRI scans (Bradley et al. 1996; Goktan et al. 2006; Marques 1982; Diogo et al. 2013b). In orangutans and common chim-
et al. 2009; Mehta et al. 2010; Standring 2016; Molina et al. panzees, there may be an insertion onto the olecranon pro-
2017). Standring (2016) and Snosek and Loukas (2016) note cess (Grönroos 1903; Ziegler 1964; Diogo et al. 2013a,b).
that sternalis may be as useful as a muscular fap in recon-
structive surgeries. Variations
Description
Dorsoepitrochlearis is typically only present in the early
DOrsOepitrOchlearis (figure 3.11)
stages of human development, usually appearing in human
Entry adapted by permission from Springer Nature embryos at CR15 mm (crump-round length of 15 mm)
Customer Service Centre GmbH: Springer Current and disappearing in embryos at CR18 mm (Haninec et al.
Molecular Biology Reports, Muscles Lost in Our Adult 2009). However, Diogo et al. (2019) state that since all other
Primate Ancestors Still Imprint in Us: on Muscle Evolution, arm muscles are differentiated earlier (CR10.5 mm), further
Development, Variations, and Pathologies. E. Boyle, V. research is needed to reveal if dorsoepitrochlearis is differ-
Mahon, R. Diogo, 2020. entiated earlier than CR15 mm.
See also: Latissimus dorsi, Triceps brachii When present in adults, dorsoepitrochlearis originates
from latissimus dorsi near to or on the tendon of inser-
Synonyms tion, passes through the axilla, and inserts onto the medial
Alternative names for this muscle include latissimo-condy- epicondyle of the humerus (Haninec et al. 2009; Bakkum
lus or latissimo-epitrochlearis (Barnard 1875), latissimo- and Miller 2016). It may also insert into the brachial and
condyloideus (Kohlbrügge 1890–1892; Chapman 1878, forearm fascia, the humerus, the lateral epicondyle and
1879; 1880, 1900; Hepburn 1892; Primrose 1899, 1900; olecranon, or the long head of triceps (Bergman et al.
Grönroos 1903; Sommer 1907; MacDowell 1910; Pira 1913; 1988; Bakkum and Miller 2016). When the dorsoepitroch-
Sullivan and Osgood 1927; Loth 1931), latissimo-tricipitalis learis inserts onto the medial epicondyle of the humerus,
(Schück 1913; Fick 1925; Kallner 1956; Preuschoft 1965), its distal part is similar to that of the chondroepitrochlearis
appendix of latissimus dorsi (Tyson 1699), accessorium (Figure 3.4), the main difference being that proximally the
tricipitis (Bergman et al. 1988) or tricipiti accessorius former is often linked to the latissimus dorsi, while the lat-
(Macalister 1875), and tensor fasciae antebrachii, anconeus ter is often linked to the pectoral muscles. Farfán et al.
accessorius, accessorius latissimus dorsi, dorso-antebrachi- (2019) report a case in which dorsoepitrochlearis origi-
alis, anconeus quintus, anconeus longus, or extensor cubiti nated via three fascicles.
sensu Jouffroy (1971).
Innervation
Typical Presentation In three of the cases described by Haninec et al. (2009),
This muscle is only present as a variation or anomaly. dorsoepitrochlearis was innervated by the thoracodorsal
nerve. This differs from the innervation observed in other Diogo and Wood 2011; Diogo et al. 2018), although dissec-
primates (see above). tions by Sommer (1907), Raven (1950), and Diogo et al.
(2010) did not fnd this muscle.
Prevalence
Haninec et al. (2009) observe the presence of this muscle in 4 Variations
out of 209 adults (1.9%). Bergman et al. (1988) and Bakkum Description
and Miller (2016) note this muscle is present in 5% of bodies. Panniculus carnosus is a cutaneous muscle sheet arising
from the pectoral muscle mass and covering various trunk
Anomalies regions (Smith et al. 2015). It is present in many mammals
Description and derives from the pectoralis muscle of amphibians and
An anomalous dorsoepitrochlearis muscle connects latissi- reptiles (Diogo et al. 2018). Bergman et al. (1988) state that
mus dorsi to pectoralis major or to the long head of the tri- panniculus carnosus is represented only by vestigial rem-
ceps (Windle 1893; Aziz 1979; Smith et al. 2015; Alghamdi nants in humans, which may present as extra muscular slips
et al. 2017). Alghamdi et al. (2017) note that in a fetus with in the pectoral region. These slips may have attachments to
craniorachischisis, dorsoepitrochlearis was fused to the the abdominal aponeurosis, rectus sheath, serratus anterior
inner side of the inferior band (costohumeralis) of pectoralis fascia, axillary fascia, the fascia between coracobrachialis
major. Alghamdi et al. (2017) note that this muscle is similar and pectoralis minor, the humerus, or the coracoid process
in presentation to dorsoepitrochlearis found on the right side (Bergman et al. 1988). Several muscles in adult humans are
of one female fetus with anencephaly by Windle (1893). considered to be remnants of panniculus carnosus, includ-
ing some craniofacial muscles, platysma, pectorodorsalis,
Prevalence sternalis, abdominal external oblique, palmaris brevis, and
Testut (1884) and Aziz (1981) suggest that this muscle is potentially several other striated muscles in the upper limb,
present in about 5% of anomalous humans. Barash et al. pectoral region, and trunk (Naldaiz-Gastesi et al. 2018).
(1970) state that this muscle is present in an infant with tri-
Innervation
somy 18, and Aziz (1980, 1981) notes the presence of dorso-
epitrochlearis in two trisomy 18 neonates, but not in three Panniculus carnosus is innervated by various nerves in dif-
trisomy 13 neonates. In the literature review conducted by ferent species, depending on where in the body the pan-
Smith et al. (2015), they noted the presence of this muscle niculus carnosus fbers are located (Naldaiz-Gastesi et al.
in 4 out of 26 individuals with trisomy 18 (15.4%). 2018).
Prevalence
N/A
Haninec et al. (2009) note that the presence of dorsoepi-
trochlearis can compress nerves in the axilla and lead to
Anomalies
symptoms such as paresthesia, pain, and muscle weakness.
Description
Dunlap et al. (1986) describe panniculus carnosus in a fetus
panniculus carnOsus (nOt illustrateD) with trisomy 18 and a fetus with trisomy 21. On the left
Entry adapted by permission from Springer Nature Customer side of the fetus with trisomy 18, panniculus carnosus arose
Service Centre GmbH: Springer Current Molecular Biology from the level of the tenth rib in the midaxillary line and
Reports, Muscles Lost in Our Adult Primate Ancestors Still inserted onto the humerus and onto the deep aspect of pec-
Imprint in Us: on Muscle Evolution, Development, Variations, toralis major. It had a third tendinous insertion into the ten-
and Pathologies. E. Boyle, V. Mahon, R. Diogo, 2020. don of latissimus dorsi, which Dunlap et al. (1986) state is
pectorodorsalis. On the right side, it arose from the level of
Synonyms the seventh rib in the midaxillary line and inserted onto the
According to the summary by Naldaiz-Gastesi et al. (2018), humerus. In the fetus with trisomy 21, panniculus carnosus
alternative names for this muscle include cutaneous trunci, was adjacent to the axillary border of latissimus dorsi and
musculus cutaneous, cutaneous maximus muscle, subcuta- inserted onto the humerus.
neous muscle, and superfcial fascia system.
Prevalence
Typical Presentation In their literature review, Smith et al. (2015) found that pan-
Remnants of panniculus carnosus are present only as varia- niculus carnosus was reported in 1 out of 26 trisomy 18
tions, and the entire muscle is present as a rare anomaly. individuals (3.8%) and 1 out of 7 individuals with trisomy
21 (14.3%).
Comparative Anatomy
This muscle is present in basal primates and several species Clinical Implications
of monkeys and can be vestigial in gorillas (Jouffroy 1971; N/A
Pectoralis major (Figures 3.4 a nd 3.5) In bonobos, pectoralis major can be fused with deltoid,
sternocleidomastoid, and/or pectoralis minor (Diogo et al.
See also: Sternalis, Sternoclavicularis, Pectoralis quartus, 2017). In orangutans, the clavicular head originates from
Pectorodorsalis, Chondroepitrochlearis, Dorsoepitrochlearis, the sternum and does not have an attachment to the clavicle
Pectoralis abdominis (Chapman 1880; Hepburn 1892; Beddard 1893; Primrose
Synonyms 1899, 1900; Michaelis 1903; Sullivan and Osgood 1927;
Stewart 1936; Kallner 1956; Diogo et al. 2013b).
This muscle may also be referred to as the great pectoral
(Macalister 1875).
Variations
Description The three heads of pectoralis major may be completely
Pectoralis major has three heads. The clavicular head distinct from one another (Macalister 1875; Knott 1883a).
originates from the medial half of the clavicle (Standring Absence or reduction of one or more heads has been observed
2016). The sternocostal head originates from the body of (Macalister 1875; Bing 1902; Bergman et al. 1988; Snosek
the sternum and the second through sixth costal cartilages and Loukas 2016; Standring 2016; Haładaj et al. 2019).
(Standring 2016). The abdominal or rectus head originates The clavicular head may be divided (Macalister 1875). It
from the external oblique aponeurosis (Standring 2016). can also extend laterally on the clavicle as far as deltoideus
The muscle inserts via a tendon onto the humerus at the (Bergman et al. 1988; Snosek and Loukas 2016; Haładaj
lateral lip of the intertubercular sulcus (Standring 2016). et al. 2019). The sternocostal head may extend laterally to
latissimus dorsi (Bergman et al. 1988). It may also be bilam-
Innervation inar (Macalister 1875). There may also be variation in the
Pectoralis major is innervated by the medial and lateral number of costal attachments of this head, with inclusion of
pectoral nerves (Standring 2016). the first, seventh, and/or eighth costal cartilages (Macalister
1875; Snosek and Loukas 2016; Standring 2016).
Comparative Anatomy Pectoralis major may be continuous with deltoid, exter-
Pectoralis major has a similar typical presentation in the nal oblique, or rectus abdominis (Macalister 1875; Bergman
apes, with clavicular, sternocostal, and abdominal heads et al. 1988). It may send an accessory slip to biceps brachii or
(Gibbs 1999; Diogo et al. 2010, 2012, 2013a,b, 2017). In latissimus dorsi (Macalister 1875; Knott 1883a; Mori 1964).
gibbons and gorillas, pectoralis major is often blended An accessory head of pectoralis major has been observed
with biceps brachii (Deniker 1885; Kohlbrügge 1890– originating from serratus anterior (Loukas et al. 2006b).
1892; Howell and Straus 1932; Stewart 1936; Raven 1950; Hammad and Mohamed (2006) observed an accessory
Preuschoft 1965; Michilsens et al. 2009; Diogo et al. 2012). tendon originating from the lateral portion of pectoralis
FIGURE 3.5 Anterior view of the pectoralis muscles and pectoral region variations.
major that had an attachment to the shoulder joint capsule. 2018), deltoid was fused with pectoralis major bilaterally,
Tubbs et al. (2008a) also note an insertion of pectoralis forming the deltopectoral complex. In the neonate, the pec-
major onto the shoulder joint capsule. toral portion did not insert directly onto the humerus but
The fbers of the left and right pectoralis major muscles instead was continuous with biceps brachii (Aziz 1979). In
may fuse over the sternum (Macalister 1875; Snosek and the fetus, the abdominal portion of pectoralis major on the
Loukas 2016; Standring 2016). In rare cases, the entire pecto- left side was absent (Alghamdi et al. 2018). The fascia on
ralis major muscle may be absent or even doubled (Macalister the deep surface of pectoralis major was fused with the fas-
1867a, 1875; Bergman et al. 1988; Upasna et al. 2015; Snosek cia of the short head of biceps brachii bilaterally.
and Loukas 2016). Pectoralis major is associated with super-
numerary muscles including sternalis, pectoralis quartus, Prevalence
pectorodorsalis, and chondroepitrochlearis (see these entries). In their literature review, Smith et al. (2015) found that
pectoralis major was fused with deltoideus in 21 out of 26
Prevalence individuals with trisomy 18 (80.8%).
In a study of 40 cadavers, Haładaj et al. (2019) observe the
fusion of the clavicular head of pectoralis major with del- Clinical Implications
toideus in two male cadavers (5%), and hypotrophy of the Haładaj et al. (2019) state that knowledge of the numerous
clavicular head in one female (2.5%). In a literature review anatomical variations of pectoralis major is important for
of 186 cases of congenital muscle absences, Bing (1902) planning and conducting surgeries in this region.
reported that the sternocostal part of pectoralis major is
defective or absent more than any other muscle (defcient
in 102 cases). Upasna et al. (2015) report the absence of
pectOralis MinOr (figure 3.5)
the pectoralis major in 1 out of 50 cases (2%), and Lee and See also: Pectoralis minimus, Costocoracoideus,
Chun (1991) report its absence in one case. Upasna et al. Sternoclavicularis, Tensor semivaginae articulationis
(2015) cite notes from Bergman and colleagues that sug- humeroscapularis
gest that this muscle was absent in 3 of 15,000 cases in
one study, the muscle was absent in 5 of 54,000 cases in Synonyms
another study. Mori (1964) reports a slip connecting pec- N/A
toralis major to biceps brachii in 15 out of 320 arms from
160 Japanese individuals (4.7%). Mori (1964) also reports Typical Presentation
a slip connecting pectoralis major to latissimus dorsi in 12 Description
out of 320 arms from 160 Japanese individuals (3.75%), but Pectoralis minor lies deep to pectoralis major (Standring
this slip is not designated as an axillary arch, simply an 2016). It originates from the third, fourth, and ffth ribs and
“abnormous slip.” from the intercostal fascia and has a tendinous insertion
into the coracoid process (Standring 2016).
Anomalies
Description Innervation
Poland syndrome, a rare congenital anomaly, can lead to Pectoralis minor is innervated by the medial and lateral
the absence of the sternocostal head, defciencies of the cla- pectoral nerves (Standring 2016).
vicular head, or the absence of the entire pectoralis major
muscle (Cingel et al. 2013; Snosek and Loukas 2016). Comparative Anatomy
In a fetus with craniorachischisis, dorsoepitrochlearis Pectoralis minor has a similar typical presentation in the
was fused to the inner side of the inferior band (costo- apes, extending from the ribs to the coracoid process, with
humeralis) of pectoralis major on the right side of the body similar variability in costal origins (Gratiolet and Alix
(Alghamdi et al. 2017). On the left side of the body, pec- 1866; Deniker 1885; Kohlbrügge 1890–1892; Hepburn
toralis major was fused with deltoideus, and an additional 1892; Beddard 1893; Primrose 1899, 1900; Sonntag 1923,
bundle was present on the inner side of this muscle. This 1924; Sullivan and Osgood 1927; Stewart 1936; Miller
bundle originated from the anterior capsule of the shoulder 1952; Kallner 1956; Gibbs 1999; Michilsens et al. 2009;
joint and attached distally to the crest of the greater tuber- Gibbs 1999; Diogo et al. 2010, 2012, 2013a,b, 2017).
cle of the humerus (Alghamdi et al. 2017). Mieden (1982) The costal attachments may include the frst rib in gib-
describes an infant with median cleft lip, hypotelorism, bons, common chimpanzees, and bonobos (Hepburn 1892;
and alobar holoprosencephaly. On the right side, pectoralis Michilsens et al. 2009; MacDowell 1910, Diogo et al. 2012,
major split into superfcial and deep layers at the costal car- 2013a, 2017). In common chimpanzees, the insertion into
tilages (Mieden 1982). the coracoid may be absent (Champneys 1872; Chapman
In a neonate with trisomy 13, the right pectoralis major 1879; Sutton 1883; Hepburn 1892; MacDowell 1910;
was divided into clavicular, sternal, and costal portions Ziegler 1964; Swindler and Wood 1973; Diogo et al. 2013a,
(Aziz 1980). In two neonates with trisomy 18 (Aziz 1979, 2017). It may have an insertion into the clavicle in gibbons
1981) and in a female fetus with trisomy 18 (Alghamdi et al. (Hepburn 1892; Sonntag 1924, Stewart 1936; Gibbs 1999;
Michilsens et al. 2009; Gibbs 1999; Diogo et al. 2012). It Alghamdi et al. (2017) note that on the left side of the body
may have additional attachments to the proximal humerus of a fetus with craniorachischisis, pectoralis minor had a nor-
or glenohumeral joint capsule in orangutans and common mal insertion but sent an abnormal slip connecting it to pec-
chimpanzees (Diogo et al. 2013a,b). toralis major. On the right side of the body, pectoralis minor
originated from ribs two through four and inserted onto the
Variations lesser tubercle of the humerus (Alghamdi et al. 2017).
Description In a neonate with trisomy 13, pectoralis minor was absent
Pectoralis minor may arise from the second to fourth ribs on the left side (Aziz 1980). In a neonate with trisomy 18, the
(Macalister 1875; Standring 2016). Other origin patterns left pectoralis minor was divided into two fascicles (Aziz
include the second to ffth ribs, second to fourth ribs, third 1979). One extended from the ffth rib to the coracoid pro-
to fourth ribs, or third to ffth ribs (Macalister 1875; Mori cess and the other extended from the fourth rib to the cora-
1964; Standring 2016), or only from the ffth rib in a rare coid process. In a female fetus with trisomy 18, the insertion
case (Turan-Özdemir and Cankur 2004). An origin from of pectoralis minor on the right side extended inferiorly to
the sixth rib has been observed (Macalister 1875; Bergman the fascia of coracobrachialis (Alghamdi et al. 2018).
et al. 1988). The tendon may cross the coracoid process into
the coraco-acromial ligament, or even extend beyond the Prevalence
ligament and attach to shoulder joint capsule or the humerus In their literature review, Smith et al. (2015) found that
(Macalister 1875; Knott 1883a; Bergman et al. 1988, Tubbs pectoralis minor was absent in 3 out of 24 individuals with
et al. 2005a; Uzel et al. 2008; Snosek and Loukas 2016; trisomy 13 (12.5%) and in 1 out of 26 trisomy 18 individu-
Standring 2016). Other potential attachments include the als (3.8%).
clavicle, costocoracoid membrane, costohumeral ligament,
or the supraspinatus tendon (Macalister 1875; Uzel et al. Clinical Implications
2008; Snosek and Loukas 2016). Aberrant lateral insertions (ectopic insertions) of the pecto-
Pectoralis minor may be connected with subclavius, ralis minor tendon beyond the coracoid process may lead to
pectoralis major, biceps brachii, latissimus dorsi, or cora- pain, clicking, or restricted range of motion (Moineau et al.
cobrachialis (Macalister 1875; Bergman et al. 1988; Snosek 2008; Low and Tan 2010; Pandey et al. 2016; Snosek and
and Loukas 2016). Pectoralis minor may be absent unilat- Loukas 2016; Ruiz Santiago et al. 2019).
erally or bilaterally, especially when pectoralis major is
absent (Macalister 1875; Bergman et al. 1988, Snosek and chOnDrOepitrOchlearis (figure 3.4)
Loukas 2016; Standring 2016). It may also be doubled when
pectoralis major is doubled (Macalister 1875). Variations See also: Pectoralis major, Pectoralis quartus
of this muscle also include slips that are defned as discrete
muscles including pectoralis minimus, costocoracoideus, Synonyms
sternoclavicularis, and tensor semivaginae articulationis This muscle may also be referred to as costoepitrochlearis,
humeroscapularis (see these entries). chondrohumeralis (Bergman et al. 1988), thoracoepicondy-
laris, chondrofascialis, chondrobrachialis, costohumeralis
Prevalence (Snosek and Loukas 2016), and chondroepicondylaris sensu
Macalister (1875) notes that the tendon of pectoralis minor (Palagama et al. 2016).
was prolonged over the coracoid process in 17 out of 106
subjects (16.04%). Upasna et al. (2015) report the absence of Typical Presentation
the pectoralis minor in 1 out of 50 cases (2%). Mori (1964) This muscle is only present as a variation or anomaly.
found that pectoralis minor originated from the frst three
ribs in 2 out of 326 cases (0.6%), ribs two through four in Comparative Anatomy
46 cases (14.1%), ribs two through fve in 115 cases (35.3%), Chondroepitrochlearis is present in rare cases in com-
ribs three and four in nine cases (2.8%), ribs three through mon chimpanzees and bonobos, originating from the ribs
fve in 153 cases (46.9%), and ribs three through six in one and inserting onto the proximal humerus at the crest of
case (0.3%). The superior most attachment was to rib one in the greater tubercle (MacDowell 1910; Diogo et al. 2013a,
two cases (0.6%), rib two in 161 cases (49.4%), and rib three 2017). Palagama et al. (2016) suggest that chondroepitroch-
in 163 cases (50%). The inferior most attachment was to rib learis, acting as an insertion point for pectoralis major on
three in two cases (0.6%), rib four in 55 cases (16.9%), rib the medial epicondyle, would impede shoulder extension
fve in 268 cases (82.2%), and rib six in one case (0.3%). in suspensory primates, likely contributing to why it is rare
in humans and our closest relatives.
Anomalies
Poland syndrome, a rare congenital anomaly, can cause the Description
underdevelopment or absence of pectoralis minor (Snosek Chondroepitrochlearis is considered a variant of pec-
and Loukas 2016; Standring 2016; Petleshkova et al. 2019). toralis major. It originates from one or more ribs, the
chondrocostal junction of pectoralis major, or the external Clinical Implications

oblique aponeurosis (Macalister 1875; Knott 1880, 1883a; Due to its position in the axilla, the presence of chondro-
Bergman et al. 1988; Snosek and Loukas 2016). It courses epitrochlearis may interfere with axillary lymphadenec-
through the axilla to insert onto the medial intermuscular tomy (Natsis et al. 2010, 2012; Palagama et al. 2016).
septum or medial epicondyle of the humerus (Macalister Furthermore, its presence can limit abduction at the
1875; Knott 1880, 1883a; Bergman et al. 1988; Snosek and shoulder joint (Lin 1988; Redler et al. 2012; Palagama
Loukas 2016). Macalister (1875) found an insertion into et al. 2016) or lead to entrapment of the ulnar nerve
the brachial aponeurosis. Standring (2016) describes chon- (Spinner et al. 1991).
droepitrochlearis as an extra head of pectoralis major that
originates deep to the sternal and abdominal heads from
the ffth through seventh ribs and their associated costal sternOclavicularis (figure 3.5)
cartilages. It can be present unilaterally or bilaterally (Lin
1988; Snosek and Loukas 2016). It may have two bellies See also: Pectoralis major, Pectoralis minor
(Macalister 1875). It frequently occurs with pectorodorsalis
(Macalister 1875; Chiba et al. 1983; Lin 1988; Spinner et al. Synonyms
1991; Snosek and Loukas 2016; Palagama et al. 2016). Sternoclavicularis is also referred to as preclavicularis
Macalister (1875) suggests that chondroepitrochlearis medialis, sternoclavicularis anterior, sternoclavicularis
may be a variant of pectoralis quartus (see the entry for anticus, or supraclavicularis (Gruber 1860; Macalister 1875;
this muscle). In a cadaver examined by Carroll et al. (2019), Knott 1883a; Eisler 1912; Snosek and Loukas 2016).
chondroepitrochlearis was present along with a third head
of biceps brachii. These authors suggest that chondro- Typical Presentation
epitrochlearis may be a remnant of panniculus carnosus
(Carroll et al. 2019). Barcia and Genovés (2009) consider
chondrofascialis to be a separate muscle from, not a syn-
Comparative Anatomy
onym of, chondroepitrochlearis, as they argue that chon-
drofascialis inserts into the fascia of the arm without direct Sternoclavicularis has been observed in one orangutan
connections to pectoralis major or the humerus and should (Michaelis 1903) but has not been recorded in any other
thus be considered intermediate in form between pectoralis orangutan dissections, nor in dissections of the other apes
quartus and chondroepitrochlearis. (Diogo et al. 2010, 2012, 2013a,b, 2017).
Variations
Innervation
Description
Chondroepitrochlearis is innervated by the medial pectoral
Sternoclavicularis is considered a variant of pectoralis
nerve (Barcia and Genovés 2009; Palagama et al. 2016).
major and pectoralis minor. It may be present when pec-
Prevalence toralis major is defcient (Macalister 1875; Snosek and
Loukas 2016). It can originate from the manubrium,
Flaherty et al. (1999) observed chondroepitrochlearis
the capsule of the sternoclavicular joint, the anterior
in only 1 out of 200 cadavers over the course of 20 years
sternoclavicular ligament, or the frst or second costal
(0.5%). Natsis et al. (2012) report a prevalence of this mus-
cartilage (Macalister 1875; Knott 1883a; Mori 1964;
cle in 1 out of 119 cadavers (0.84%), based on dissections
Bergman et al. 1988; Sakuma et al. 2007; Sontakke et al.
done by them and Natsis et al. (2010).
2013; Snosek and Loukas 2016). It may insert into the
middle or distal portions of the clavicle (Macalister 1875;
Anomalies Knott 1883a; Mori 1964; Smith et al. 2015; Bergman
Description et al. 1988; Snosek and Loukas 2016). When sterno-
In a neonate with trisomy 13, Aziz (1980) describes chon- clavicularis is present bilaterally, a tendinous band can
droepitrochlearis as a derivative of the sternal portion of connect the muscles, which is referred to as musculus
pectoralis major that became aponeurotic near its humeral interclavicularis (Knott 1883a) or interclavicularis anti-
insertion. The aponeurosis was divided into superfcial and cus digastricus (Sakuma et al. 2007; Snosek and Loukas
deep faps. The superfcial fap attached to the lateral crest 2016). It attaches onto the manubrium via a tendinous
of the bicipital groove, and then became muscular along the intersection and connects the two clavicles (Bergman
humeral shaft. This muscle, chondroepitrochlearis, inserted et al. 1988). If sternoclavicularis inserts into the cora-
onto the medial epicondyle. coid process, it may be referred to as coraco-clavicularis
anticus (Knott 1883a).
Prevalence
In their literature review, Smith et al. (2015) found that Innervation
chondroepitrochlearis was present in 5 out of 24 individu- Sternoclavicularis is innervated by the lateral pectoral
als with trisomy 13 (20.8%). nerve (Sakuma et al. 2007; Sontakke et al. 2013).
Prevalence Souza et al. 2019). This muscle can also arise from the
Gruber (1860) suggests it is present in about 3% of bodies. rectus sheath (Bascho 1906; Bonastre et al. 2002; Snosek
Mori (1964) found this muscle in 2 out of 1050 cadavers and Loukas 2016). Pectoralis quartus can also insert onto
(0.2%). the fascia of the arm, the short head of the biceps, the
bicipital groove of the humerus, or the coracoid process
(Birmingham 1889; Arican et al. 2006; Snosek and Loukas
Anomalies 2016; Potau et al. 2018). Hunt (2017) reports a case in which
Description a conjoined tendon of pectoralis major and pectoralis quar-
As in karyotypically normal humans, sternoclavicularis tus passed superiorly through the intertubercular groove to
originates from the anterior surface of the manubrium and insert over the glenohumeral joint capsule. Pectoralis quar-
capsule of the sternoclavicular joint and inserts onto the tus can be connected to the axillary arch or sternalis when
anterior surface of the clavicle (Smith et al. 2015). Barash these muscles are present (Knott 1883a; Bergman et al.
et al. (1970) report the unilateral presence of sternoclavicu- 1988; Bonastre et al. 2002; Snosek and Loukas 2016).
laris in a trisomy-18 infant. Pectoralis quartus is similar in presentation to a muscle
that some refer to as pectoralis tertius, which runs between
Prevalence the lower ribs and the humerus or coracoid process (Snosek
In their literature review, Smith et al. (2015) found that and Loukas 2016). This muscle appears to be equivalent to
sternoclavicularis was present in 1 out of 17 individuals the xiphihumeralis muscle that runs between the xiphoid
with trisomy 18 (5.9%). and coracoid process in rodents (Snosek and Loukas 2016;
Diogo et al. 2018). Pectoralis quartus is also similar in
Clinical Implications presentation to costocoracoideus (see the entry for this
Sontakke et al. (2013) note that sternoclavicularis muscle muscle).
could be mistaken for a mass or tumor in medical imaging,
and Sakuma et al. (2007) suggest that it could interfere with Innervation
subclavian-vein catheter insertion. Pectoralis quartus is innervated by the medial pecto-
ral nerve (Birmingham 1889; Snosek and Loukas 2016).
Arican et al. (2006) observe that this muscle is innervated
pectOralis quartus (figure 3.5) by the fourth intercostal nerve in one specimen.
See also: Pectoralis major
Prevalence
Synonyms Pectoralis quartus has been reported to have a prevalence
This muscle is also referred to as pectoralis abdominis between 2.8% and 11%–16% (Diogo and Wood 2012a;
(Potau et al. 2018) or pectoralis abdominalis (Dunlap et al. Testut 1884; Wagenseil 1937; Bonastre et al. 2002; Snosek
1986). and Loukas 2016; Hunt 2017; Cardoso Souza et al. 2019).
Anomalies
Description
Comparative Anatomy As in karyotypically normal humans, pectoralis quartus
In one gibbon, pectoralis quartus originated from between may arise below pectoralis major and extend to the humerus
the abdominal head of pectoralis major and the rectus (Howell and Straus 1933; Dunlap et al. 1986).
sheath and inserted into the humerus (Van den Broek 1909;
Diogo et al. 2012). Pectoralis quartus (pectoralis abdomi- Prevalence
nis) has been described in gorillas, but Diogo et al. (2010) In their literature review, Smith et al. (2015) found that pec-
argue that these structures correspond to the abdominal toralis quartus (“pectoralis abdominalis”) was present in
head of pectoralis major. one out of seven individuals with trisomy 21 (14.3%).
Variations Clinical Implications

Hunt (2017) suggests that the presence of pectoralis quartus
Description could obscure the long head of the biceps during imaging
Pectoralis quartus is a variant of pectoralis major, extending studies, leading to the misdiagnosis of pathology to this
laterally along the inferior border of this muscle (Bergman structure. Furthermore, Hunt (2017) notes that pain attrib-
et al. 1988; Arican et al. 2006; Snosek and Loukas 2016). uted to the long head of the biceps could actually be due
It usually arises near the costochondral junction of ribs fve to pectoralis quartus. Bonastre et al. (2002) state that pec-
and six, crosses the axilla, and inserts on the tendon of pectoralis quartus can contribute to axillary thrombosis and
toralis major (Macalister 1875; Knott 1883a; Birmingham may complicate the removal of lymph nodes during axil-
1889; Bergman et al. 1988; Arican et al. 2006; Cardoso lary lymphadenectomy.
pectOrODOrsalis (figure 3.4) Anomalies

See also: Pectoralis major, Latissimus dorsi Description
Aziz (1980, 1981) reports the presence of pectorodorsalis,
Synonyms a muscle bundle connecting latissimus dorsi to pectoralis
Pectorodorsalis is also referred to as the axillary arch, ach- major, in three trisomy 13 neonates. In a female fetus with
selbogen, Langer’s arch, Langer’s muscle, axillopectoral trisomy 18, Langer’s axillary arch was present on the left
muscle, and arcus axillans (Langer 1846; Bergman et al. side, extending from the lateral side of latissimus dorsi to
1988; Bakkum and Miller 2016; Rai et al. 2018). the inferior aspect of pectoralis major near its insertion onto
the humerus (Alghamdi et al. 2018). In a fetus with trisomy
Typical Presentation 18, Dunlap et al. (1986) describe a pectorodorsalis that pres-
This muscle is only present as a variation or anomaly. ents as a tendinous insertion of panniculus carnosus into the
latissimus dorsi tendon. However, they do not include this
Comparative Anatomy as a case of pectorodorsalis in their summary tables.
Dorsoepitrochlearis is considered to be the manifestation
Prevalence
of axillary arch muscles in the apes. Pectorodorsalis is
therefore not present in the apes (Diogo et al. 2010, 2012, In their literature review, Smith et al. (2015) found that pec-
2013a,b, 2017). torodorsalis was present in 16 out of 24 individuals with
trisomy 13 (66.7%).
Variations
Description
Ucerler et al. (2005) note that the presence of pectorodor-
Pectorodorsalis is present as a slip of muscle that crosses salis can lead to entrapment of the axillary vein or median
the axilla and connects latissimus dorsi to pectoralis major nerve, diffculty staging lymph nodes in malignancy cases,
or its tendon (Macalister 1875; Mori 1964; Bergman et al. problems with axillary surgery, and shoulder instability
1988; Chavan and Wabale 2014; Bakkum and Miller 2016; (Chavan and Wabale 2014). Chavan and Wabale (2014) also
Standring 2016). It can also attach to the axillary fascia, suggest that the presence of this muscle can lead to neuro-
biceps brachii or its fascia, coracobrachialis or its fas- vascular symptoms in the upper limb or the development of
cia, pectoralis minor, teres major, or the coracoid process upper limb lymph edema after breast surgery. Regarding
(Ramsay 1812; Macalister 1875; Mori 1964; Bergman et al. neurovascular compression, Rai et al. (2018) state that
1988; Bakkum and Miller 2016; Standring 2016). It may be pectorodorsalis can lead to brachial plexus compression,
present bilaterally (Chavan and Wabale 2014). thoracic outlet syndrome, and hyperabduction syndrome.
Innervation
Pectorodorsalis may be innervated by the medial pectoral pectOralis MiniMus (figure 3.5)
nerve or the thoracodorsal nerve (Chavan and Wabale 2014;
Bakkum and Miller 2016). See also: Pectoralis minor
Prevalence Synonyms
Ramsay (1812) observed the presence of this muscle in 1 Pectoralis minimus is also referred to as chondrocoracoi-
out of 30 cadavers (3.3%). Macalister (1875) notes that it deus ventralis, sternocostocoracoidian, sternochondrocora-
has been found by Struthers in 8 out of 105 cases (7.6%) and coideus, or sternochondrocoracoideus ventralis (Snosek
by Wood in 6 out of 106 cases (5.7%). In a review by Khan and Loukas 2016). Sternochondroscapularis is potentially
et al. (2008) they list a prevalence ranging as low as 0.2% another synonym (Aziz 1980).
(Serpell and Baum 1991) to as high as 10% (Clarys et al.
1996). Wagenseil (1927) reports a prevalence of 43.8% in Typical Presentation
a Chinese population. Mori (1964) reports an axillary arch This muscle is only present as a variation or anomaly.
in 52 out of 1050 sides from Japanese cadavers (5%). Mori
(1964) also reports a slip from pectoralis major to coraco- Comparative Anatomy
brachialis in 3 out of 50 arms (6%) and suggests it is a type N/A
of axillary arch. Kasai and Chiba (1977) suggest a preva-
lence of 9.1% in a Japanese population. Miguel et al. (2001) Variations
observe a presence of 3% in Spain. Turgut et al. (2005) Description
provide a prevalence of 1.9% in a Turkish population. A Pectoralis minimus (Gruber) originates from the carti-
prevalence of 3.6% in a Bulgarian population is provided lage of the frst, second, or potentially the third rib, and
by Georgiev et al. (2007). Turki and Adds (2017) found only inserts onto the coracoid process (Macalister 1875; Knott
three axillary arch muscles from the examination of 280 1883a; Bergman et al. 1988; Turgut et al. 2000; Rai et al.
Caucasian cadavers spanning 14 years of dissection (1%). 2008; Soni et al. 2008; Ebenezer and Rathinam 2013;
Khizer Hussain Afroze et al. 2015; Snosek and Loukas Clinical Implications
2016). It may also have origins from the manubrium or The presence of pectoralis minimus can cause impingement
costoclavicular ligament (Macalister 1875; Knott 1883a). or compression of nearby thoracoacromial vessels (Turgut
Turgut et al. (2000) observed insertions into the shoulder et al. 2000; Rai et al. 2008; Ebenezer and Rathinam 2013;
joint capsule, the lateral clavicle, and fascia of subclavius. It Khizer Hussain Afroze et al. 2015; Snosek and Loukas
may be present bilaterally (Turgut et al. 2000). 2016).
Innervation
Pectoralis minimus is innervated by the lateral pectoral
cOstOcOracOiDeus (figure 3.6)
nerve (Soni et al. 2008; Khizer Hussain Afroze et al. 2015).
See also: Pectoralis minor
Prevalence
Khizer Hussain Afroze et al. (2015) observed the presence Synonyms
of this muscle in 3 out of 56 cadavers in India (5.4%). This muscle is also known as chondrocoracoideus (Bergman
et al. 1988).
Anomalies
Description Typical Presentation

Pectoralis minimus arises from the frst rib, superior to pec- This muscle is only present as a variation.
toralis minor, and inserts onto the coracoid process (Smith
et al. 2015). Barash et al. (1970) report the unilateral pres- Comparative Anatomy
ence of pectoralis minimus in a trisomy-18 infant. Aziz Although Jouffroy (1971) states that costocoracoideus may
(1980) reports a “sternochondroscapularis” on the right side be present in some primates, dissections and literature
of a neonate with trisomy 13 that originated from the manu- reviews by Diogo et al. (2010, 2012, 2013a,b, 2017) do not
brium and frst rib and inserted onto the coracoid process fnd costocoracoideus present as a distinct muscle in the
and humeral head. apes. Rather, all these species present a costocoracoid liga-
ment, as in humans, which corresponds to the costocora-
Prevalence coid muscle of monotremes (Diogo et al. 2013a). A distinct
In their literature review, Smith et al. (2015) found that pec- costocoracoideus muscle may be present in common chim-
toralis minimus was present in 9 out of 26 individuals with panzees as a variant, similar to the condition in humans
trisomy 18 (34.6%). (Diogo et al. 2013a).
FIGURE 3.6 Thoracoscapular variations in anterior view.

Variations In this specimen, it originated from the sternum and carti-

Description lage of the frst and second ribs.
Humans typically only have a costocoracoid ligament. Variations
When present as a distinct muscle, costocoracoideus typi-
cally originates via one or more slips from ribs six to eight Description
or the rectus sheath and inserts onto the coracoid process Tensor semivaginae articulationis humeroscapularis (Gruber)
(Macalister 1875; Bergman et al. 1988; Snosek and Loukas originates from one or two of the upper rib cartilages and
2016). It thus has a similar origin and insertion to pecto- from the sternum (Macalister 1875; Knott 1883a; Bergman
ralis quartus (see the entry for this muscle). It may also et al. 1988; Snosek and Loukas 2016). It inserts into the
originate from ribs fve through seven (Macalister 1875). shoulder joint capsule, the fascia deep to deltoid, or the sub-
Wood (1864) reports a “chondro-coracoid muscle” that acromial bursa (Macalister 1875; Huntington 1904; Knott
arose from latissimus dorsi at the tenth rib and inserted 1883a; Bergman et al. 1988; Snosek and Loukas 2016).
into the apex of the coracoid process. Venieratos et al. Snosek and Loukas (2016) note that this muscle may be a
(2017) report a “chondrocoracoideus” arising via three displaced portion of pectoralis major, and it is often present
slips from ribs six, seven, and eight and from the aponeu- when pectoralis major is defcient.
rosis of the external oblique, fusing with the sternocostal
Innervation
head of pectoralis major at the level of rib fve and six, and
fusing with the tendon of the short head of biceps brachii Tensor semivaginae articulationis humeroscapularis is
before insertion onto the coracoid process. Sonne (2020) innervated by the lateral pectoral nerve (Huntington 1904).
reports a “chondrocoracoideus” muscle originating from
Prevalence
the costal cartilage of the sixth rib.
N/A
Innervation
Anomalies
Costocoracoideus is innervated by the medial pectoral
nerve (Venieratos et al. 2017; Sonne 2020). Description
Smith et al. (2015) state that “tensor semivaginae scapulo-
Prevalence humeralis” passes from the manubrium to the humeral head
Sonne (2020) found this muscle in 1 out of 36 cadavers and/or the scapula. This is based on the description from
(2.8%). Ramirez-Castro and Bersu (1978), who describe this muscle
as passing from the frst sternochondral joint to the tendi-
Anomalies nous attachments over the glenohumeral joint capsule. In a
N/A neonate with trisomy 18, tensor semivaginae articulationis
humeroscapularis arose from the costal surfaces of ribs one
Clinical Implications and two, coursed below the clavicle, and ended as an apo-
Venieratos et al. (2017) suggest that “chondrocoracoideus” neurosis over the glenohumeral joint that inserted into the
can potentially compress the axillary artery and branches scapula (Aziz 1979).
of the brachial plexus, contributing to thoracic outlet
syndrome. Prevalence
“tensor semivaginae scapulohumeralis” was present in 1
tensOr seMivaginae articulatiOnis out of 24 individuals with trisomy 13 (4.2%) and 10 out of
huMerOscapularis (figure 3.6) 26 individuals with trisomy 18 (38.5%).
See also: Pectoralis minor Clinical Implications
N/A
Synonyms
Alternative names for this muscle include sternohumeralis
and sternochondrohumeralis (Snosek and Loukas 2016), infraclavicularis (figure 3.7)
and tensor semivaginae scapulohumeralis (Ramirez-Castro
and Bersu, 1978; Smith et al. 2015). Synonyms
N/A
This muscle is only present as a variation or anomaly. Typical Presentation
Comparative Anatomy
Macalister (1871) found tensor semivaginae articulationis Comparative Anatomy
humeroscapularis on the left side of a female chimpanzee. N/A
FIGURE 3.7 Deltoideus and clavicular region variations in anterior view.
Infraclavicularis (Bardeleben) is a rare muscle situated infe- Costoclavicularis is situated deep to the upper part of pec-
rior to the clavicle (Knott 1883a; Snosek and Loukas 2016). toralis minor and inferior to subclavius (Mori 1964; Snosek
It arises from the middle third of the clavicle or the anterior and Loukas 2016). It arises from the second rib and attaches
sternoclavicular ligament and inserts onto the lateral end of to the inferior aspect of the middle portion of the clavicle
the clavicle and deltopectoral fascia (Knott 1883a; Ingalls (Mori 1964; Snosek and Loukas 2016).
1913; Snosek and Loukas 2016; Wehrli et al. 2017). Snosek
and Loukas (2016) note that this muscle may be a displaced Innervation
portion of the clavicular head of pectoralis major. It may be Mori (1964) states that Nishi reports this muscle may be
present bilaterally (Wehrli et al. 2017). innervated by a branch of the anterior thoracic nerve (either
a branch of the medial or lateral pectoral nerve).
Innervation
Infraclavicularis is innervated by the lateral pectoral nerve Prevalence
(Snosek and Loukas 2016). Mori (1964) observed costoclavicularis in 3 out of 367 cases
(0.8%) and reports that Nishi found this muscle in 1 out of
Prevalence 12 cadavers (8.3%).
N/A Anomalies
Anomalies N/A
N/A Clinical Implications
N/A
rOOs banDs/Muscles (figure 3.7)
See also: Scalenus minimus
cOstOclavicularis (figure 3.7)
Synonyms Synonyms
N/A N/A
Comparative Anatomy
Comparative Anatomy To our knowledge, most Roos bands have not been
N/A described in the apes (Diogo et al. 2010, 2012, 2013a,b,
2017). Scalenus minimus muscles corresponding to a type Clinical Implications

5 band have been found in gorillas and bonobos (see the The presence of these bands creates a tunnel through
entry for this muscle). which the T1 nerve root passes, which also compresses the
T1 nerve root against the frst rib, therefore predisposing
Variations individuals with these muscles to thoracic outlet syndrome
Description (Roos 1976; Spears et al. 2011).
Roos bands or muscles are fbro-muscular structures that
originate in the neck, anterior to the T1 root, and insert
onto the frst rib (Roos 1976; Spears et al. 2011; Snosek DeltOiDeus (DeltOiD) (figure 3.7)
and Loukas 2016). Roos (1976) classifes seven types of Synonyms
presentations for these bands and recorded their preva- N/A
lence in 241 patients that underwent operations for tho-
racic outlet syndrome and in 58 thoracic outlet dissections Typical Presentation
from 29 cadavers. A type 1 band is classifed as a tough
Description
fbrous ligament extending from the top of an incomplete
cervical rib to the midsection of the frst rib. A type 2 band Deltoid originates from the lateral third of the clavicle, the
extends from an elongated seventh cervical rib vertebra to acromion, and the crest of the scapular spine (Standring
the frst rib. A type 3 band is the smallest and deepest and 2016). It inserts via a tendon onto the deltoid tubercle on the
presents as a muscular band extending between the neck humeral midshaft (Standring 2016).
of the frst rib to the inner surface of the posterior curve of Innervation
this rib. A type 4 band is a muscle connection between the
Deltoid is innervated by the axillary nerve (Standring 2016).
anterior and middle scalene muscles that attach to the frst
rib. A type 5 band is the scalenus minimus muscle attach- Comparative Anatomy
ing onto the frst rib between the subclavian artery and the
Deltoid has a similar typical presentation in the apes, with cla-
brachial plexus. A type 6 band is the scalenus minimus
vicular, acromial, and spinal portions that converge to insert
muscle attaching to the cupula of the pleura beneath the
into the humerus (Gratiolet and Alix 1866; Primrose 1899,
frst rib. A type 7 band extends between the middle sca-
1900; Sonntag 1923, 1924; Sullivan and Osgood 1927; Miller
lene muscle to the costal cartilage of the frst rib. Spears
1952; Kallner 1956; Swindler and Wood 1973; Gibbs 1999;
et al. (2011) list three other band types, including a band
Diogo et al. 2010, 2012, 2013a,b, 2017). The insertion of del-
extending from the anterior scalene muscle to subclavius
toid into the humerus in gibbons is more distal than in the other
and the costal cartilage (type 8), a band running along
apes (Diogo et al. 2012). In bonobos, deltoid may be fused with
the posterior inner surface of the frst rib (type 9), and a
pectoralis major or infraspinatus (Diogo et al. 2017).
double fbrous band that extends over the cupula of the
lung (type 10). Variations
Description
Innervation Each part of the deltoid, or the entire muscle, can be reduced
N/A or absent (Otto 1830; Gruber 1872; Macalister 1875; Bing
1902; Bergman et al. 1988; Lamb 2016). The muscle sheet of
Prevalence the deltoid can split into multiple parts (Bergman et al. 1988),
Out of 241 patients that underwent surgery for thoracic out- including a variation in which the posterior-most fbers are
let syndrome, Roos (1976) found type 1 bands in 18 cases separated from the rest of the muscle sheet by a layer of fas-
(7.5%), type 2 bands in 10 cases (4.1%), type 3 bands in 146 cia (Kayikçioglu et al. 1993; Kamburoğlu et al. 2008; Lamb
cases (60.6%), type 4 bands in 13 cases (5.4%), type 5 bands 2016). The muscle may also split into numerous fascicles
in 40 cases (16.6%), type 6 bands in fve cases (2.1%), type (Macalister 1875; Lamb 2016). The completeness of the divi-
7 bands in two cases (0.8%), and no bands were present in sion between the clavicular, acromial, and spinous portions
seven cases (2.9%). Out of 58 thoracic outlet dissections can vary (Macalister 1875; Mori 1964; Lamb 2016).
in 29 cadavers, Roos (1976) found type 3 bands present in Deltoid can fuse with pectoralis major (deltopectoral
10 cases (17.2%), a type 4 band present in one case (1.7%), complex, see anomalous description below), or may have
type 5 bands present in eight cases (13.8%), and no bands connections to trapezius, infraspinatus, supraspinatus, latis-
were present in 39 cases (67.2%). Spears et al. (2011) found simus dorsi, teres major, biceps brachii, brachialis, brachio-
type 3 bands in 29 out 70 cadavers (41.4%) and in 35 out of radialis, the infraspinous fascia, or the lateral scapular border
100 (35%) frst ribs from these cadavers. Type 5 bands were (Macalister 1875; Mori 1964; Bergman et al. 1988; Lamb
found in 11 ribs (11%). 2016; Standring 2016). Acromioclavicularis or prae-clavicu-
laris lateralis extends between the acromion and the clavicular
Anomalies portion of the deltoid (Gruber 1865; Macalister 1875; Knott
N/A 1883a; Lamb 2016). Basiodeltoideus or fasciculus infraspinata
deltoideus (Gruber) extends from the infraspinatus fascia or fetus with trisomy 18 (Alghamdi et al. 2018), deltoid was
axillary border of the scapula to the lower fbers of the deltoid fused with pectoralis major bilaterally, forming the deltopec-
(Knott 1883a; Lamb 2016). Costodeltoideus (Calori) refers to toral complex. Alghamdi et al. (2017) note that in a fetus with
a slip that originates from the scapula between teres minor craniorachischisis, a deltopectoral complex was present bilat-
and infraspinatus, or between teres minor and teres major, erally. In a neonate with trisomy 13, deltoid was divided into
that connects deltoid to the clavicle or to the acromion pro- clavicular, acromial, and spinous portions (Aziz 1980).
cess (Macalister 1875; Knott 1883a; Bergman et al. 1988;
Prevalence
Lamb 2016). Fraser et al. (2014) report an accessory slip con-
necting deltoid with teres minor that they do not believe cor- In their literature review, Smith et al. (2015) found a bilateral
responds to costodeltoideus. Infraspinatohumeralis is a slip deltopectoral complex, in which there was a complete fusion of
that originates from the scapula and ends in the fascia of the deltoid and pectoralis major, in 21 out of 26 fetuses/neonates
arm (Calori 1867; Bergman et al. 1988; Lamb 2016). with trisomy 18 (80.8%). The deltoid was divided into three
distinct parts in 1 out of 24 individuals with trisomy 13 (4.2%).
Prevalence
Mori (1964) studied the deltoid in 50 arms. The acromial
Understanding variations in deltoid anatomy and insertions
portion was fully separated from the rest of the muscle
is important for successful shoulder surgery (Kamburoğlu
in 12 arms (24%), partially separated from the rest of the
et al. 2008; Fraser et al. 2014).
muscle in 19 arms (38%), and not separated from the rest
of the muscle in 19 arms (38%). The clavicular portion was
fully separated from the rest of the muscle in two arms infraspinatus (figure 3.8)
(4%), partially separated from the rest of the muscle in two
Synonyms
arms (4%), and not separated from the rest of the muscle in
46 arms (92%). The spinous portion originated from only N/A
the scapular spine in 16 arms (32%), from the spine and
the infraspinous fascia in 31 arms (62%), and from only the
infraspinous fascia in three arms (6%). Description
Infraspinatus originates from the infraspinous fossa and
Anomalies infraspinous fascia and has a tendinous insertion onto the
Description greater tubercle of the humerus (Standring 2016).
Macalister (1875) states that Haller found deltoid with an
Innervation
insertion into brachialis into an abnormal individual. In two
neonates with trisomy 18 (Aziz 1979, 1981) and in a female Infraspinatus is innervated by the suprascapular nerve
(Standring 2016).
FIGURE 3.8 Scapulohumeral muscles in posterior view.


Infraspinatus has a similar typical presentation in the Variant bundles of infraspinatus may interfere with suc-
apes, extending from the infraspinous fossa and fas- cessful posterior portal placement for shoulder arthroscopy
cia to the greater tubercle of the humerus and often (Ashaolu et al. 2015).
also to the capsule of the glenohumeral joint (Beddard
1893; Miller 1952; Gibbs 1999; Diogo et al. 2010, 2012,
supraspinatus (figure 3.8)
2013a,b, 2017).
Synonyms
Variations N/A
Description
Infraspinatus can be completely absent as a rare variation Typical Presentation
(Bing 1902; Lamb 2016). Infraspinatus can be fused with Description
teres minor (Macalister 1875; Knott 1883a; Aasar 1947; Supraspinatus originates from the supraspinous fossa and
Mori 1964; Bergman et al. 1988; Lamb 2016; Standring the supraspinous fascia and has a tendinous insertion into
2016). It may also fuse with the insertion of supraspinatus or the greater tubercle of the humerus (Standring 2016).
connect with deltoid (Macalister 1875; Knott 1883a; Lamb
2016; Standring 2016). Infraspinatus may be bilaminar Innervation
(Macalister 1875). Scapulohumeralis digastricus singularis Supraspinatus is innervated by the suprascapular nerve
of Gruber refers to an accessory muscle that originates from (Standring 2016).
the acromion and scapular spine, courses between deltoid
and infraspinatus, and inserts into the humerus (Macalister Comparative Anatomy
1875; Lamb 2016).
Supraspinatus has a similar typical presentation in the
Two other accessory fascicles have been named; infra-
apes, extending from the supraspinous fossa and fascia to
spinatus minor (see below) and infraspinatus superfcialis
the greater tubercle of the humerus and often also to the
extends from the scapular spine to the greater tubercle of
capsule of the glenohumeral joint (Gibbs 1999; Diogo et al.
the humerus (Knott 1883a; Aasar 1947; Bergman et al.
2010, 2012, 2013a,b, 2017). In orangutans, supraspinatus
1988; Lamb 2016). Ashaolu et al. (2015) report the pres-
may have additional origins from the scapular spine and/or
ence of infraspinatus minor together with a teres minor
acromion and may also be fused with infraspinatus (Gibbs
that partially divided into three bundles upon inser-
1999; Diogo et al. 2013b). In common chimpanzees, it may
tion onto the humerus on the left side of a Nigerian
have an additional insertion into the acromioclavicular liga-
male cadaver. The authors’ description of infraspinatus
ments (Gratiolet and Alix 1866).
minor—arising inferior to infraspinatus “major” and
extending between the lateral aspect of the infraspinous
Variations
fossa and the greater tubercle of the humerus—seems to
correspond to the defnition of infraspinatus superfcialis Description
provided by Bergman et al. (1988). Typically, infraspi- Macalister (1875), Bergman et al. (1988), and Lamb (2016)
natus minor is defned as the superior or upper part of a note that the presentation of supraspinatus is largely similar
divided infraspinatus (Aasar 1947; Knott 1883a; Bergman among humans. Supraspinatus may be absent in rare cases
et al. 1988; Lamb 2016). (Bing 1902). Other rare variations of supraspinatus include
a division of the muscle belly into two portions, connec-
Prevalence tions to the tendons of pectoralis minor, pectoralis major,
Mori (1964) found infraspinatus fused with teres minor in or subscapularis, and an attachment to a tendinous lamella
10 out of 50 arms (20%). below pectoralis major (Macalister 1875; Le Double 1897;
Bergman et al. 1988; Uzel et al. 2008; Lamb 2016). It
Anomalies may also receive a slip from the suprascapular ligament
Description (Macalister 1875). Supraspinatus can also blend with the
Poland syndrome can cause the underdevelopment or insertion of infraspinatus (Lamb 2016).
absence of infraspinatus (Kutluk and Metin 2017). In a
Prevalence
fetus with craniorachischisis, infraspinatus was fused with
teres minor on both the left and right sides (Alghamdi et al. N/A
2017). This fusion was also observed bilaterally in a fetus
Anomalies
with trisomy 18 (Alghamdi et al. 2018).
Description
Prevalence Poland syndrome can cause the underdevelopment or
N/A absence of supraspinatus (Kutluk and Metin 2017).
N/A Mori (1964) found teres minor fused with infraspinatus in
10 out of 50 arms (20%) and absent in 2 arms (4%). Sato
Clinical Implications (1968c) found teres minor absent in 33 out of 346 sides
N/A (9.5%) in Kyushu-Japanese males and absent in 27 of 214
sides (12.6%) in females. Teres minor was partially fused
with infraspinatus in 74 out of 268 sides in males (27.6%)
teres MinOr (figure 3.8)
and in 54 out of 166 sides in females (32.5%).
Synonyms
N/A Anomalies
Description
Teres minor was fused bilaterally with infraspinatus in both
Description a fetus with craniorachischisis (Alghamdi et al. 2017) and a
Teres minor originates from the lateral margin of the scap- fetus with trisomy 18 (Alghamdi et al. 2018).
ula and inserts via a tendon onto the greater tubercle of the
humerus (Standring 2016). Its insertion blends with the cap- Prevalence
sule of the shoulder joint (Standring 2016). N/A

Teres minor is innervated by the axillary nerve (Standring Variable slips of teres minor may be surrounded by vascu-
2016). lar structures (Ashaolu et al. 2015). In their case, a branch
of the posterior circumfex artery lied between the supe-
Comparative Anatomy
rior and middle bundles of the teres minor insertion, thus
Teres minor has a similar typical presentation in the apes, having the potential to complicate surgery in that region
extending from the lateral border of the scapula to the (Ashaolu et al. 2015).
greater tubercle of the humerus (Gibbs 1999; Diogo et al.
2010, 2012, 2013a,b, 2017). The insertion often extends
to the shaft of the humerus in all apes except orangutans teres MajOr (figure 3.8)
(Gratiolet and Alix 1866; Sonntag 1924; Raven 1950; Synonyms
Swindler and Wood 1973; Gibbs 1999; Diogo et al. 2010,
N/A
2012, 2013a,b, 2017). Teres minor may be blended with
infraspinatus in gibbons, gorillas, and orangutans (Gibbs
1999; Diogo et al. 2012). It may have an additional inser-
tion into the shoulder joint capsule in bonobos (Miller 1952; Description
Gibbs 1999; Diogo et al. 2017). Teres major originates from the inferior scapular angle and
inserts via a tendon onto the intertubercular sulcus of the
Variations humerus (Standring 2016).
Teres minor may be absent entirely (Macalister 1875; Knott Teres major is innervated by the lower subscapular nerve
1883a; Mori 1964; Sato 1968c; Bergman et al. 1988; Lamb (Standring 2016).
2016). A common variation of teres minor is fusion with
infraspinatus (Macalister 1875; Knott 1883a; Mori 1964; Comparative Anatomy
Sato 1968c; Bergman et al. 1988; Lamb 2016; Standring Teres major has a similar typical presentation in the apes,
2016). Teres minor may send a slip to deltoideus (Aasar extending from the lateral border and inferior angle of the
1947; Lamb 2016). There may be an insertion onto the sur- scapula to the proximal humerus (Gibbs 1999; Diogo et al.
gical neck of the humerus (Jain et al. 2012; Lamb 2016). 2010, 2012, 2013a,b, 2017). Fusion with the tendon of latis-
Ashaolu et al. (2015) report the presence of a teres minor simus dorsi is observed as a common variant in all apes
that partially divided into three bundles upon insertion onto (Gratiolet and Alix 1866; Champneys 1872; Barnard 1875;
the humerus on the left side of a Nigerian male cadaver. Kohlbrügge 1890–1892; Hepburn 1892; Beddard 1893;
The superior portion inserted onto the posterior facet of Primrose 1899, 1900; Sonntag 1923, 1924; Sullivan and
the greater tubercle, the middle portion inserted onto the Osgood 1927; Stewart 1936; Miller 1952; Kallner 1956;
inferior facet of the greater tubercle, and the inferior por- Gibbs 1999; Michilsens et al. 2009; Diogo et al. 2010, 2012,
tion inserted onto the antero-inferior surface of the greater 2013a,b, 2017). In one bonobo fetus, teres major was fused
tubercle (Ashaolu et al. 2015). with triceps brachii and subscapularis (Diogo et al. 2017).
Variations inserts via a tendon onto the lesser tubercle of the humerus
Description (Standring 2016).
Teres major may be absent (Macalister 1867a, 1875;
Bergman et al. 1988; Lamb 2016). Teres major can fuse Innervation
with latissimus dorsi or its tendon, send a slip to the long Subscapularis is innervated by the upper and lower sub-
head of the triceps or the brachial fascia, or fuse with rhom- scapular nerves (Standring 2016).
boid major (Macalister 1875; Le Double 1897; Aasar 1947;
Bergman et al. 1988; Iamsaard et al. 2012; Standring 2016;
Lamb 2016). Slips connecting teres major to biceps brachii Comparative Anatomy
or coracobrachialis have also been observed (Macalister Subscapularis has a similar typical presentation in the apes,
1875; Aggarwal et al. 2009; Lamb 2016). It may connect extending from the subscapular fossa to the lesser tuber-
with the shoulder joint capsule (Macalister 1875). cle of the humerus (Gibbs 1999; Diogo et al. 2010, 2012,
2013a,b, 2017). The insertion can sometimes extend to the
Prevalence shaft of the humerus as a variant in all apes except gibbons
In a sample of 52 arms, Mori (1964) found that the tendon (Beddard 1893; Sonntag 1923; Gibbs 1999; Diogo et al.
of teres major was fused with the tendon of latissimus dorsi 2010, 2012, 2013a,b, 2017). It may insert onto the shoulder
in 28 arms (53.8%), the insertions of teres major and latis- joint capsule in bonobos (Miller 1952; Diogo et al. 2017).
simus dorsi were parallel on the humerus in 13 arms (25%),
the insertions of teres major and latissimus dorsi made a v Variations
shape on the humerus in 10 arms (19.2%), and a slip passed Description
from the infraglenoid tubercle of the scapula to the tendon
Standring (2016) reports that variation in subscapularis is
of teres major in one arm (1.9%).
not common, while Lamb (2016) states that this muscle has
the most reported variations of any of the rotator cuff mus-
Anomalies cles. In addition to the lesser tubercle, its tendon can insert
Description into the bicipital groove or greater tubercle (MacDonald
On the left side of a fetus with craniorachischisis, a muscu- et al. 2007; Cash et al. 2009; Lamb 2016). Subscapularis
lar slip connected teres major to triceps brachii (Alghamdi can divide into two discrete parts or numerous slips
et al. 2017). In a fetus with trisomy 18 and cyclopia, teres (Macalister 1867a, 1875; Lamb 2016). Subscapularis
major fused with latissimus dorsi near their humeral inser- may connect with biceps brachii, pectoralis major, or tri-
tion at the intertubercular sulcus (Smith et al. 2015). In a ceps brachii (Macalister 1875; Le Double 1897; Bergman
male fetus with triploidy, teres major was doubled on the et al. 1988; Lamb 2016). Mori (1964) found an abnormal
right side (Moen et al. 1984). slip that extended from levator scapulae to subscapularis.
Coracobrachialis profundus may insert into subscapularis
or its tendon (Macalister 1875; Knott 1883a). Le Double
Prevalence (1897) describes a slip that originated from the subscapular
N/A tendon and attached to the axillary fascia or skin. An acces-
sory slip may extend from the shoulder joint capsule and
lesser tubercle and insert onto latissimus dorsi (Namking
Iamsaard et al. (2012) suggest that variations in teres major et al. 2013).
insertions (such as the insertion these authors observed Subscapularis is associated with several named varia-
onto the tendon of latissimus dorsi) may cause unusual tions. Glenobrachialis (Gruber) refers to a slip that origi-
movement of the arm, and knowledge of such variations nates with the long head of biceps brachii and attaches onto
is important when planning tendon transfer to treat rotator the surgical neck of the humerus (Macalister 1875; Knott
cuff tears. 1883a; Bergman et al. 1988). Subscapulo-capsularis may
insert into the bicipital groove, shoulder joint capsule, or
subscapularis (figure 3.8) neck of the humerus (Macalister 1867a, 1875; Lamb 2016).
Subscapularis-teres-latissimus is an accessory muscle that
See also: Subscapularis minor has three different presentations and fuses with the sub-
scapularis insertion (Kameda 1976; Lamb 2016). Depressor
Synonyms
tendinis subscapularis majoris (also referred to as deltoidius
N/A profundus, tensor capsulae humeralis, or retinaculum mus-
culare tendinis subscapularis majoris (Gruber) originates
from the subscapularis tendon and inserts into the surgi-
Description cal neck of the humerus (Macalister 1875; Knott 1883a).
Subscapularis occupies the subscapular fossa. It primar- Subscapularis minor may also be present as a variation (see
ily originates from the costal surface of the scapula and the entry for this muscle).
Prevalence or just below the lesser tubercle (Knott 1883a; Aasar 1947;
Mori (1964) found a slip from the medial margin of levator Bergman et al. 1988; Standring 2016; Pires et al. 2017).
scapulae to the ventral surface of subscapularis in 22 out of Although Staniek and Brenner (2012) suggested that the
100 sides (22%). Kameda (1976) observed the presence of infraglenoid muscle described by them (and by Lee et al.
subscapularis-teres-latissimus in 3.8% of cases. In an MRI 2019a) should be distinguished from subscapularis minor,
study of 50 individuals, Cash et al. (2009) found that sub- this muscle is similar in morphology, origin, and insertion.
scapularis primarily inserted onto the lesser tubercle in ten
cases (20%), the bicipital groove in 33 cases (66%), and the Innervation
greater tubercle in seven cases (14%).
The infraglenoid muscles described by Staniek and Brenner
(2012) were innervated by the axillary nerve.
Anomalies
An anomalous presentation of subscapularis includes the Gruber (1859) noted the presence of subscapularis minor in
insertion splitting the plane of muscle (Smith et al. 2015). ten out of 200 limbs (5%). Knott (1883a) found this muscle in
four out of 39 subjects (10.3%). Le Double (1897) reports that
Prevalence Krause found this muscle in 3 out of 35 specimens (8.6%) and
In the literature review conducted by Smith et al. (2015), Testut found this muscle in 3 out of 18 specimens (16.7%).
they found that subscapularis with insertion splitting the
plane of muscle was present in 1 out of 26 individuals with Anomalies
trisomy 18 (3.8%). N/A
N/A
Breisch (1986) states that an accessory subscapularis muscle
can create a myotendinous tunnel through which the axil-
subscapularis MinOr (figure 3.8) lary and subscapular nerves pass, which may lead to nerve
entrapment and its associated neurological symptoms. Pires
See also: Subscapularis
et al. (2017) similarly state that an accessory subscapularis
Synonyms passing over the axillary nerve can contribute to quadran-
gular space compression syndrome.
This muscle is also referred to as scapularis secundus or
accessory subscapularis (Gruber 1859; Le Double 1897;
Bergman et al. 1988), infraspinatus secundus, subscapulo- latissiMus DOrsi (figures 3.4, 3.9, anD 3.11)
humeralis, subglenoidalis (Knott 1883a), and potentially
See also: Pectorodorsalis, Dorsoepitrochlearis, Levator ten-
the infraglenoid muscle (Lee et al. 2019a).
dinis musculi latissimus dorsi
Synonyms
Latissimus dorsi is also referred to as the grand dorsal
Comparative Anatomy (Bakkum and Miller 2016).
In a common chimpanzee dissected by Ziegler (1964), an
accessory head arose from the superior-most part of the axil- Typical Presentation
lary border of subscapularis. A few fbers of this head inserted Description
onto the capsular ligament while the rest attached to the Latissimus dorsi originates from the spines of the lower six
humerus distal to the lesser tubercle. In an orangutan dissected thoracic vertebrae, the thoracolumbar fascia and therefore the
by Sullivan and Osgood (1927), there was an upper bundle of spines of the lumbar and sacral vertebrae, the posterior portion
subscapularis that was distinct from the rest of the muscle. of the iliac crest, and the lower three or four ribs (Standring
2016). Its fbers converge into a tendon that inserts onto the
Variations intertubercular sulcus of the humerus (Standring 2016).
Description
This muscle is a variation of subscapularis. When present, Innervation
subscapularis minor originates from the upper axillary bor- Latissimus dorsi is innervated by the thoracodorsal nerve
der of the scapula (Knott 1883a; Bergman et al. 1988) or the (Standring 2016).
anterior and lateral surface of subscapularis (Breisch 1986;
Pires et al. 2017). Its insertion is variable, with possible Comparative Anatomy
attachments to the shoulder joint capsule, the crest of the Latissimus dorsi has a similar typical presentation in
lesser tubercle of the humerus, the intertubercular sulcus, the apes, extending from the vertebrae, ribs, pelvis, and
FIGURE 3.9 Coracobrachialis and associated variations in anterior view.
thoracolumbar fascia to the intertubercular sulcus of the It can vary in the number of costal attachments (Wood
humerus (Gibbs 1999; Michilsens et al. 2009; Diogo et al. 1868; Macalister 1875; Mori 1964; Bakkum and Miller
2010, 2012, 2013a,b, 2017). Fusion of the latissimus dorsi 2016; Standring 2016). It may originate only from the
tendon with teres major is observed as a common variant lumbar vertebrae (Rickenbacher et al. 1985). It can also
in all apes (see the entry for teres major). In orangutans, originate only from the ribs, instead of a more extensive
latissimus dorsi is often divided into two or more bundles attachment, or it can be divided into multiple fascicles
or sends accessory slips to other structures (Beddard 1893; (Macalister 1875; Bergman et al. 1988; Bakkum and Miller
Primrose 1899, 1900; Schück 1913; Stewart 1936; Diogo 2016). Common variations include additional attachments
et al. 2013b). In gorillas, latissimus dorsi often blends with to the inferior angle of the scapula; the presence of pectoro-
trapezius (Raven 1950; Preuschoft 1965; Gibbs 1999; Diogo dorsalis (axillary arch), which connects latissimus dorsi to
et al. 2010). Latissimus dorsi was also partially blended pectoralis major or other nearby muscles; and the presence
with trapezius in a fetal bonobo (Diogo et al. 2017). An of dorsoepitrochlearis, which connects latissimus dorsi the
additional origin from the inferior angle of the scapula has upper arm (see the entries for these muscles) (Macalister
been observed in an adult bonobo (Miller 1952). 1875; Mori 1964; Bergman et al. 1988; Standring 2016;
Bakkum and Miller 2016). Latissimus dorsi or its tendon
Variations may be continuous with, or send a slip to, teres major
(Macalister 1875; Mori 1964; Aasar 1947; Bergman et al.
Description 1988; Iamsaard et al. 2012; Bhatt et al. 2013; Bakkum
Latissimus dorsi may be absent on one or both sides of and Miller 2016; Lamb 2016; Standring 2016). A slip can
the body (Bergman et al. 1988; Bakkum and Miller 2016). also pass to the long head of the triceps (Macalister 1875;
Standring 2016; Bakkum and Miller 2016) or to the frst rib Variations
(Miyauchi 1982a). It may overlap with the external oblique Description
(Macalister 1875).
Levator tendinis musculi latissimus dorsi (Gruber) origi-
Prevalence nates from the coracoid process and shoulder joint capsule
and has a tendinous insertion onto the tendon of latissimus
Mori (1964) studied the attachments of latissimus dorsi
dorsi (Gruber 1844; Wood 1868; Macalister 1875; Knott
on 100 sides from 50 cadavers. This author found that
1883a; Le Double 1897; Bergman et al. 1988; Del Sol and
the upper limit of the aponeurosis from the vertebral col-
Olave 2005; Moore and Rice 2018). Moore and Rice (2018)
umn was the spinous process of the ffth thoracic verte-
suggest that the presence of this muscle may represent a
bra on two sides (2%), sixth thoracic vertebra on 16 sides
failed migration of a coracobrachialis longus variant.
(16%), seventh thoracic vertebra on 44 sides (44%), eighth
thoracic vertebra on 30 sides (30%), and ninth thoracic Innervation
vertebra on eight sides (8%). Mori (1964) studied the
Innervation of levator tendinis musculi latissimus dorsi
costal origins of this muscle on 60 sides and found that
has not been described, but it is found in close proxim-
latissimus dorsi had origins from ribs 9–12 on 25 sides
ity to the nerves passing through the axilla (Del Sol and
(41.7%) and ribs 10–12 on 35 sides (58.3%). This author
Olave 2005).
also observed an attachment to the inferior angle of the
scapula in 53.3% of cases. Prevalence
Anomalies Del Sol and Olave (2005) observed the presence of this
muscle in 1 out of 108 adult male cadavers (0.9%).
Description
In a cadaver with a severe case of congenital scoliosis, Anomalies
Stevenson et al. (2014) found signifcant asymmetry in the
Description
latissimus dorsi muscles. In a fetus with craniorachischisis,
the right latissimus dorsi was reduced compared to the left On the left side of a neonate with trisomy 13, levator ten-
side (Alghamdi et al. 2017). Smith et al. (2015) observed dinis musculi latissimus dorsi originated just beneath the
the fusion of teres major with latissimus dorsi near their coracoid from the fascia covering the shoulder joint cap-
humeral insertion at the intertubercular sulcus in one fetus sule and inserted into the latissimus dorsi tendon (Pettersen
with trisomy 18 and cyclopia. On both sides of a male infant et al. 1979).
with triploidy, latissimus dorsi was divided into scapular
Prevalence
and humeral heads (Moen et al. 1984).
In their literature review, Smith et al. (2015) found that this
Prevalence muscle was present in 1 out of 24 individuals with trisomy
N/A 13 (4.2%).
Clinical Implications Clinical Implications

Understanding variations in latissimus dorsi is important Del Sol and Olave (2005) suggest that the presence of this
for the planning and execution of the numerous reconstruc- muscle may lead to compression of branches of the brachial
tive and cosmetic surgeries that utilize parts of this muscle plexus or axillary vessels.
(Bhatt et al. 2013).
cOracObrachialis (figure 3.9)
levatOr tenDinis Musculi latissiMus DOrsi (figure 3.9) See also: Coracobrachialis profundus, Coracobrachialis
See also: Coracobrachialis longus, Latissimus dorsi longus
Synonyms Synonyms
This muscle is also referred to as coracobrachialis acces- N/A
sorius, coracobrachialis superior, coracobrachialis minor or
secundus, le court coracobrachialis (Bergman et al. 1988;
Del Sol and Olave 2005; Akita and Nimura 2016a), or cora- Description
cobrachialis brevis s. rotator humeri (Wood 1864). Coracobrachialis originates from the coracoid process
along with the tendon of the short head of the biceps and
Typical Presentation inserts onto the shaft of the humerus (Standring 2016).
Innervation
Comparative Anatomy Coracobrachialis is innervated by the musculocutaneous
N/A nerve (Standring 2016).
Comparative Anatomy along the medial border of the long head of the biceps (Catli
Coracobrachialis has a similar typical presentation in the et al. 2012). Piagkou et al. (2019) report a coracobrachialis
apes, extending from the coracoid process to the proximal with a superficial and deep head that was connected with an
humerus (Gibbs 1999; Diogo et al. 2010, 2012, 2013a,b, 2017). accessory muscle bundle. This muscle was associated with a
“three-headed” biceps, with the third head originating from
the tendon of the short head and the coracobrachialis to
Variations
insert onto the radial tuberosity (Piagkou et al. 2019).
Description
Coracobrachialis is comprised of three parts: the proximal
Prevalence
(deep) portion, the middle portion (corresponding to cora-
cobrachialis proprius/medius), and the distal (superficial) In a study of 50 arms, Mori (1964) found that coracobra-
portion, and only in some cases do the parts appear distinct chialis was completely separated into superficial and deep
from one another (Mori 1964; Bergman et al. 1988; Akita layers in eight arms (16%), partially separated into super-
and Nimura 2016a). The proximal portion occasionally pres- ficial and deep layers in four arms (8%), and not separated
ents as a muscle referred to as coracobrachialis profundus, into layers in 38 arms (76%).
and the distal portion occasionally presents in the form of
coracobrachialis longus (see the entries for these muscles) Anomalies
(Bergman et al. 1988; Akita and Nimura 2016a). Georgiev Description
et al. (2017a) report a coracobrachialis muscle in a male Macalister (1875) notes that Barkow observed the absence of
cadaver in which coracobrachialis brevis, coracobrachialis, coracobrachialis in an anomalous individual. In a fetus with
and a variant of coracobrachialis longus (their “coracoepi- craniorachischisis dissected by Alghamdi et al. (2017), they
trochlearis”) are distinct from each other and well-developed. found that coracobrachialis on the right side of the body was
Coracobrachialis can send additional slips to the lesser fused with the short head of biceps brachii at its proximal
tubercle, medial epicondyle, supracondylar process of the end and fused with dorsoepitrochlearis at its distal end. The
humerus, brachialis, triceps brachii, or the medial inter- fusion of the proximal end of this muscle with the short head
muscular septum (Macalister 1875; Bergman et al. 1988; of biceps brachii was also observed on the left side of the
Standring 2016; Akita and Nimura 2016a). Its origin may body. Alghamdi et al. (2017) note that coracobrachialis was
connect with the insertion of pectoralis minor (Macalister associated with the median nerve in this specimen.
1875). Its tendon may receive slips from the tendons of In the female fetus with trisomy 18 dissected by
latissimus dorsi or teres major (Macalister 1875). Garbelotti Alghamdi et al. (2018), the insertion of pectoralis minor
et al. (2017) report a slip of coracobrachialis that inserted on the right side extended inferiorly to the fascia of cora-
into the intermuscular septum in the proximal humerus, cobrachialis. The proximal end of coracobrachialis was
trapping the lateral cord of the brachial plexus against it and completely fused with the proximal end of the short head
coracobrachialis. Coracobrachialis can also be absent as a of biceps brachii (side not indicated). In a dissection of a
variation (Bergman et al. 1988; Akita and Nimura 2016a). trisomy 18 and cyclopic fetus completed by Smith et al.
Several authors report variations in coracobrachialis (2015), they observed that coracobrachialis on the right side
associated with variations in biceps brachii. El-Naggar and of the body was fused with the short head of the biceps and
Zahir (2001) reported a coracobrachialis with two bellies inserted through its entire length along the anteromedial
that formed inferior to its origin on the coracoid, the first proximal humerus.
inserting onto the middle of the humerus and the second onto Bersu et al. (1976) describe a male infant with Hanhart
the medial head of the triceps, with the musculocutaneous syndrome. On the left side of this specimen, the upper limb
nerve passing in between them. This muscle was associated was absent below the elbow, and rudiments of the radius and
with a “three-headed” biceps, with the third head originat- ulna were present. Coracobrachialis bifurcated and had a
ing mostly from the humerus and partly from the tendinous deep medial portion that represented the usual muscle and a
origin of the medial head of the triceps to insert onto the superficial portion that attached to the capsule of the elbow
common tendon for the biceps (El-Naggar and Zahir 2001). joint, the latter of which sent a slip to the biceps tendon. On
Catli et al. (2012) observed the presence of a coracobrachi- the left arm of a male neonate with Meckel syndrome, cor-
alis with a superficial and deep head that both originated acobrachialis had two heads and the superficial head was
from the coracoid process and inserted onto the middle third encircled by the two roots of the median nerve (Pettersen
of the humerus, and a “capsular” head that originated from 1984). A partial fusion of coracobrachialis with the short
the articular capsule of the glenohumeral joint. This muscle head of biceps brachii has been observed in infants with
was associated with a “four-headed” biceps brachii that had Neu-Laxova syndrome (Shved et al. 1985).
two supernumerary heads (Catli et al. 2012). The first super-
numerary head arose from the insertion of coracobrachialis Prevalence
as a united tendon and inserted onto the medial aspect of the In their literature review, Smith et al. (2015) found that
short head of biceps brachii (Catli et al. 2012). The second coracobrachialis was split into proximal and middle por-
arose from the upper third of the humerus and was fused tions in 1 out of 24 individuals with trisomy 13 (4.2%).
Coracobrachialis had otherwise anomalous presentations Prevalence

in 3 out of 26 individuals with trisomy 18 (11.5%); in two Howell and Straus (1932) observed the presence of coraco-
female neonates, coracobrachialis was represented by small brachialis profundus in 3 out of 39 arms that they dissected
bellies in addition to the short head of the biceps, and in (7.7%).
another female neonate, coracobrachialis was fused with
the short head of the biceps (Smith et al. 2015). In a study
of six upper limbs from three infants with Neu-Laxova syn- Anomalies
drome, Shved et al. (1985) found that coracobrachialis was Description
partially fused with the short head of biceps brachii in all In a fetus with trisomy 18 and cyclopia, Smith et al. (2015)
six upper limbs (100%). found coracobrachialis profundus on the left side of the
body. This muscle was also present on the left side of a neo-
Clinical Implications nate with trisomy 13 (Aziz 1980).
Division of coracobrachialis into distinct parts can lead
Prevalence
to the compression of the musculocutaneous nerve as the
nerve passes between the parts (El-Naggar and Zahir 2001). In their literature review, Smith et al. (2015) found that this
muscle was present in 2 out of 24 individuals with trisomy
13 (8.3%).
cOracObrachialis prOfunDus (figure 3.9)
See also: Coracobrachialis
N/A
Synonyms
This muscle is also referred to as coracocapsularis, coraco- cOracObrachialis lOngus (figure 3.9)
brachialis superior, or coracobrachialis brevis (Macalister
1875; Bergman et al. 1988; Smith et al. 2015; Akita and See also: Coracobrachialis
Nimura 2016a).
Synonyms
Typical Presentation This muscle is also referred to as coracobrachialis inferior
This muscle is only present as a variation or anomaly. or coracobrachialis superfcialis (Wood 1864; Bergman
et al. 1988; Smith et al. 2015).
Comparative Anatomy
Coracobrachialis profundus is typically not present as a dis-
tinct structure in the apes (Diogo et al. 2010, 2012, 2013a,b, This muscle is only present as a variation or anomaly.
2017). Kallner (1956) reports the presence of this muscle
Comparative Anatomy
in one orangutan, and Hepburn (1892) reports the presence
of some fbers corresponding to this muscle in one gorilla. Coracobrachialis longus is typically not present as a dis-
Coracobrachialis profundus may be present as a variation tinct structure in the apes (Diogo et al. 2010, 2012, 2013a,b,
in common chimpanzees (Parsons 1898a,b; Howell and 2017). Kallner (1956) describes a coracobrachialis longus
Straus 1932). in one orangutan, but the presence of this muscle is not cor-
roborated by the illustrations provided by the author (Diogo
Variations et al. 2013b). In one gorilla, Hepburn (1892) reports the pres-
Description ence of some fbers potentially corresponding to this mus-
cle inserting onto the intermuscular septum of the arm, but
Coracobrachialis profundus refers to a deep accessory
Howell and Straus (1932) suggest that this is just an exten-
bundle near the proximal part of coracobrachialis that
sion of the coracobrachialis proprius. Coracobrachialis
originates from the coracoid process and inserts into the
longus may be present as a rare variation in common chim-
shoulder joint capsule or into the surgical neck of the
panzees (Howell and Straus 1932; Oishi et al. 2009).
humerus below the lesser tubercle (Macalister 1875; Knott
1883a; Bergman et al. 1988; Akita and Nimura 2016a). It
Variations
may also insert into the tendon of subscapularis, fbers of
subscapularis, or the capsular ligament of the shoulder Description
(Macalister 1875; Knott 1883a). Macalister (1875) notes Coracobrachialis longus is a superfcial and distal bundle of
that coracobrachialis profundus may be a variation of the coracobrachialis and extends from coracoid process to the
deltoid or subscapularis. distal humerus, potentially inserting onto the medial epi-
condyle or medial intermuscular septum of the arm (Wood
Innervation 1864; Macalister 1875; Knott 1883a; Akita and Nimura
This muscle is likely innervated by the musculocutaneous 2016a; Paraskevas et al. 2016). It may be continuous with
nerve. brachialis (Macalister 1875). Georgiev et al. (2017a) use
coracoepitrochlearis to refer to a variant of coracobrachialis

longus that originates from the coracoid process to insert onto
the medial epicondyle. Georgiev et al. (2018a) use humero-
epitrochlearis to refer to a variant of coracobrachialis longus
that originates from the medial surface of the middle part of
the humerus to insert onto the medial epicondyle.
Innervation
Coracobrachialis longus is innervated by the musculocuta-
neous nerve (Georgiev et al. 2017a, 2018a).
Prevalence
N/A
Anomalies
Description
Bersu et al. (1976) describe a male infant with Hanhart syn-
drome. On the left side of this specimen, the upper limb was
absent below the elbow, and rudiments of the radius and
ulna were present. Coracobrachialis bifurcated and had a
deep medial portion that represented the usual muscle and a
superfcial portion that attached to the capsule of the elbow
joint, the latter of which sent a slip to the biceps tendon. In a
neonate with trisomy 18, coracobrachialis longus extended
from the coracoid process to the medial epicondyle of the
humerus (Aziz 1979).
Prevalence
they found that this muscle was present in 2 out of 26 indi-
viduals with trisomy 18 (7.7%). FIGURE 3.10 Brachialis and biceps brachii in anterior view. V1
indicates an additional bundle of biceps brachii that originates
Clinical Implications from brachialis and joins the biceps insertion tendon.
El-Naggar and Al-Saggaf (2004) noted that the presence Innervation
of coracobrachialis longus can contribute to compression
Brachialis is typically innervated by the musculocutaneous
of the median nerve and brachial artery, as well as palsy
nerve and radial nerve (Standring 2016). It may sometimes
hypoxia of the forearm and hand. Paraskevas et al. (2016)
be supplied by the median nerve (Machida 1961; Akita and
suggest that the presence of this muscle could induce symp-
Nimura 2016a).
toms of median, ulnar, or medial antebrachial cutaneous
neuropathy. Georgiev et al. (2017a) note that an accessory Comparative Anatomy
belly of coracobrachialis inserting onto the medial epicon-
Brachialis has a similar typical presentation in the apes,
dyle of the humerus may be a good candidate to be used as
extending from the humeral shaft to the ulnar tuberos-
a graft for surgery, but its presence may lead to confusion
ity, and sometimes the coronoid process (Raven 1950;
during medical imaging of the arm.
Miller 1952; Gibbs 1999; Diogo et al. 2010, 2012, 2013a,b,
2017). Many of the same variations are also present in the
brachialis (figure 3.10) apes, including division into multiple parts, the presence
of accessory slips, or fusion with other muscles (Sonntag
Synonyms 1924; Sullivan and Osgood 1927; Howell and Straus
Other names for brachialis include brachialis anticus and 1932; Ziegler 1964; Gibbs 1999; Diogo et al. 2010, 2012,
fexor brachii brevis (Hepburn 1892; Parsons 1898b). 2013a,b, 2017).
Typical Presentation Variations

Brachialis originates from the lower half of the humeral Brachialis may send slips or otherwise have connections to
shaft and the associated intermuscular septa and inserts via the radius near the tuberosity, both the radius and ulna, the
a tendon onto the coronoid process and the tuberosity of the bicipital aponeurosis, the forearm fascia, or the semilunar
ulna (Standring 2016). fascia (Macalister 1875; Mori 1964; Bergman et al. 1988;
Standring 2016; Akita and Nimura 2016a). Other common alobar holoprosencephaly. On the right side, brachialis was
variations include division into multiple parts, complete divided into a superfcial and deep head. A low origin of
absence, and fusion with biceps brachii, brachioradialis, brachialis was observed in infants with Neu-Laxova syn-
coracobrachialis, extensor carpi radialis, or pronator teres drome (Shved et al. 1985).
(Wood 1864, 1866, 1867a, 1868; Macalister 1875; Machida
1961; Mori 1964; Bergman et al. 1988; Standring 2016; Prevalence
Akita and Nimura 2016a). Brachialis may have three or In their literature review, Smith et al. (2015) found that
four heads (Wood 1864; Macalister 1875; Akita and Nimura brachialis had two accessory slips joining biceps in 2 out
2016a). Macalister (1875) notes that in rare cases, brachialis of 24 individuals with trisomy 13 (8.3%). Brachialis was
may send a slip to fexor digitorum superfcialis. otherwise anomalous (high origin, divided into two por-
Brachialis lateralis minor is a supernumerary muscle tions) in 3 out of 26 individuals with trisomy 18 (11.5%). An
associated with brachialis that inserts into the ulnar tuberos- accessory brachialis was present in 1 out of 26 individu-
ity, while brachialis internus minor is medially situated and als with trisomy 18 (3.8%). In a study of six upper limbs
also inserts into the ulna (Macalister 1875; Knott 1883a). from three infants with Neu-Laxova syndrome, Shved et al.
Capsularis sub-brachialis (Portal) or artecularis cubiti (1985) found that a low origin of brachialis was present in
anterior refers to one or two deep bundles of brachialis that two upper limbs (33%).
attach to the capsule of the elbow joint (Macalister 1875;
Nishi 1966; Bergman et al. 1988; Akita and Nimura 2016a). Clinical Implications
Supinator radii brevis accessorius extends from brachialis Loukas et al. (2006c) suggest that an accessory brachialis
to the radial tuberosity (Knott 1883a). Pai et al. (2008a) can contribute to median nerve palsy. Pai et al. (2008a) state
report an accessory brachialis that originated from the lat- that an accessory brachialis may contribute to radial-tunnel
eral part of brachialis and the lateral intermuscular septum syndrome. Vadgaonkar et al. (2008) suggest that an acces-
and split into two slips, the medial slip inserting onto the sory brachialis can lead to various compression syndromes
deep fascia of pronator teres and the lateral slip inserting involving the median nerve or the brachial artery.
onto the fascia of supinator. Vadgaonkar et al. (2008) report
an accessory brachialis that inserted on the shaft of the
biceps brachii (figure 3.10)
radius and formed a fbro-muscular tunnel above the cubi-
tal fossa containing the median nerve, brachial artery, and Synonyms
other structures. This muscle is also referred to as biceps fexor cubiti (Owen
1868; Hepburn 1892).
Prevalence
Knott (1883a) found an accessory slip between brachialis Typical Presentation
and biceps brachii in 3 out of 49 subjects (6.1%). Knott Description
(1883a) also found brachialis internus minor in 3 out of 49 The short head of biceps brachii originates via a tendon
subjects (6.1%). Mori (1964) found that in 12 out of 50 arms from the coracoid process (Standring 2016). The long head
(24%), brachialis was divided into two distinct heads. In of biceps brachii originates within the shoulder joint cap-
one arm (2%), an accessory slip originated from brachialis sule via a tendon from the supraglenoid tubercle of the scap-
to insert onto the radial tuberosity (Mori 1964). ula (Standring 2016). Both bellies converge into a tendon
to insert onto the radial tuberosity (Standring 2016). The
Anomalies
tendon has a medial expansion, termed the bicipital apo-
Description neurosis (Standring 2016).
In a neonate with trisomy 13, brachialis sent two accessory
slips to biceps brachii on the right side (Aziz 1980). In a Innervation
neonate with trisomy 18, the right brachialis was divided Biceps brachii is innervated by the musculocutaneous nerve
into a thick part that originated below the deltoid tuberos- (Standring 2016).
ity and a deep, thin sheet that originated from the distal
humerus (Aziz 1979). In a fetus with craniorachischisis, Comparative Anatomy
Alghamdi et al. (2017) observed that brachialis on the right Biceps brachii has a similar typical presentation in the apes,
upper limb was absent, while the left brachialis was con- arising from the coracoid process or proximal humerus
nected to an extra head of triceps brachii. In this specimen, and the supraglenoid tubercle and inserting into the radial
Alghamdi et al. (2017) note that brachialis was associated tuberosity (Gibbs 1999; Diogo et al. 2010, 2012, 2013a,b,
with the median nerve. 2017). Accessory heads are occasionally present in com-
Bersu et al. (1976) describe a male infant with Hanhart mon chimpanzees (Howell and Straus 1932; Diogo et al.
syndrome. On the left side of this specimen, the upper limb 2013a). The bicipital aponeurosis is usually absent in bono-
was absent below the elbow, and rudiments of the radius and bos and orangutans (Diogo et al. 2013b, 2017). In gibbons,
ulna were present. Brachialis was absent. Mieden (1982) the distal portion of biceps brachii is blended with fexor
describes an infant with median cleft lip, hypotelorism, and digitorum superfcialis (Kohlbrügge 1890–1892; Howell
and Straus 1932; Gibbs 1999; Michilsens et al. 2009; Diogo by Theile in one in nine subjects (11.1%), by Hallett in 1 out
et al. 2012). of 15 subjects (6.7%), and by Wood in 18 out of 175 subjects
(10.3%). In a sample of 49 subjects, Knott (1883a) found
Variations an accessory slip between brachialis and biceps brachii in
Description three subjects (6.1%) and an additional head between biceps
The short head and the long head can sometimes be fused or and coracobrachialis in fve subjects (10.2%). Sato (1969)
one of them could be absent (Macalister 1875; Knott 1883a; found a third head in 35 out of 286 limbs from Kyushu-
Bergman et al. 1988; Akita and Nimura 2016a). When the Japanese males (9.07%) and in 17 out of 250 limbs (6.8%)
long head is defcient or absent, it may be replaced by a from females.
singular or doubled tendon that originates from the proxi- Mori (1964) found the third head of the biceps in 10 out
mal humerus, shoulder joint capsule, or the pectoralis major of 50 arms (20%). The third head originated from the dis-
tendon (Bergman et al. 1988; Akita and Nimura 2016a; tal aspect of the deltoid tuberosity in four arms (8%), the
Katsuki et al. 2018). Cutler et al. (2018) report a long head distal aspect of the insert site of coracobrachialis in three
of the biceps tendon with a trifurcate origin. Collett et al. arms (6%), the tendon of pectoralis major in two arms (4%),
(2018) report a short head of biceps brachii with an origin and the lesser tubercle in one arm (2%). Ravi et al. (2020)
from an aponeurotic sheet that extended over the coracoid reported supernumerary heads of the biceps in 5 out of 50
process, coracoacromial ligament, the inferior surface arms (10%). The biceps in one of the specimens had four
of the acromion, and the greater tubercle of the humerus. heads, and the muscle in the other specimens had three
Biceps brachii may be entirely absent (Macalister 1875; heads (Ravi et al. 2020).
Bergman et al. 1988). Several authors report variations
in biceps brachii associated with variations in coracobra- Anomalies
chialis (see coracobrachialis entry for more details). Other Description
variations of biceps brachii recorded by Macalister (1875) In a fetus with craniorachischisis, Alghamdi et al. (2017)
include information on absence, variable origins and inser- observed that biceps brachii on the left upper limb demon-
tions, supernumerary heads, and accessory slips. strated a fusion between the short head and coracobrachia-
A third or humeral head may be present, which typi- lis. There was also an abnormal slip connecting teres major
cally arises from brachialis (Macalister 1875; Mori 1964; to the long head. A fusion of the short head and coracobra-
Sato 1969; Bergman et al. 1988; Akita and Nimura 2016a; chialis was also observed on the right side. The short head
Standring 2016; Ravi et al. 2020). Moore and Rice (2018) of the right biceps brachii also had an abnormal origin at the
report a “four-headed” biceps with both an inferomedial lesser tubercle of the humerus, and an abnormal insertion
and inferolateral humeral head. Accessory slips may also be onto the lateral condyle and medial supracondylar ridge of
present and give biceps brachii the appearance of having up the humerus. The long head of the right biceps brachii was
to fve heads (Macalister 1875; Bergman et al. 1988; Akita missing. In this specimen, Alghamdi et al. (2017) note that
and Nimura 2016a). An accessory slip may originate from biceps brachii was associated with the median nerve.
the humerus at the insertion of coracobrachialis, extend to In a neonate with trisomy 13, the right biceps brachii
brachialis, and connect with the short head of biceps bra- had two accessory heads (Aziz 1980). One arose from the
chii and the semilunar fascia (Bergman et al. 1988). Other medial crest of the bicipital groove, and the other arose
accessory slips can arise from the neck of the humerus and from the distal half of the humerus. Biceps brachii on this
connect to the short and long head, or from the intertuber- side also received two slips from brachialis. On the left side,
cular sulcus, lesser tubercle, deltoid tuberosity, lower end the short head of the biceps was absent, but there were also
of the humerus, shoulder joint capsule, tendon of pectoralis two accessory heads. One arose from the lesser tubercle,
major, ulna, radius, or antebrachial fascia (Macalister 1875; and the other arose from the proximal humerus.
Mori 1964; Bergman et al. 1988; Akita and Nimura 2016a; In a neonate with trisomy 18, the superior portion of
Standring 2016). Muscular slips can connect biceps brachii biceps brachii was divided into a long head, short head,
to other muscles including the deltoid, brachialis, brachio- and a fascicle that was continuous with the deltopectoral
radialis, pronator teres, fexor carpi radialis, or fexor digi- complex (Aziz 1979). In a female fetus with trisomy 18,
torum profundus (Macalister 1875; Knott 1883a; Bergman the proximal end of coracobrachialis was completely fused
et al. 1988; Akita and Nimura 2016a). Brachiofascialis with the proximal end of the short head of biceps brachii
(Struthers), also referred to as brachialis accessorius or (side not indicated) (Alghamdi et al. 2018). In a fetus with
supinator brevis accessorius, is a slip extending between trisomy 18 and cyclopia, Smith et al. (2015) observed the
biceps brachii and fascia over pronator teres (Wood 1864; presence of a tendon connecting biceps brachii to pectoralis
Macalister 1875; Knott 1883a; Akita and Nimura 2016a). major, as well as the bilateral absence of the bicipital apo-
neurosis. The short head of the left biceps brachii was miss-
Prevalence ing, and the short head of the right biceps brachii was fused
Macalister (1875) found a third head present in about one in with coracobrachialis. In an adult with trisomy 21, the long
ten subjects (10%) and reports that a third head was found head of biceps brachii was absent bilaterally (Bersu 1980).
Bersu et al. (1976) describe a male infant with Hanhart triceps brachii (figure 3.8)
syndrome. On the left side of this specimen, the upper limb
was absent below the elbow, and rudiments of the radius Synonyms
and ulna were present. The insertion tendon of biceps bra- Triceps brachii is also referred to as multiceps extensor
chii inserted onto the radial rudiment and the long head was cubiti (Barnard 1875) or triceps extensor cubiti (Hepburn
defcient in fbers. In a male fetus with triploidy, Moen et al. 1892).
(1984) found an extra muscle belly between the short head
of biceps and the supraglenoid tubercle on the right side. In Typical Presentation
a female fetus with triploidy, the long head of biceps brachii Description
was absent bilaterally (Moen et al. 1984). Doubling of the The long head of triceps brachii originates via a tendon
short head of biceps brachii has been observed in infants from the infraglenoid tubercle of the scapula, the lateral
with Neu-Laxova syndrome (Shved et al. 1985). head originates via a tendon the posterior humeral shaft and
Mieden (1982) describes an infant with median cleft lip, the lateral intermuscular septum, and the medial head orig-
hypotelorism, and alobar holoprosencephaly (case I) and inates from the entire posterior surface of the humeral shaft
two male fetuses with cyclopia and alobar holoprosenceph- and the medial and lateral intermuscular septa (Standring
aly (cases II and III). On the right side of case I, an accessory 2016). The heads converge into a tendon that inserts onto
muscle belly originated from the biceps tendon and inserted the olecranon process of the ulna (Standring 2016).
onto the radius proximal to the attachment of supinator.
In case III, an accessory muscle belly was associated with Innervation
biceps brachii on both sides of the body. On the right side, Triceps brachii is innervated by the radial nerve (Standring
it originated from the coracoid process and attached to the 2016).
humerus proximal to brachialis. On the left, it originated
from the humerus lateral to coracobrachialis and fused with Comparative Anatomy
biceps brachii at its distal insertion (Mieden 1982).
Triceps brachii has a similar typical presentation in the
Prevalence apes, but the long head has a more extensive origin along
the axillary border of the scapula in all apes (Diogo et al.
In a study of six upper limbs from three infants with Neu-
2010, 2012, 2013a,b, 2017). Articularis cubiti (subanconeus)
Laxova syndrome, Shved et al. (1985) found that the short
is present in some common chimpanzees (Champneys 1872;
head of biceps brachii was doubled in two upper limbs
Sonntag 1923; Diogo et al. 2013a).
(33%). In their literature review, Smith et al. (2015) found
that biceps brachii muscle was variable (with accessory
Variations
heads, abnormal connections to other muscles, extra ten-
dons, and variable origins and insertions) in 12 out of 24 Description
individuals with trisomy 13 (50%), 15 out of 26 individuals Fibers of the long head of triceps brachii may extend to
with trisomy 18 (57.7%), and 1 out of 7 individuals with the capsule of the shoulder joint or to the axillary bor-
trisomy 21 (14.3%). These authors also found that an extra der of the scapula, and the medial head may sometimes
biceps brachii tendon inserting into pectoralis major was extend to form an arch across the ulnar groove (Macalister
present in 9 out of 26 individuals with trisomy 18 (34.6%). 1875; Mori 1964; Bergman et al. 1988; Akita and Nimura
A muscle belly extending between the biceps and the pec- 2016a). Prabhu et al. (2012) report a muscular connection
toralis major tendon was present in 4 out of 26 individuals between the long head and the lateral head. There may
with trisomy 18 (15.4%). The short head of biceps bra- be a tendinous connection between the tendon of the long
chii was absent in 2 out of 24 specimens with trisomy 13 head of the triceps and the latissimus dorsi tendon (Knott
(8.3%). Fusion of the short head with coracobrachialis was 1883a; Koizumi 1934; Ishimi 1950; Akita and Nimura
observed in 3 out of 26 individuals with trisomy 18 (11.5%) 2016a). There may also be a feshy slip connecting triceps
and in one out of seven individuals with trisomy 21 (14.3%). brachii with latissimus dorsi or subscapularis (Macalister
The long head of biceps brachii was absent in 1 out of 26 1875; Knott 1883a). Triceps brachii may be fused with
individuals with trisomy 18 (3.8%) and in one 1 of 7 indi- anconeus (Macalister 1875; Knott 1883a; Bergman et al.
viduals with trisomy 21 (14.3%) (Smith et al. 2015). 1988; Akita and Nimura 2016a). It may also fuse with
extensor carpi ulnaris (Macalister 1875). Subanconeus or
Clinical Implications articularis cubiti refers to the deep layer/medial head of
Nakatani et al. (1998) suggest that a biceps brachii with triceps attaching to the joint capsule of the elbow (Tubbs
four heads can contribute to median nerve palsy or brachial et al. 2006a; Akita and Nimura 2016a; Standring 2016).
artery compression. El-Naggar and Zahir (2001) state that An accessory or fourth head of the triceps can originate
the presence of a third head could lead to musculocutane- from the medial surface of the humerus, the axillary bor-
ous nerve compression. Ravi et al. (2020) state that super- der of the scapula, the coracoid process, or the capsule of
numerary heads of the biceps may be useful for muscle the shoulder joint (Macalister 1875; Sato 1969; Bergman
transfer surgeries. et al. 1988).
Prevalence epitrOchleOancOneus (figure 3.11)

Mori (1964) reports that the long head originated from the Entry adapted by permission from Springer Nature
capsule of the shoulder joint and the axillary border of the Customer Service Centre GmbH: Springer Current
scapula in 49 out of 50 arms (98%), while in one arm (2%) Molecular Biology Reports, Muscles Lost in Our Adult
the long head originated only from the axillary border of Primate Ancestors Still Imprint in Us: on Muscle Evolution,
the scapula. The medial head was proximal to the lateral Development, Variations, and Pathologies. E. Boyle, V.
head in 36 arms (72%) while the two heads originated at Mahon, R. Diogo, 2020.
an equal level in 14 arms (28%). In one arm (2%), a slip
originated between the medial and lateral heads and trav- Synonyms
elled distally to fuse with the lateral head. In another arm Alternative names for this muscle include anconeus quin-
(2%), a slip originated from the capsule of the shoulder tus, anconeus minimus, anconeus epitrochlearis muscle
joint and traveled distally to fuse with the medial head of (Wood) (Knott 1883a); fexor antebrachii ulnaris (Kallner
the triceps. Sato (1969) found a fourth head of triceps bra- 1956), epitrochleo-olecranonis, accessory anconeus, ulna-
chii present in 9 out of 364 limbs from Kyushu-Japanese ris internis, cubital anterieur (Wilson et al. 2016), anconeus
males (2.47%) and present in 9 out of 438 limbs (2.05%) quartus, anconeus medialis, anconeus sextus, anconeus
from females. parvus, and tensor fasciae antebrachii (Jouffroy 1971), epi-
trochleocubital or artecularis cubiti posterior (Nishi 1966).
Anomalies
Description Typical Presentation
In a fetus with craniorachischisis, Alghamdi et al. (2017) This muscle is only present as a variation or anomaly.
observed that both the left and right triceps brachii had
Comparative Anatomy
four heads, with the fourth head originating from the lat-
eral aspect of the humerus inferior to the deltoid, with fbers While typically absent in adult humans, epitrochleoanco-
extending to the medial condyle and the olecranon process neus is present often in several primate species including
of the ulna on the right side. These authors also observed common chimpanzees and bonobos (Gratiolet and Alix
slips between triceps brachii and teres major and latissimus 1866; Macalister 1871, 1875; Miller 1952; Ziegler 1964;
dorsi on the left side. Swindler and Wood 1973; Diogo and Wood 2011, 2012a;
In a male fetus with triploidy, Moen et al. (1984) found Diogo et al. 2013a, 2017). The presentation and innervation
that the long head of triceps brachii was doubled on the in nonhuman primates are the same as in humans. It is usu-
right side. Triceps brachii has been observed to have a low ally not present as a distinct muscle in gibbons, orangutans,
origin of the medial head in infants with Neu-Laxova syn- or gorillas (Diogo et al. 2010, 2012, 2013b).
drome (Shved et al. 1985). In a study of six upper limbs
Variations
from three infants with Neu-Laxova syndrome, Shved et al.
(1985) found that triceps had a low origin of the medial Description
head in all six upper limbs (100%). Epitrochleoanconeus (Gruber) is typically only present in
the early stages of human development, appearing as a dis-
Prevalence tinct muscle at CR25 mm (crown-rump length of 25 mm)
and persisting until at least CR33.5 mm (Diogo et al. 2019).
In their literature review, Smith et al. (2015) found that tri-
When present as a distinct muscle in adults, epitrochleoan-
ceps brachii had a more proximal origin than usual in 2 out
coneus extends from the medial epicondyle of the humerus
of 24 individuals with trisomy 13 (8.3%). Triceps brachii
to the medial side of the olecranon process of the ulna
was also variable (with extensive and/or variable origins,
(Gruber 1867a; Galton 1874; Macalister 1875; Knott 1883a;
or accessory slips) in 18 out of 26 individuals with trisomy
Jouffroy 1971; Lewis 1989; Akita and Nimura 2016a,b).
18 (69.2%). In a study of six upper limbs from three infants
Its origin may present as narrow, wide and feshy, or ten-
with Neu-Laxova syndrome, Shved et al. (1985) found that
dinous (Macalister 1875). It may sometimes be fused with
triceps had a low origin of the medial head in all six upper
anconeus (Bergman et al. 1988; Akita and Nimura 2016a).
limbs (100%).
It may also be doubled (Macalister 1875).
Clinical Implications Innervation

The presence of a fourth head of triceps brachii can con- Epitrochleoanconeus is innervated by the ulnar nerve
tribute to radial nerve palsy, vascular compromise due to (Howell and Strauss 1932; Miller 1952; Kallner 1956;
the compression of the deep radial artery, profunda bra- Ziegler 1964; Akita and Nimura 2016a,b).
chii artery compression, and snapping elbow (Fabrizio and
Clemente 1997; Tubbs et al. 2006b). A muscular fascicle Prevalence
connecting the long head and the lateral head may lead to Gruber (1867a) reports a prevalence of 34 out of 100 cadavers
radial nerve entrapment (Prabhu et al. 2012). (34%). Galton (1874) suggests that the epitrochleoanconeus
FIGURE 3.11 Posterior view of latissimus dorsi and posterior humeral variations. (Figure adapted by permission from Springer
Nature Customer Service Centre GmbH: Springer Current Molecular Biology Reports, Muscles Lost in Our Adult Primate Ancestors
Still Imprint in Us: on Muscle Evolution, Development, Variations, and Pathologies. E. Boyle, V. Mahon, R. Diogo, 2020.)
is found in about 53 out of 200 human upper limbs (26.5%). Clinical Implications
Macalister (1875) found this muscle in 16 out of 63 subjects The presence of epitrochleoanconeus can lead to ulnar
(25.4%). Le Double (1897) reports its presence in 32 out compression neuropathy at the elbow (Masear et al. 1988).
of 102 cadavers (31%). Clemens (1957) found the muscle Wilson et al. (2016) suggest that the presence of this muscle
in 4 out of 100 cadavers (4%). From a sample of 96 arms, may protect against the development of cubital tunnel syn-
Mori (1964) reported a prevalence of 5%. In a study of 218 drome, as it may decrease the rigidity of the entrance into
Brazilian adults, Nascimento and Ruiz (2018) found that the cubital tunnel, replacing Osborne’s ligament as the roof
this muscle was present in 29 cases (13.3%). of this tunnel.
Anomalies
Description ANTERIOR FOREARM MUSCLES
Anomalous presentation is same as above, running from the
back of the medial condyle to the medial side of the olecra- flexOr pOllicis lOngus (figure 3.12)
non (Smith et al. 2015). See also: Gantzer’s muscle
Prevalence
In their literature review, Smith et al. (2015) found that epi- Synonyms
trochleoanconeus was reported in 2 out of 26 individuals This muscle is also referred to as fexor longus proprius
with trisomy 18 (7.7%). pollicis (Todd 1839).
FIGURE 3.12 Deep anterior forearm muscles in anterior view.
Typical Presentation extends from the radius and interosseous membrane to digit
Description one and/or digit two (Diogo et al. 2012, 2017).
Flexor pollicis longus originates from the radius and the
interosseous membrane and inserts via a tendon onto the Variations
base of the distal phalanx of the frst digit (Standring 2016). Description
Flexor pollicis longus can also arise via a variable acces-
Innervation sory head from the coronoid process or from the medial
Flexor pollicis longus is innervated by the anterior interos- epicondyle of the humerus (Macalister 1875; Knott 1883a;
seous branch of the median nerve (Standring 2016). Sato 1969; Akita and Nimura 2016b; Standring 2016).
Flexor pollicis longus may also originate from brachia-
Comparative Anatomy lis (Macalister 1875; Bergman et al. 1988). Other varia-
Although it has been suggested that fexor pollicis longus tions include connections to fexor digitorum superfcialis,
is unique to humans (e.g., Bergman et al. 1988; Akita and fexor digitorum profundus, pronator teres, and the syno-
Nimura 2016b), it is also present in gibbons as a distinct vial bursa (a slip termed tensor bursae mucosae tendinum)
muscle separate from fexor digitorum profundus that (Macalister 1875; Knott 1883a; Mori 1964; Bergman et
1988; Akita and Nimura 2016b). Flexor pollicis longus distal phalanx. This muscle was also connected to flexor
may send a tendon to the first lumbrical (Macalister 1875; digitorum profundus. In a trisomy 18 cyclopic fetus, flexor
Bergman et al. 1988). Flexor pollicis longus can be partially pollicis longus had a bifurcated tendon with insertions into
or completely absent (Macalister 1875; Akita and Nimura the base of the proximal phalanx and the distal phalanx of
2016b; Standring 2016). Other variations of flexor pollicis digit one (Smith et al. (2015).
longus described by Macalister (1875) include the presence Mieden (1982) describes two male fetuses with cyclopia
of accessory slips and doubling of the muscle or its tendon. and alobar holoprosencephaly. On the left side of one speci-
Linburg-Comstock variation is described as a tendinous men, an extra muscle belly originated from flexor pollicis
connection between flexor pollicis longus and the digit two longus and fused with the tendon of flexor digitorum pro-
slip of flexor digitorum profundus, or the tendons of these fundus going to digit two. On the left arm of a male neonate
muscles (Linburg and Comstock 1979). Flexor pollicis lon- with Meckel syndrome, Pettersen (1984) observed a small
gus is also associated with Gantzer’s muscle (see the entry slip that extended between the tendon of brachioradialis
for this muscle). and flexor pollicis longus at the wrist.
In a sample of 34 subjects, Knott (1883a) found an acces- A presentation similar to that of the trisomy 18 cyclopic
sory head from the medial epicondyle in two cases (5.9%). fetus examined by Smith et al. (2015, see above) was also
A head from the coronoid was present in some form in 18 recorded in 2 out of 24 individuals with trisomy 13 (8.3%)
cases (52.9%): arising separately in nine cases (26.5%), and 4 out of 26 individuals with trisomy 18 (15.4%). Flexor
arising with the deep head of pronator teres in three cases pollicis longus was otherwise variable (e.g., tendinous or
(8.8%), closely associated with the flexor digitorum super- aponeurotic attachments to the first metacarpal, split ten-
ficialis in four cases (11.8%), and via a slip common to all dons, tendon to the second digit) in 2 out of 24 individuals
three muscles in two cases (5.9%). with trisomy 13 (8.3%), in 13 out of 26 individuals with
Mori (1964) found that there was a muscular bundle trisomy 18 (50%), and in 1 out of 7 individuals with trisomy
from flexor pollicis longus inserting into flexor digitorum 21 (14.3%). Flexor pollicis longus was also absent in 1 out
profundus in 12.5% of cases. These two muscles were con- of 26 individuals with trisomy 18 (3.8%).
nected with a muscular bundle in 4.5% of cases. The ten-
dons of these muscles were fused upon insertion into digit Clinical Implications
two in 7.5% of cases. The origin of flexor pollicis longus Shrewsbury et al. (2003) explain that the clinical evidence of
from the interosseous membrane was absent in 7.5% of restrictive thumb/index tendosynovitis and pain in humans
cases. Sato (1969) found an accessory head of flexor pol- is associated with connections between the flexor pollicis
licis longus present in 83 out of 356 limbs (23.31%) from longus and the flexor digitorum profundus (e.g., Linburg-
Kyushu-Japanese males and present in 68 out of 248 limbs Comstock variations).
(27.42%) from females.
Linburg and Comstock (1979) note that in a large sample Gantzer’s Muscle (Figure 3.12)
of humans, only 69% of them had a flexor pollicis longus
fully independent from the flexor digitorum profundus. Synonyms
Yurasakpong et al. (2018) studied 130 upper limbs and This muscle is also known as Gantzer’s (also Gantzner’s
found that the tendons of flexor pollicis longus and flexor or Ganzer’s) fasciculus, accesorius ad pollicem, flexor
digitorum profundus were connected in 32 limbs (24.6%) digitorum profundus accessorius, or accessorius ad flexo-
and varied between fibrous, tendinous, and musculotendi- rem profundus (Macalister 1875; Knott 1883a; Mori 1964;
nous connections. Erić et al. (2019) clinically diagnosed Bergman et al. 1988).
Linburg-Comstock variation via two tests (inability to flex
the thumb without simultaneous flexion of the index finger; Typical Presentation
pain during active flexion of the thumb while long fingers This muscle is only present as a variation or anomaly.
were fully extended, and the wrist was in supination) in 130
out of 215 subjects (60.5%). Comparative Anatomy
N/A
Anomalies
Bersu et al. (1976) describe a male infant with Hanhart syn- Description
drome. On the right side of this specimen, the wrist and Gantzer’s muscle is associated with flexor pollicis longus,
hand were deficient. Flexor pollicis longus inserted onto flexor digitorum profundus, and flexor digitorum super-
the trapezium. In a fetus with craniorachischisis, Alghamdi ficialis. It is generally described as an accessory head of
et al. (2017) observed that flexor pollicis longus on the left flexor pollicis longus. It can arise from the medial epicon-
upper limb had an insertion onto the proximal phalanx of dyle, the coronoid process, from both of these structures,
the first digit, in addition to the normal attachment to the or from flexor digitorum superficialis (Macalister 1875;
Knott 1883a; Mori 1964; Bergman et al. 1988; Akita and Pai et al. 2008b; Kulkarni et al. 2014; Caetano et al. 2015;
Nimura 2016b). Zdilla et al. (2019) reported an origin from Zdilla et al. 2019). It may also simulate a soft-tissue tumor
the brachialis fascia. It inserts onto the tendons or muscle or a ganglion (Kulkarni et al. 2014).
bellies of fexor pollicis longus and/or fexor digitorum pro-
fundus (Macalister 1875; Knott 1883a; Mori 1964; Bergman
flexOr DigitOruM prOfunDus (figure 3.12)
et al. 1988; Caetano et al. 2015; Akita and Nimura 2016b).
See also: Gantzer’s muscle
Innervation
Gantzer’s muscle is innervated by the anterior interosseous Synonyms
nerve (Kulkarni et al. 2014; Caetano et al. 2015; Zdilla et al. This muscle is also known as fexor digitorum perforans
2019). (Wood) (Macalister 1875), musculus perforans, cubito-
phalangettien commun (Todd 1839).
Prevalence
Macalister (1875) reports that Gantzer’s muscle can be Typical Presentation
found in about two out of fve subjects (40%). Mori (1964) Description
found this muscle in 50% of cases. In a study of 80 limbs Flexor digitorum profundus originates from the anterior
from 40 cadavers, Caetano et al. (2015) observed that this and medial surfaces of the ulna and the interosseous mem-
muscle was present in 54 limbs (67% of limbs). Kulkarni brane (Standring 2016). The muscle converges into four
et al. (2014) state that prevalence ranges from 39% to 90% tendons that pass through the tendons of fexor digitorum
in the literature, with an average of about 67%. superfcialis to insert onto the bases of the distal phalanges
Mori (1964) found that Gantzer’s muscle arose from of digits two through fve (Standring 2016).
the medial epicondyle in 24% of cases, from the coronoid
process in 68% of cases, and from both these structures in Innervation
8% of cases. In a sample of 80 limbs, Caetano et al. (2015) Innervation for this muscle is split; the portion that sends
observed that this muscle originated from the medial epi- tendons to digits four and fve is innervated by the ulnar
condyle in about 9% of limbs, from the coronoid process in nerve while the portion that sends tendons to digits two and
10% of limbs, and from the deep surface of fexor digitorum three is innervated by the anterior interosseous branch of
superfcialis in about 53% of limbs. the median nerve (Standring 2016).
Mori (1964) also observed variation in how the insertion
tendon of fexor pollicis longus relates to Gantzer’s muscle. Comparative Anatomy
The tendon arose from Gantzer’s muscle in 40% of cases, Flexor digitorum profundus has a similar typical presenta-
arose from both fexor pollicis longus and Gantzer’s muscle tion in the apes, extending from the radius, ulna, and inter-
in 40% of cases, arose from the point of fusion of fexor pol- osseous membrane to digits two through fve (Gibbs 1999;
licis longus and Gantzer’s muscle in 16% of cases, and both Diogo et al. 2010, 2012, 2013a,b, 2017). In some cases, a
muscles fused to form one tendon in 4% of cases. Caetano poorly developed tendon to digit one is present in common
et al. (2015) observed that this muscle inserted into fexor chimpanzees, orangutans, and gorillas (Gibbs 1999; Diogo
pollicis longus in 26 out of 80 limbs (45%) and into fexor et al. 2010, 2013a,b, 2017). In bonobos, the tendon to digit
digitorum profundus in 21 out of 80 limbs (26.3%). one was well-developed, stout and continuous with the main
body of the muscle in two specimens, and absent entirely in
Anomalies another specimen (Diogo et al. 2017). Bergman et al. (1988)
Description suggest that the distinct separation of the part of the muscle
Anomalous presentation is similar to the above, running that acts on the second digit is unique to humans, relating to
from the medial epicondyle to fexor pollicis longus (Smith the functional specialization of the index fnger.
et al. 2015).
Variations
Smith et al. (2015) noted that fexor digitorum superfcialis The part of the muscle that inserts onto the second digit
sent an accessory slip to fexor pollicis longus in 3 out of 24 is distinct from origin to insertion (Bergman et al. 1988;
individuals with trisomy 13 (12.5%), 8 out of 26 individu- Standring 2016). When completely distinct, it forms fexor
als with trisomy 18 (30.8%), and 1 out of 7 individuals with indicis profundus (Macalister 1875; Knott 1883a; Bergman
trisomy 21 (14.3%). et al. 1988). The tendon to the third digit may be absent
(Bergman et al. 1988). Flexor digitorum profundus is fre-
Clinical Implications quently joined by slips from other structures including
Gantzer’s muscle can contribute to median nerve and/or fexor digitorum superfcialis, fexor pollicis longus, the
anterior interosseous nerve palsy, entrapment, or pressure medial epicondyle, the coronoid process, or the radius
neuritis, as well as Kiloh-Nevin syndrome (Oh et al. 2000; (Macalister 1875; Mori 1964; Akita and Nimura 2016b;
Standring 2016). Other variations of fexor digitorum pro- Bersu et al. (1976) describe a male infant with Hanhart
fundus described by Macalister (1875) include the distinct syndrome. On the right side of this specimen, the wrist and
separation of fexor indicis profundus, accessory slips to hand were defcient. Flexor digitorum profundus inserted
and from other structures, and cases in which other fore- via a single tendon onto the capsule of the fused carpal
arm or hand muscles arose from this muscle or its tendons. mass with the single anomalous tendon of fexor digitorum
Linburg-Comstock variation is described as a tendinous superfcialis. The ffth tendon of fexor digitorum profun-
connection between fexor pollicis longus and the digit two dus arising from the fourth tendon has been observed in
slip of fexor digitorum profundus, or the tendons of these infants with Neu-Laxova syndrome (Shved et al. 1985). On
muscles (Linburg and Comstock 1979). Flexor digitorum the right hand of a male neonate with Meckel syndrome,
profundus is also associated with Gantzer’s muscle (see the Pettersen (1984) observed that fexor digitorum profundus
entry for this muscle). sent tendons to all six digits.
See fexor pollicis longus entry for prevalence information According to the literature review conducted by Smith
about Linburg-Comstock variation. Knott (1883a) found et al. (2015), fexor digitorum profundus was variable (e.g.,
fexor indicis profundus in 2 out of 34 subjects (5.9%). diminutive or missing tendons, fusion with fexor digitorum
Mori (1964) observed that fexor digitorum profundus had superfcialis, tendon to sixth digit when present, and vari-
a radial origin in 84% of cases. Mori (1964) also noted that able presentation in how and which tendons insert onto each
out of 205 arms, this muscle received a slip from the coro- digit) in 5 out of 24 individuals with trisomy 13 (20.8%) and
noid process in 40% of arms, from the medial epicondyle 6 out of 26 individuals with trisomy 18 (23.1%). In a study
in one case (about 0.5%), from the fexor pollicis longus in of six upper limbs from three infants with Neu-Laxova syn-
12.5% of arms, and its tendon was fused that of with fexor drome, Shved et al. (1985) found that the tendon of fexor
digitorum superfcialis in 11% of arms. Flexor digitorum digitorum profundus going to the ffth fnger arose from the
profundus and fexor pollicis longus were connected with tendon to the fourth fnger in two upper limbs (33%).
a muscular bundle in 4.5% of cases. The tendons of these
muscles were fused upon insertion into digit two in 7.5% of Clinical Implications
cases. Furnas (1965) found that the muscle belly and tendon Shrewsbury et al. (2003) explain that the clinical evidence of
going to digit two was doubled in 4 out of 117 cadavers restrictive thumb/index tendosynovitis and pain in humans
(3.4%). is associated with connections between the fexor pollicis
longus and the fexor digitorum profundus (e.g., Linburg-
Anomalies Comstock variations).
Description
In a fetus with craniorachischisis, Alghamdi et al. (2017) flexOr DigitOruM superficialis (figure 3.13)
observed that the tendon of the left fexor digitorum pro-
fundus that attached to the distal phalanx of the ffth digit See also: Gantzer’s muscle
also sent a small tendon to the middle phalanx of this digit.
Synonyms
Also, a slip connected this muscle to fexor pollicis longus.
On the right side of the body, fexor digitorum profundus This muscle is also referred to as fexor digitorum subli-
only sent tendons to digits four and fve. There were two mis (Standring 2016), musculus perforates, or epi-trochlo-
deeper bundles of fexor digitorum profundus deep to this phalanginien commun (Todd 1839).
main belly. The frst bundle sent one tendon to the forearm
fascia and a second tendon that joined the tendons of the
second bundle and fexor digitorum superfcialis going to Description
digit three. Windle (1893) described a slip running from the The humero-ulnar head of fexor digitorum superfcialis
inner condyle of the humerus to fexor digitorum profundus originates from the medial epicondyle of the humerus,
in two female fetuses with anencephaly. the coronoid process, the anterior band of the ulnar col-
In a neonate with trisomy 13, the left fexor digitorum lateral ligament, and the intermuscular septa (Standring
profundus was divided into three fascicles (Aziz 1980). In 2016). The radial head originates from the middle third
a female fetus with trisomy 18, the tendons of fexor digito- of the radius (Standring 2016). The heads converge into a
rum profundus on the left side split more proximally than muscle with two layers; the superfcial layer sends tendons
is normal (Alghamdi et al. 2018). On the right side, fexor to the third and fourth digits, while the deep layer sends
digitorum profundus had two bellies that were fused proxi- tendons to the second and ffth digits (Standring 2016).
mally. The lateral belly sent one tendon to digit two and The tendons for each digit enter the fexor sheaths, split
the medial belly sent three tendons to the three medial dig- to course around the tendons of fexor digitorum profun-
its. The lateral belly was fused proximally to fexor pollicis dus, and insert onto the intermediate phalanx of each digit
longus. (Standring 2016).
FIGURE 3.13 Flexor digitorum superfcialis, pronator quadratus, radiocarpus, and accessory radiocarpus in anterior view.
Innervation tendons may be replaced by slips from other structures

Flexor digitorum superfcialis is innervated by the median (Macalister 1875; Bergman et al. 1988; Akita and Nimura
nerve (Standring 2016). 2016b; Standring 2016). There is also frequent variation in
how the tendons supply each digit (Macalister 1875; Furnas
Comparative Anatomy 1965; Kaplan 1969; Shrewsbury and Kuczynski 1974;
Bergman et al. 1988; Akita and Nimura 2016b). Flexor digi-
Flexor digitorum superfcialis has a similar typical pre-
torum superfcialis can be connected via slips to fexor pol-
sentation in the apes, extending from the radius, ulna, and
licis longus, palmaris longus, the brachioradialis tendon, or
medial epicondyle to the intermediate phalanges of dig-
the ulnar tuberosity, or be fused with fexor pollicis longus,
its two through fve (Gibbs 1999; Diogo et al. 2010, 2012,
fexor digitorum profundus and its tendons, or pronator
2013a,b, 2017). In all apes, the ulnar origin may be occasion-
teres (Macalister 1875; Mori 1964; Bergman et al. 1988;
ally absent (Macalister 1873; Deniker 1885; Beddard 1893;
Akita and Nimura 2016b). Other variations of fexor digi-
Loth 1931; Kallner 1956; Gibbs 1999; Diogo et al. 2010,
torum profundus described by Macalister (1875) include
2012, 2013a,b, 2017). The radial origin may be absent in
variation in tendon insertion, tendon doubling or absence,
gibbons, orangutans, and common chimpanzees (Gratiolet
the presence of accessory slips, and presentation as a digas-
and Alix 1866; Deniker 1885; Michaelis 1903; Diogo et al.
tric muscle.
2012, 2013b).
Flexor digitorum superfcialis is associated with three
supernumerary muscles. Radiopalmaris originates from
Variations
the radius and attaches to the palmar aponeurosis and digi-
Description tal fexor sheath (Bergman et al. 1988; Akita and Nimura
The radial head of fexor digitorum superfcialis and 2016b). Palmar fexor digitorum superfcialis accessorius
the insertions onto the third or ffth digit may be absent originates from the palmar fascia and transverse carpal
(Macalister 1875; Knott 1883a; Mori 1964; Furnas 1965; ligament and ends in a tendon that joins the fexor tendon
Shrewsbury and Kuczynski 1974; Bergman et al. 1988; going to digit two (Bergman et al. 1988; Akita and Nimura
Akita and Nimura 2016b; Standring 2016; Yammine and 2016b). Flexor digitorum superfcialis is also associated
Erić 2018). Occasionally, the fexor digitorum superfcialis with Gantzer’s muscle (see the entry for this muscle).
Prevalence and hand were defcient. Flexor digitorum superfcialis

Mori (1964) noted that the radial head of fexor digitorum inserted via a single tendon associated with palmaris lon-
superfcialis was absent in 7.5% of his subjects and diminu- gus onto the connective tissue capsule of the fused carpal
tive in 12.5%. Mori (1964) also observed that this muscle mass. Mieden (1982) describes an infant with median cleft
was fused with pronator teres in 2.5% of the individuals lip, hypotelorism, and alobar holoprosencephaly. The ten-
examined by him, while it was fused with fexor digitorum don of fexor digitorum superfcialis to the ffth digit was
profundus in 8%, and with fexor pollicis longus in 45%. absent on the right side (Mieden 1982). On the left side of
Furnas (1965) found that the tendon and muscle belly of a male neonate with Meckel syndrome, Pettersen (1984)
fexor digitorum superfcialis going to digit fve was absent observed that the fexor digitorum superfcialis tendon to
in 6 out of 117 arms (5.1%). Kaplan (1969) found that in 21 the ffth digit was absent. An accessory muscle associated
out of 68 hands the fexor superfcialis tendons to digit fve with fexor digitorum superfcialis and ffth tendon arising
were connected with the tendon going to digit two (30.8%), from the fourth tendon has been observed in infants with
while in 23 hands the fexor superfcialis tendons to digit Neu-Laxova syndrome (Shved et al. 1985).
fve were connected with the tendon going to digit four
(33.8%). Shrewsbury and Kuczynski (1974) found that the Prevalence
tendon to digit fve was absent in seven of 23 hands (30%) Overall, fexor digitorum superfcialis was anomalous in
from 15 cadavers. many individuals included in the literature review conducted
A meta-analysis of 34 studies with a total sample of by Smith et al. (2015). This muscle was variable (e.g., dimin-
12,213 forearms by Yammine and Erić (2018) reveals that utive radial head, doubled muscle bellies or tendons, varia-
the clinical prevalence of functional absence of the fexor tion in presentation or absence of tendons, tendons replaced
digitorum superfcialis tendon to digit fve is 7.45%, the by other structures) in 6 out of 24 individuals with trisomy
clinical prevalence of the “common type” of the fexor digi- 13 (25%), 11 out of 26 individuals with trisomy 18 (42.3%),
torum superfcialis tendon to digit fve is 37.5%, and the and in 2 out of 7 individuals with trisomy 21 (28.6%). An
cadaveric prevalence of muscle belly absence for fexor absent radial head of this muscle was noted in 4 out of 24
digitorum superfcialis to digit fve is 2.5%. individuals with trisomy 13 (16.7%) and 2 out of 26 individ-
uals with trisomy 18 (7.7%). Flexor digitorum superfcialis
Anomalies sent a slip to fexor digitorum profundus in 3 out of 24 indi-
Description viduals with trisomy 13 (12.5%), 7 out of 26 individuals with
trisomy 18 (26.9%), and 2 out of 7 individuals with trisomy
In a neonate with trisomy 13, fexor digitorum superfcialis
21 (28.6%). These authors also noted that fexor digitorum
was divided into four fascicles bilaterally, and the tendon
superfcialis sent an accessory slip to fexor pollicis longus
to the ffth digit was absent on both sides (Aziz 1980). In a
in 3 out of 24 individuals with trisomy 13 (12.5%), 8 out
trisomy 18 cyclopic fetus dissected by Smith et al. (2015),
of 26 individuals with trisomy 18 (30.8%), and 1 out of 7
the authors observed that the left fexor digitorum superf-
individuals with trisomy 21 (14.3%). An accessory slip from
cialis lacked both its radial origin and its insertion onto the
fexor digitorum superfcialis to radiocarpus was present in
ffth digit. On the right side of this individual, fexor digi-
1 out of 24 individuals with trisomy 13 (4.2%) and 1 out of
torum superfcialis sent a slip to fexor digitorum profun-
26 individuals with trisomy 18 (3.8%).
dus (Smith et al. 2015). In a female fetus with trisomy 18,
In a study of six upper limbs from three infants with
the tendon of fexor digitorum superfcialis to the ffth digit
Neu-Laxova syndrome, Shved et al. (1985) found that an
was absent bilaterally (Alghamdi et al. 2018). The proximal
accessory muscle associated with fexor digitorum superf-
end of this muscle was also fused with that of fexor carpi
cialis was present in two upper limbs (33%) and the tendon
radialis on both sides of the body. Among individuals with
going to the ffth fnger arose from the tendon to the fourth
trisomy 21, the tendon to the ffth digit was absent on the
fnger in two upper limbs (33%).
left side in one fetus, both sides of another fetus, and on the
right side of an adult (Bersu 1980). There was a double belly
of fexor digitorum superfcialis to the ffth digit in a child
with trisomy 21 (Bersu 1980). A diminutive or absent fexor superfcialis tendon may
In one anencephalic fetus, Windle (1893) observed that impair grip strength and can also functionally present as
fexor digitorum superfcialis did not send a tendon to the a tendon rupture, which can be assessed by clinical tests
ffth digit. In a fetus with craniorachischisis, the right fexor (Bowman et al. 2003; Yammine and Erić 2018; Belbl et al.
digitorum superfcialis was diminutive and only sent ten- 2020).
dons to digits three and four (Alghamdi et al. 2017). On the
left side of the body, this muscle only inserted onto digits prOnatOr quaDratus (figure 3.13)
two, three, and four, and its proximal end was fused with
pronator teres and fexor carpi ulnaris. Synonyms
Bersu et al. (1976) describe a male infant with Hanhart This muscle is also referred to as pronator radii quadratus
syndrome. On the right side of this specimen, the wrist (Knott 1883a) or cubito-radial (Todd 1839).
Typical Presentation Anomalies

Pronator quadratus extends from the distal portion of the In a fetus with craniorachischisis, Alghamdi et al. (2017)
ulnar shaft to the distal portion of the radial shaft (Standring noted that pronator quadratus was absent on the right side.
2016). In a neonate with trisomy 18, “fexor carpi ulnaris brevis”
was present on the left side, extending from the distal ulna
Innervation to the hamate (Aziz 1979).
Pronator quadratus is innervated by the median nerve
(Standring 2016). Prevalence
In their literature review, Smith et al. (2015) found that pro-
Comparative Anatomy nator quadratus was absent in 1 out of 26 individuals with
Pronator quadratus has a similar typical presentation in the trisomy 18 (3.8%).
apes but appears to have a more oblique orientation in gib-
bons and orangutans (Gibbs 1999; Diogo et al. 2010, 2012, Clinical Implications
2013a,b, 2017). Dwight (1895) observed that pronator qua- N/A
dratus can be separated into multiple layers as a variation in
common chimpanzees.
raDiOcarpus (figure 3.13)
Variations See also: Pronator quadratus, Flexor carpi radialis brevis
Description
Pronator quadratus can be split into two or more parts Synonyms
or layers, and its fbers may run in different directions This muscle is also referred to as radiocarpeus, radiocar-
(Macalister 1875; Knott 1883a; Inoue 1934; Mori 1964; pien (Fano), or court radial anterieur (Le Double) (Bergman
Johnson and Shrewsbury 1976; Sakamoto et al. 2015; Akita et al. 1988; Akita and Nimura 2016b). Though Knott (1883a)
and Nimura 2016b; Das et al. 2008). Pronator quadratus and Akita and Nimura (2016b) list fexor carpi radialis bre-
may distally expand into the wrist or metacarpal region vis as a synonym of radiocarpus, we treat these muscles as
(Bergman et al. 1988; Akita and Nimura 2016b). It may also separate structures due to (1) the described tendinous and
expand proximally to connect with pronator teres or fexor muscular connections between fexor carpus radialis brevis
carpi radialis (Macalister 1875; Bergman et al. 1988; Akita and fexor carpi radialis and (2) the possibility of radiocar-
and Nimura 2016b). Its fbers can insert into the thenar pus extending proximally to the ulna or medial epicondyle.
eminence and act as an accessory adductor of the frst digit
(Bergman et al. 1988; Akita and Nimura 2016b). Pronator
quadratus can also be absent as a rare variation (Macalister This muscle is only present as a variation or anomaly.
1875; Bergman et al. 1988; Akita and Nimura 2016b).
Comparative Anatomy
Pronator quadratus is in close proximity to several super-
numerary muscles. Radiocubitocarpien (radialis internus bre- N/A
vis biceps) has an ulnar head and radial head and ends on the
Variations
wrist (Calori 1870; Bergman et al. 1988; Akita and Nimura
2016b). Tensor capsulae radiocubitalis (inferioris) originates Description
from the distal radius and courses over the anterior surface Radiocarpus is a supernumerary muscle that originates
of pronator quadratus to insert onto the radioulnar capsule from the radius proximal to pronator quadratus and may
(Bergman et al. 1988; Akita and Nimura 2016b). Ulnocarpus extend proximally to the ulna or medial epicondyle of the
brevis (fexor carpi ulnaris brevis or cubitocarpeus) origi- humerus (Macalister 1875; Bergman et al. 1988; Akita and
nates from the distal end of the ulna and inserts onto the pisi- Nimura 2016b). It inserts onto one or more carpal or meta-
form, hamate, fourth metacarpal, ffth metacarpal, capsule of carpal bones, or bones from both regions (Macalister 1875;
carpal articulations, or abductor digiti minimi (Knott 1883a; Bergman et al. 1988; Akita and Nimura 2016b). It may be
Mori 1964; Bergman et al. 1988; Akita and Nimura 2016b). divided into slips (Macalister 1875).
Knott (1883a) found that pronator quadratus was split com- N/A
pletely into two parts in 2 out of 34 subjects (5.9%). Mori
(1964) recorded the presence of supernumerary muscle Prevalence
“fexor carpi ulnaris brevis” in 4 out of 205 arms (1.9%). Macalister (1875) notes that Wood has reported radiocarpus
Mori (1964) also classifed pronator quadratus into eight present in 4 out of 34 subjects in one study (11.8%) and in 2
types and found that the typical presentation of this muscle out of 36 subjects (5.6%) in two studies. Over the course of
was only present in 39% of cases. three studies, Macalister (1875) found radiocarpus in 2 out
of 53 subjects (3.8%), 1 out of 60 subjects (1.7%), and 4 out 1892; Sommer 1907; Pira 1913; Sonntag 1923; Preuschoft
of 64 subjects (6.3%). 1965; Gibbs 1999; Diogo et al. 2010, 2013a, 2017).
In a neonate with trisomy 18, radiocarpus was present on the Macalister (1875), Knott (1883a), Bergman et al. (1988),
right side, extending from the proximal radius to the fexor and Akita and Nimura (2016b) note that palmaris longus is
retinaculum (Aziz 1979). Radiocarpus was present bilater- likely the most variable muscle in the human body. It can
ally in a neonate with trisomy 13 and extended from just be absent bilaterally or unilaterally, is more often absent
below the radial tuberosity to the trapezium (Aziz 1980). in females, and is more often absent on the left side of
Some anomalous cases exhibit an accessory radiocar- the body (Macalister 1875; Knott 1883a; Le Double 1897;
pus that originates from the underside of fexor digitorum Reimann et al. 1944; George 1953; Sato 1969; Hall 1984;
superfcialis and inserts onto the belly of the radiocarpus Bergman et al. 1988; Akita and Nimura 2016b; Standring
(Figure 3.13) (Smith et al. 2015). 2016; Georgiev et al. 2017b). Wilde et al. (2021) suggest that
the absence of palmaris longus in adults is the result of the
Prevalence
muscle failing to form early in development.
In their literature review, Smith et al. (2015) found that Both its origin and insertion are variable. Palmaris lon-
radiocarpus was present in 5 out of 24 individuals with tri- gus can originate from the medial intermuscular septum,
somy 13 (20.8%) and in 5 out of 26 individuals with trisomy biceps, brachialis, the fascia of the forearm, the coronoid
18 (19.2%). The accessory radiocarpus was present in 2 out process, or the radius (Macalister 1875; Knott 1883a;
of 24 individuals with trisomy 13 (8.3%). An accessory slip Bergman et al. 1988; Akita and Nimura 2016b). It can
from fexor digitorum superfcialis to radiocarpus was pres- insert onto the interosseous membrane, forearm fascia,
ent in 1 out of 24 individuals with trisomy 13 (4.2%) and 1 the fexor carpi ulnaris tendon, abductor pollicis brevis,
out of 26 individuals with trisomy 18 (3.8%). the hypothenar eminence, any one of the fexor tendons, the
pisiform, scaphoid, or digit four (Macalister 1875; Knott
1883a; Bergman et al. 1988; Akita and Nimura 2016b). The
N/A proportion of feshy muscle belly to tendon in palmaris lon-
gus is variable (Macalister 1875; Knott 1883a; Bergman
palMaris lOngus (figure 3.14) et al. 1988; Akita and Nimura 2016b). The tendon and mus-
cle can be inverted so that palmaris longus is feshy distally
Synonyms and the tendon is proximal, in which the muscle would be
This muscle corresponds to palmaris longus externus sensu termed palmaris longus inversus (Macalister 1875; Mori
(Jouffroy 1971) and is also referred to as epitrochlo-pal- 1964; Bergman et al. 1988; Akita and Nimura 2016b;
maire (Todd 1839). Georgiev et al. 2017b).
Palmaris longus can be doubled or tripled (accessory
Typical Presentation palmaris longus), or either the muscle belly or tendon can
Description be split (Macalister 1875; Reimann et al. 1944; Bergman
Palmaris longus originates from the medial epicondyle of et al. 1988; Akita and Nimura 2016b; Georgiev et al. 2017b).
the humerus (Standring 2016). Some fbers attach onto the It is also associated with two supernumerary muscles.
fexor retinaculum as its tendon of insertion passes into the Palmaris profundus originates from the middle third of the
hand to join with the palmar aponeurosis (Standring 2016). radius and courses beneath the fexor retinaculum to insert
with the palmar aponeurosis (Bergman et al. 1988; Akita
Innervation and Nimura 2016b). Accessorius ad fexorem digiti minimi
Palmaris longus is innervated by the median nerve refers to a muscle that arises from the palmaris longus
(Standring 2016). tendon to insert onto the ffth metacarpal (Bergman et al.
1988).
Comparative Anatomy
Palmaris longus is typically present in gibbons and orang- Prevalence
utans, extending from the medial epicondyle of the humerus Macalister (1875) notes that palmaris longus deviates from
to the palmar aponeurosis (Chapman 1880; Kohlbrügge the typical presentation in about every one out of four
1890–1892; Hepburn 1892; Beddard 1893; Primrose 1899, cases (25%). The rate of absence of palmaris longus has
1900; Grönroos 1903; Michaelis 1903; Sonntag 1924; Loth been reported to be anywhere between about 2%–25%
1931; Kallner 1956; Michilsens et al. 2009; Oishi et al. 2009; (Hall 1984; Bergman et al. 1988). Bergman et al. report an
Diogo et al. 2012, 2013b). It is sometimes absent in common absence of 11.2% across all bodies and cite a study of 800
chimpanzees and bonobos and typically absent in gorillas subjects in which it was absent on both sides in 7.7% of
(Chapman 1878; Bischoff 1880; Deniker 1885; Hepburn cases, absent on the right side in 4.5% of cases, and absent
FIGURE 3.14 Superfcial anterior forearm muscles in anterior view. This fgure displays the anomalous presentation of fexor carpi
radialis brevis as described by Smith et al. (2015).
on the left side in 5.2% of cases. Bergman et al. (1988) also muscle was absent in 205 arms (12.8%), which is close to
note that prevalence of absence is lower in Asian popula- the prevalence of 12.7% (178 out of 1,400 cases) reported
tions, as it is absent in 3.4% of Japanese individuals (30 out by Gruber (Reimann et al. 1944). These authors also found
of 884 arms; Adachi 1909) and 2.2% of Chinese individuals that palmaris longus showed variations other than absence
(two out of 95 arms; Nakano 1923). Knott (1883a) found the in 46 out of 530 arms (9%). Palmaris longus was doubled in
muscle absent in 4 out of 34 subjects (11.8%) and found an 4 out of 530 arms (0.8%). Mori (1964) noted that an inverted
accessory head from the coronoid process in two subjects palmaris longus was present in 3 out of 360 arms (0.8%),
(5.9%). Le Double (1897) reports an absence in 91 out of the tendon of insertion ended in the forearm fascia in 3 out
520 cases (17.5%). In a study of 276 cadavers by George of 205 arms (1.5%), the tendon of insertion was split into
(1953), palmaris longus was absent in 15.2% of the 552 two fascicles in three or 4 out of 40 arms (10%), and the
limbs examined. Sato (1969) found palmaris longus absent tendon of insertion was divided into three fascicles in 1 out
in 33 out of 406 limbs from Kyushu-Japanese males (8.13%) of 40 arms (2.5%).
and absent in 6 out of 262 limbs (2.29%) from females. Through clinical testing of 7,000 teenagers in Turkey,
Reimann et al. (1944) conducted an extensive examina- Ceyhan and Mavt (1997) report an absence of 63.9%.
tion of palmaris longus in 1,600 arms. They found that this Clinical testing of 300 Caucasian subjects revealed
that palmaris longus was absent unilaterally in 49 sub- upper limbs from three infants with Neu-Laxova syndrome,
jects (16.3%) and absent bilaterally in 26 subjects (8.6%) Shved et al. (1985) found that palmaris longus was absent in
(Thompson et al. 2001). Through clinical testing of 800 four upper limbs (66%) and an “anomalous structure” was
subjects in Serbia, Erić et al. (2010) determined that pal- present in two upper limbs (33%).
maris longus was absent unilaterally in 173 subjects (21.6%) According to Smith et al. (2015), palmaris longus was
and absent bilaterally in 127 subjects (15.9%). Clinical test- absent in 20 out of 24 individuals with trisomy 13 (83.3%),
ing of 200 subjects in Marathwada, India, found that this 23 out of 26 individuals with trisomy 18 (88.5%), and in 6
muscle was absent unilaterally in 20 individuals (10%) out of 7 individuals with trisomy 21 (85.7%). This muscle
and absent bilaterally in 11 individuals (6%) (Hussaini and was also variable (diminutive in one specimen and variable
Deshmukh 2018). insertion onto metacarpal fve in another) in 2 out of 26
Georgiev et al. (2017b) studied palmaris longus anatomy individuals with trisomy 18 (7.7%) and was variable (dimin-
in 112 limbs from 56 cadavers. They found that this muscle utive in one specimen and distally located belly in another)
was only variable in nine cases (8%). Palmaris longus was in 2 out of 7 individuals with trisomy 21 (28.6%) (Smith
absent in three cases (2.7%), doubled in two cases (1.8%), et al. 2015).
had an intermediate muscle belly in two cases (1.8%), was
digastric in one case (0.9%), and was reversed and associ- Clinical Implications
ated with an accessory abductor digiti minimi in one case Palmaris longus inversus can lead to forearm pain and
(0.9%) (Georgiev et al. 2017b). swelling, compression and paresthesia of the median or
ulnar nerve, and carpal tunnel syndrome (Backhouse and
Anomalies
Churchill-Davidson 1975; Still and Kleinert 1973; Depuydt
Description et al. 1998; Bhashyam 2017). Tiengo et al. (2006) note that
Palmaris longus was absent bilaterally in two neonates with the presence of an accessory palmaris longus can emulate
trisomy 18 and three neonates with trisomy 13 (Aziz 1979, a tumor in medical images and also contribute to median
1980, 1981). In a female fetus with trisomy 18, palmaris lon- nerve compression or carpal tunnel syndrome. Palmaris
gus was absent on the right side (Alghamdi et al. 2018). In profundus can cause compressive neuropathy of the ante-
a trisomy 18 cyclopic fetus, palmaris longus was absent on rior interosseous nerve and the median nerve (Akita and
one side (Smith et al. (2015). Palmaris longus was absent Nimura 2016b).
bilaterally in four individuals with trisomy 21 described by
Bersu (1980). In a ffth case (a fetus), the muscle belly was
flexOr carpi ulnaris (figure 3.14)
situated at the wrist in the right forearm and at the middle
of the forearm in the left forearm. Synonyms
In a male fetus with triploidy, Moen et al. (1984) found This muscle is also referred to as ulnaris internus, cubital
that palmaris longus was absent on the left side. The interne, or cubito-carpien (Todd 1839).
absence or “anomalous structure” of palmaris longus
has been observed in infants with Neu-Laxova syndrome Typical Presentation
(Shved et al. 1985). In a fetus with craniorachischisis,
Description
Alghamdi et al. (2017) observed that the palmaris longus
was absent on the left side. On the right side of the body, The humeral head of fexor carpi ulnaris originates from
palmaris longus was fused with fexor carpi ulnaris at its the medial epicondyle, and the ulnar head originates the
proximal end. proximal portion of the posterior ulna, including the olec-
Bersu et al. (1976) describe a male infant with Hanhart ranon (Standring 2016). It has insertions into the pisiform,
syndrome. On the right side of this specimen, the wrist hamate, and ffth metacarpal (Standring 2016).
and hand were defcient. Palmaris longus inserted onto the
Innervation
radial side of the reduced fexor digitorum superfcialis ten-
don just proximal to the fexor retinaculum. Mieden (1982) Flexor carpi ulnaris is innervated by the ulnar nerve
describes an infant with median cleft lip, hypotelorism, and (Standring 2016).
alobar holoprosencephaly (case I) and two male fetuses
with cyclopia and alobar holoprosencephaly (cases II and Comparative Anatomy
III). In cases I and II, palmaris longus was absent bilaterally The insertion of fexor carpi ulnaris is confned to the pisi-
(Mieden 1982). form in gibbons, orangutans, and gorillas (Diogo et al.
2010, 2012, 2013a,b, 2017). The presentation of fexor carpi
Prevalence ulnaris in common chimpanzees and bonobos is more
Among the limbs of ten anencephalic fetuses, Windle (1893) similar to that of humans, as it can have attachments to
noted that palmaris longus was absent unilaterally in two structures beyond that of the pisiform including the ffth
males and missing bilaterally in one male (30%), while this metacarpal and the hamate (Gratiolet and Alix 1866; Miller
muscle was doubled in one female (10%). In a study of six 1952; Diogo et al. 2013a, 2017).
Variations Flexor carpi radialis (Figure 3.14)

A slip may connect flexor carpi ulnaris to the coronoid This muscle is also referred to as flexor carpi radialis lon-
process (Bergman et al. 1988; Akita and Nimura 2016b; gus (Mori 1964), radialis internus, grand palmaire, or radial
Standring 2016). Flexor carpi ulnaris can also exhibit attach- anterieur (Todd 1839).
ments to the flexor retinaculum, the third or fourth metacar-
pals, the metacarpophalangeal joint capsule of the fifth digit,
or the abductor digiti minimi (Macalister 1875; Knott 1883a;
Bergman et al. 1988; al-Qattan and Duerksen 1992; Akita and Description
Nimura 2016b; Standring 2016). It may originate from triceps Flexor carpi radialis originates from the medial epicondyle
brachii (Macalister 1875). Bergman et al. (1988) describe a of the humerus (Standring 2016). It inserts via a long tendon
rare variation in which an accessory belly is present infe- onto the base of the second metacarpal and sends a slip to
rior to the main belly of this muscle and is innervated by the the third metacarpal (Standring 2016).
median nerve. Both Sakthivel and Verma (2017) and Kunc
et al. (2019) report an accessory flexor carpi ulnaris associated Innervation
with an absent palmaris longus, though in the former study Flexor carpi radialis is innervated by the median nerve
the accessory muscle was innervated by the ulnar nerve and (Standring 2016).
no innervation information was provided in the latter study.
Comparative Anatomy
Prevalence An origin from the radius is frequently present in all apes
Mori (1964) did not find any variation in origin or (Gibbs 1999; Diogo et al. 2010, 2012, 2013a,b, 2017). Some
insertion of flexor carpi ulnaris but noted that in 6% of cases individuals may lack an attachment to the third metacarpal
the muscular part extended distally past the radiocarpal joint. (Gratiolet and Alix 1866; Kohlbrügge 1890–1892; Beddard
1893; Dwight 1895; Primrose 1899, 1900; Sullivan and
Osgood 1927; Gibbs 1999; Miller 1952; Michilsens et al.
Anomalies
2009; Diogo et al. 2010, 2012, 2013a,b, 2017). An inser-
Description tion into the trapezium may be found in gorillas (Preuschoft
Macalister (1875) notes that Koster observed an insertion 1965; Diogo et al. 2010). In bonobos, flexor carpi radialis
of flexor carpi ulnaris into the interosseous membrane in an may fuse with pronator teres (Miller 1952) or the flexor
individual with a congenital deformity and that Friedlowsky digitorum superficialis tendon to digit four (Diogo et al.
found a slip from flexor carpi ulnaris to the annular ligament 2017).
in a hand that had only a thumb and two fingers. Bersu et al.
(1976) describe a male infant with Hanhart syndrome. On Variations
the right side of this specimen, the wrist and hand were defi- Description
cient. Flexor carpi ulnaris inserted onto the ulnar aspect of Flexor carpi radialis may be completely absent (Mori 1964;
the fused carpal mass. In a fetus with craniorachischisis dis- Bergman et al. 1988; Akita and Nimura 2016b; Standring
sected by Alghamdi et al. (2017), the authors observed that 2016). It may be doubled (Mori 1964; Akita and Nimura
the left flexor carpi ulnaris had two heads like the normal 2016b). Accessory slips or accessory heads may connect
condition, but the ulnar nerve separated abnormally between this muscle to brachialis, the biceps tendon, the bicipital
the two heads. Additionally, the proximal end of flexor carpi aponeurosis, coronoid process, or radius (Macalister 1875;
ulnaris was fused with pronator teres and flexor digitorum Knott 1883a; Mori 1964; Bergman et al 1988; Akita and
superficialis. On the right side of the body, flexor carpi ulna- Nimura 2016b; Standring 2016). It may have insertions into
ris was abnormally fused with palmaris longus near where the flexor retinaculum, scaphoid, trapezium, or fourth meta-
these muscles arise from the medial epicondyle. Smith carpal (Macalister 1875; Knott 1883a; Bergman et al 1988;
et al. (2015) noted the presence of an accessory belly, termed Akita and Nimura 2016b; Standring 2016).
flexor carpi ulnaris accessorius, which originated from the
middle of the ulna and inserted with flexor digiti minimi Prevalence
onto the fifth digit in a fetus with trisomy 18 and cyclopia. In a sample of 34 specimens, Knott (1883a) found that flexor
carpi radialis inserted into the second and third m etacarpals
Prevalence in 19 cases (55.9%), into the trapezium in one case (2.9%), into
Flexor carpi ulnaris accessorius was present in 9 out of 26 the trapezium and second metacarpal in one case (2.9%), and
individuals with trisomy 18 (34.6%). into the third and fourth metacarpal in one case (2.9%). Mori
(1964) noted that flexor carpi radialis had a radial origin in
Clinical Implications 7.5% of cases, received a slip from the coronoid process in
Variations of flexor carpi ulnaris and its tendon of inser- 5% of cases, was doubled in 2% of cases, and was entirely
tion can cause ulnar nerve compression (O’Hara and Stone absent in 2.5% of cases. Mori (1964) also observed that
1988; al-Qattan and Duerksen 1992). flexor carpi radialis inserted onto the second metacarpal in
90% of cases and sent a slip to the third metacarpal in 7.5% Typical Presentation
of cases. Mori (1964) noted a flexor carpi radialis accessorius This muscle is only present as a variation or anomaly.
inserting onto the scaphoid in 2.5% of cases (unclear if this is
synonymous with flexor carpi radialis brevis).
Comparative Anatomy
Anomalies According to Diogo and Wood (2012a), Lewis (1989) states
that in chimpanzees (and humans), if flexor carpi radialis
Description originates from the radius via two tendons, one is referred
Macalister (1875) notes that Koster observed an insertion to as flexor carpi radialis brevis and the other is referred to
of flexor carpi radialis into the distal radius in an individual flexor carpi radialis longus.
with a congenital deformity and that Friedlowsky found an
insertion into the scapho-trapezium in a hand that had only a
Variations
thumb and two fingers. In a trisomy 18 cyclopic fetus, Smith
et al. (2015) observed that on both sides of the body, flexor Description
carpi radialis was fused with pronator teres. In a fetus with Flexor carpi radialis brevis is a variant of flexor carpi radia-
craniorachischisis, Alghamdi et al. (2017) observed that the lis. It is a small muscle that originates from the radius and
left flexor carpi ulnaris had an insertion onto the trapezoid. typically inserts onto the tendon of flexor carpi radialis, or
This muscle was absent on the right side. In a female fetus with the flexor carpi radialis tendon onto the base of the
with trisomy 18, the proximal end of this muscle was fused second metacarpal (Knott 1883a; Nakahashi and Izumi
with that of flexor digitorum superficialis on both sides of 1987; Bergman et al. 1988). This muscle can also insert
the body (Alghamdi et al. 2018). Absence or anomalous onto the retinacular septum, trapezium, capitate, or bases
insertion of flexor carpi radialis has been observed in infants of the third or fourth metacarpals (Knott 1883a; Nakahashi
with Neu-Laxova syndrome (Shved et al. 1985). Bersu and Izumi 1987; Hongsmatip et al. 2019).
et al. (1976) describe a male infant with Hanhart syndrome.
On the right side of this specimen, the wrist and hand were Innervation
deficient. Flexor carpi radialis inserted onto the trapezoid. Flexor carpi radialis brevis is innervated by the anterior
interosseous nerve (Dodds 2006; Hongsmatip et al. 2019).
Prevalence
In their literature review, Smith et al. (2015) found that Prevalence
flexor carpi radialis was absent in 3 out of 26 individuals
Gruber observed flexor carpi radialis brevis in 1 out of 400
with trisomy 18 (11.6%), and that this muscle had vari-
limbs (0.25%), and Wood (1867a) noted its presence in 6 out
able attachments (to the flexor retinaculum, to the base of
of 70 limbs (8.6%). Knott (1883a) found this muscle in 1 out
metacarpal one) in 2 out of 26 individuals with trisomy
of 34 subjects (2.9%).
18 (7.7%). Fusion of pronator teres and flexor carpi radia-
lis was noted in 2 out of 24 individuals with trisomy 13
(8.3%). In a study of six upper limbs from three infants with Anomalies
Neu-Laxova syndrome, Shved et al. (1985) found that flexor Description
carpi radialis was absent in two upper limbs (33%) and an According to Smith et al. (2015), an anomalous flexor carpi
anomalous insertion was present in four upper limbs (66%). radialis brevis can originate from the flexor carpi radialis
belly and insert onto the trapezium.
N/A Prevalence
they found that this muscle was present in 1 out of 26 indi-
Flexor carpi radialis brevis (Figure 3.14) viduals with trisomy 18 (3.8%).
See also: Flexor carpi radialis, Pronator quadratus
Synonyms Hongsmatip et al. (2019) suggest that the interaction
This muscle is also referred to as flexor carpi radialis vel between the tendons of flexor carpi radialis and flexor carpi
profundus (Wood) and the short radiocarpal flexor muscle radialis brevis could lead to tendon intersection syndrome
(Hongsmatip et al. 2019). Though Knott (1883a) and Akita or tenosynovitis.
and Nimura (2016b) list flexor carpus radialis brevis as a
synonym of radiocarpus, we treat these muscles as sepa- Pronator teres (Figure 3.14)
rate structures due to (1) the described tendinous and mus-
cular connections between flexor carpus radialis brevis Synonyms
and flexor carpi radialis and (2) the possibility of radiocar- This muscle is also referred to as pronator radii teres (Knott
pus extending proximally to the ulna or medial epicondyle. 1883a).
Typical Presentation frequencies are similar to the ones provided by Vymazalová

Description et al. (2015), who studied pronator teres in 68 adults and 3
fetuses and noted an origin from both the medial epicon-
The humeral head of pronator teres originates from the
dyle and medial intermuscular septum in 70.6% of cases
medial supracondylar ridge of the humerus, the intermuscu-
and an origin from only the medial epicondyle in 29.4% of
lar septum it shares with flexor carpi radialis, and the ante-
cases. Olewnik et al. (2018a) observed that the ulnar head
brachial fascia (Standring 2016). The ulnar head originates
was present in 43 limbs (86%), therefore yielding an absence
from the coronoid process of the ulna (Standring 2016).
prevalence of 14%. Vymazalová et al. (2015) found that the
The muscle inserts via a tendon onto the lateral radial shaft
ulnar head was absent in 4.4% of cases. Nebot-Cegarra et al.
(Standring 2016).
(1991–1992) observed an absence of the ulnar head of prona-
Innervation tor teres in 22% of cases.
Pronator teres is innervated by the median nerve (Standring
2016). Anomalies
Description
Comparative Anatomy
Macalister (1875) notes that Koster observed an insertion
Pronator teres has a similar typical presentation in the of pronator teres into the distal radius in an individual with
apes, but the ulnar head is frequently absent as a variation a congenital deformity. In a fetus with craniorachischisis,
(Gratiolet and Alix 1866; Chapman 1878; Bischoff 1880; Alghamdi et al. (2017) observed that the right pronator teres
Deniker 1885; Hepburn 1892; Chapman 1900; Loth 1931; only had a humeral head, and some of its fibers had an abnor-
Preuschoft 1965; Jouffroy 1971; Lewis 1989; Gibbs 1999; mal insertion onto the fascia at the level of the hand. On the
Michilsens et al. 2009; Diogo et al. 2010, 2012, 2013a,b, left side, the proximal end of pronator teres was fused with
2017). flexor carpi ulnaris and flexor digitorum superficialis. In a
male infant with triploidy, Moen et al. (1984) found that pro-
Variations
nator teres was absent bilaterally. Absence of the ulnar head
Description and fusion of pronator teres with flexor carpi radialis has
The ulnar head can be reduced or absent (Macalister 1875; been observed in infants with Neu-Laxova syndrome (Shved
Mori 1964; Bergman et al. 1988; Nebot-Cegarra et al. et al. 1985).
1991–1992; Vymazalová et al. 2015; Standring 2016; In a neonate with trisomy 13, pronator teres partially
Olewnik et al. 2018a). The two heads may be partially or originated from flexor carpi radialis (Aziz 1980). On the
completely divided (Macalister 1875; Knott 1883a; Akita right side, it inserted into radiocarpus. In a trisomy 18
and Nimura 2016b). Both heads may also be doubled cyclopic fetus, Smith et al. (2015) observed that on both
(Macalister 1875; Bergman et al. 1988). An accessory slip sides of the body, pronator teres was fused with flexor
may originate from a supracondylar process of the humerus, carpi radialis and there was a second deep belly of prona-
or a ligament can connect pronator teres to the medial epi- tor teres that had an insertion that extended more distally
condyle (Bergman et al. 1988; Standring 2016; Akita and onto the radius.
Nimura 2016b). A third head may arise from the ulna from
below the coronoid process (Macalister 1875; Knott 1883a;
Akita and Nimura 2016b). Slips can extend to the fascia of Prevalence
the arm, the medial intermuscular septum, or the humerus In their literature review, Smith et al. (2015) found that fusion
(Macalister 1875; Bergman et al. 1988; Akita and Nimura of pronator teres and flexor carpi radialis, and pronator teres
2016b; Standring 2016). This muscle can also be connected with two bellies, were both noted in 2 out of 24 individuals
to biceps, brachialis, flexor carpi radialis, flexor pollicis with trisomy 13 (both at a prevalence of 8.3%). Altogether,
longus, or flexor digitorum superficialis (Macalister 1875; pronator teres was variant (accessory slips, doubled superfi-
Mori 1964; Bergman et al. 1988; Akita and Nimura 2016b; cial head, variable origins and insertions) in 4 out of 24 indi-
Standring 2016). viduals with trisomy 13 (16.7%). In 2 out of 26 individuals
with trisomy 18, the ulnar head of pronator teres was missing
Prevalence (7.7%). In a study of six upper limbs from three infants with
In a sample of 80 arms, Mori (1964) found that pronator Neu-Laxova syndrome, Shved et al. (1985) found that both
teres received a slip from brachialis in 15% of cases and a absence of the ulnar head and fusion of pronator teres with
slip from biceps brachii in 17.5% of cases. The ulnar head flexor carpi radialis occurred in four upper limbs (66%).
was poorly developed in 10% of cases. The insertion was
divided into two parts in 4% of cases. In a study of 50 Clinical Implications
limbs, Olewnik et al. (2018a) found that the humeral head As variations of pronator teres may affect the course of the
of pronator teres originated from both the medial epicon- median nerve, variations can lead to median nerve compres-
dyle and medial intermuscular septum in 36 cases (72%), but sion neuropathy or “pronator teres syndrome” (Hartz et al.
only from the medial epicondyle in 14 cases (28%). These 1981; Nebot-Cegarra et al. 1991–1992; Olewnik et al. 2018a).
POSTERIOR FOREARM MUSCLES shaft (Kohlbrügge 1890–1892; Hepburn 1892; Miller

1952; Gibbs 1999; Michilsens et al. 2009; Diogo et al.
brachiOraDialis (figure 3.15) 2010, 2012, 2013a,b, 2017). The origin may extend to
Synonyms the level of the deltoid insertion in gorillas and common
chimpanzees (Sonntag 1923; Swindler and Wood 1973;
Brachioradialis is also referred to as supinator longus, supi-
Diogo et al. 2010). The insertion usually reaches the sty-
nator primus, or grand supinateur (Todd 1839; Barnard
loid process of the radius in gorillas (Diogo et al. 2010).
1875; Parsons 1898b; Jouffroy 1971).
It sometimes reaches the styloid process in gibbons, com-
Typical Presentation mon chimpanzees, bonobos, and orangutans (Hepburn
1892; Primrose 1899, 1900; Ziegler 1964; Swindler and
Description Wood 1973; Michilsens et al. 2009; Diogo et al. 2012,
Brachioradialis originates from the lateral supracondylar 2013a,b, 2017). Brachioradialis may be fused with bra-
ridge of the humerus and lateral intermuscular septum and chialis in gibbons, common chimpanzees, and bonobos
inserts via a tendon onto the distal radius proximal to the (Hepburn 1892; Straus 1941; Miller 1952; Diogo et al.
styloid process (Standring 2016). 2012, 2017).
Innervation
Variations
Brachioradialis is innervated by the radial nerve (Standring
2016). Description
Brachioradialis may insert more proximally on the radius
Comparative Anatomy (Bergman et al. 1988; Akita and Nimura 2016b; Standring
Brachioradialis has a similar typical presentation in the 2016). In other cases, the insertion can extend distally to
apes, extending from the distal humerus to the radial attach onto the scaphoid, trapezium, or base of the third
FIGURE 3.15 Superfcial posterior forearm muscles in posterior view.

metacarpal (Macalister 1875; Bergman et al. 1988; Akita abductor pollicis longus, and then rejoining as one tendon
and Nimura 2016b). Its tendon may be bifurcated or tri- distally. Opponens pollicis had an abnormal attachment
furcated, or the entire muscle may be doubled (Macalister to this tendon near the base of the first metacarpal.
1875; Mori 1964; Bergman et al. 1988; Turkof et al. In a female fetus with trisomy 18, brachioradialis was
1994, 1995; Akita and Nimura 2016b; Standring 2016). thin on both sides of the body (Alghamdi et al. 2018). In a
Brachioradialis can be partially fused with or connected trisomy 18 cyclopic fetus, Smith et al. (2015) observed that
via slips to abductor pollicis longus, biceps brachii, bra- brachialis was fused with extensor carpi radialis longus and
chialis, deltoid, extensor carpi radialis longus, supinator, extensor carpi radialis brevis at their proximal ends on the
or the forearm fascia (Macalister 1875; Inoue 1934; Mori left side of the body. In a study of specimens with triploidy,
1964; Bergman et al. 1988; Akita and Nimura 2016b; Moen et al. (1984) found that brachioradialis was doubled
Standring 2016). In rare cases, brachioradialis can be on the right side of a male fetus and doubled bilaterally in a
entirely absent (Macalister 1875; Bergman et al. 1988; female fetus. Partial fusion of brachioradialis with brachia-
Standring 2016). lis has been observed in infants with Neu-Laxova syndrome
An accessory brachioradialis may also be referred to as (Shved et al. 1985).
supinator radii longus accessorius, brachioradialis brevis, On the left arm of a male neonate with Meckel syndrome,
or brachioradialis minor (Knott 1883a). It originates next to a small slip extended between the tendon of brachioradia-
or from brachioradialis and attaches to the radial tuberosity lis and flexor pollicis longus (Pettersen 1984). Bersu et al.
(Lauth 1830; Gruber 1867b; Knott 1883a; Le Double 1897; (1976) describe a male infant with Hanhart syndrome. On
Bergman et al. 1988; Rodríguez-Niedenführ et al. 2001; the right side of this specimen, the wrist and hand were
Akita and Nimura 2016b). This muscle can also insert onto deficient. Brachioradialis inserted onto the radius proximal
the supinator, pronator teres tendon, or the ulna (Bergman to the styloid process.
et al. 1988; Rodríguez-Niedenführ et al. 2001; Akita and
Nimura 2016b).
Prevalence
Prevalence In their literature review, Smith et al. (2015) found that
brachioradialis was variable (e.g., more distal attachment,
Mori (1964) found that brachioradialis was connected
diminutive muscle) in 3 out of 26 individuals with trisomy
(via muscular slips) with brachialis in 2.5% of cases,
18 (11.5%). In a study of six upper limbs from three infants
abductor pollicis longus in 12.5% of cases, extensor carpi
with Neu-Laxova syndrome, Shved et al. (1985) found that
radialis longus in 5% of cases, and biceps brachii in 2.5%
brachioradialis was partially fused with brachialis in all six
of cases. The insertion tendon of brachioradialis attached
upper limbs (100%).
to the dorsal radiocarpal ligament in 2.5% of cases (Mori
1964). In another 2.5% of cases, the insertion tendon was
doubled, and the superficial branch of the radial nerve Clinical Implications
passed between the slips (Mori 1964). An accessory brachioradialis muscle or a split brachiora-
In a study of 176 upper limbs from 88 cadavers, dialis muscle/tendon can create possible entrapment sites
Rodríguez-Niedenführ et al. (2001) found an accessory for the radial nerve (Turkof et al. 1994, 1995; Spinner and
brachioradialis present in five upper limbs (2.8%) from Spinner 1996; Akita and Nimura 2016b).
three cadavers. Turkof et al. (1994) found that the super-
ficial branch of the radial nerve emerged between a split
brachioradialis tendon in 5 out of 150 arms (3.3%) from
4 out of 75 cadavers. Turkof et al. (1995) found that in 143 Supinator (Figure 3.16)
patients with Wartenberg’s syndrome (radial sensory nerve Synonyms
entrapment at the forearm), seven of them (4.9%) exhibited
This muscle is also referred to as supinator brevis (Parsons
a superficial branch of the radial nerve passing between a
1898b) or epicondylo-radial (Todd 1839).
split brachioradialis tendon.

Windle (1893) observed a slip connecting brachioradialis Supinator has two heads/two layers of fibers (Standring
and supinator on the right side of one male fetus with 2016). The humeral (superficial) head and ulnar (deep)
anencephaly. On the left side of the body of a fetus with head originate together from the lateral epicondyle of the
craniorachischisis, Alghamdi et al. (2017) observed that humerus, radial collateral ligament, annular ligament of
brachioradialis had a normal origin but it had two ten- the superior radioulnar joint, and from the supinator crest
dons. The first attached to the distal end of the radius. on the ulna (Standring 2016; Akita and Nimura 2016b).
The other attached to the base of the first metacarpal, Supinator inserts onto the proximal third of the radius
exhibiting bifurcation as it surrounded the tendon of (Standring 2016).
FIGURE 3.16 Deep posterior forearm muscles in posterior view.
Innervation parts of supinator have been named as variations, includ-

Supinator is innervated by the posterior interosseous nerve ing the lateral and medial tensors of the annular ligament
(Standring 2016). (Macalister 1875; Standring 2016). Supinator may be
doubled or divided (Macalister 1875). It may send acces-
Comparative Anatomy sory bundles to the biceps tendon, the bursa under the ten-
Supinator has a similar typical presentation in the apes, don, the radial tuberosity, or pronator teres (Clason 1869;
extending from the lateral epicondyle of the humerus and Macalister 1875; Bergman et al. 1988; Akita and Nimura
from the proximal ulna to the proximal radius (Gibbs 1999; 2016b).
Diogo et al. 2010, 2012, 2013a,b, 2017). The humeral origin
may be absent in some orangutans and bonobos (Barnard Prevalence
1875; Straus 1941; Kallner 1956; Diogo et al. 2013b, 2017). Mori (1964) found that the distal end of the supinator inser-
The ulnar origin may be absent in gibbons (Michilsens tion was distal to the middle point of the radius in 16% of
et al. 2009; Diogo et al. 2012). cases.
Variations Anomalies
The degree of division into two layers can vary (Bergman Windle (1893) observed a slip connecting brachioradialis
et al. 1988; Akita and Nimura 2016b). Its insertion may and supinator on the right side of one male fetus with anen-
shift more distally along the radius (Mori 1964). Small cephaly. On the left side of a fetus with craniorachischisis
(Alghamdi et al. 2017), supinator was fused with extensor Anomalies

carpi ulnaris. Description
Anconeus had a normal presentation and innervation in a
Prevalence fetus with craniorachischisis dissected by Alghamdi et al.
N/A (2017).
Clinical Implications Prevalence

If the most proximal portion of the humeral (superfcial) N/A
head of supinator is tendinous and forms a fbrous arch
(arcade of Frohse), this can cause paralysis of the posterior Clinical Implications
interosseous nerve (Spinner 1968).
N/A
ancOneus (figure 3.15) extensOr carpi raDialis lOngus (figure 3.15)

Synonyms See also: Extensor carpi radialis brevis, Extensor carpi
Alternative names for anconeus include extensor ante- radialis accessorius
brachii ulnaris, epicondylo-cubitalis, anconeus brevis,
anconeus parvus or anconeus quartus (Jouffroy 1971). Synonyms
This muscle is also referred to radialis externus longior or
Typical Presentation humero sus-metacarpien (Todd 1839).
Description
Anconeus originates via a tendon from the lateral epicon- Typical Presentation
dyle of the humerus and inserts onto the lateral surface of Description
the olecranon and the posterior surface of the proximal Extensor carpi radialis longus originates from the lateral
ulnar shaft (Standring 2016). supracondylar ridge of the humerus and the lateral inter-
muscular septum and inserts via a tendon onto the base of
Innervation the second metacarpal (Standring 2016).
Anconeus is innervated by the radial nerve (Standring
2016). Innervation
Extensor carpi radialis longus is innervated by the radial
Comparative Anatomy nerve (Standring 2016).
Anconeus has a similar typical presentation in the great
apes, extending from the lateral epicondyle to the olecra- Comparative Anatomy
non process and adjacent areas of the ulna (Miller 1952; Extensor carpi radialis longus has a similar typical presen-
Gibbs 1999; Diogo et al. 2010, 2013a,b, 2017). In gibbons, tation in the apes, sometimes with an additional attachment
it is undifferentiated from extensor carpi ulnaris and does to the frst metacarpal (Kohlbrügge 1890–1892; Hepburn
not appear as a distinct muscle (Kohlbrügge 1890–1892; 1892; Straus 1941; Bojsen-Møller 1978; Gibbs 1999;
Hepburn 1892; Gibbs 1999; Michilsens et al. 2009; Diogo Michilsens et al. 2009; Diogo et al. 2010, 2012, 2013a,b,
et al. 2012). 2017). It may also attach to the third metacarpal in com-
mon chimpanzees and bonobos (Diogo et al. 2013a, 2017).
Variations It may be blended with extensor carpi radialis brevis in
Description common chimpanzees, gibbons, and orangutans (Deniker
1885; Beddard 1893; Sonntag 1923; Gibbs 1999; Diogo
Anconeus can have connections to, or be partially fused
et al. 2012, 2013b).
with, epitrochleoanconeus, extensor carpi ulnaris, or tri-
ceps brachii (Macalister 1875; Bergman et al. 1988; Akita
Variations
and Nimura 2016a; Standring 2016). This muscle may be
divided into two portions (Macalister 1875). It can also be Description
entirely absent (Bergman et al. 1988). Anconeus likely has Extensor carpi radialis longus shares a developmental ori-
developmental origins from the triceps brachii due to its gin with extensor carpi radialis brevis so these muscles may
position, propensity to be blended with this muscle, and the be completely or partially fused (Macalister 1875; Knott
shared innervation between anconeus and the medial head 1883a; Mori 1964; Albright and Linburg 1978; Bergman
of triceps (Bergman et al. 1988; Akita and Nimura 2016a). et al. 1988; Akita and Nimura 2016b; Standring 2016). The
muscle belly or tendon of extensor carpi radialis longus can
Prevalence split into multiple slips prior to insertion (Macalister 1875;
N/A Knott 1883a; Mori 1964; Bergman et al. 1988; Tountas
and Bergman 1993; Akita and Nimura 2016b; Standring 1978). A tendon originating from extensor carpi radia-
2016). This muscle may send a slip to the frst metacar- lis brevis and inserting onto the second metacarpal next
pal, third metacarpal, fourth metacarpal, or the trapezium to extensor carpi radialis longus was present in 37 limbs
(Macalister 1875; Knott 1883a; Albright and Linburg 1978; (21.4%). Extensor carpi radialis intermedius was present
Bojsen-Møller 1978; Bergman et al. 1988; Lewis 1989; in 42 limbs (24.3%).
Akita and Nimura 2016b; Standring 2016). Its tendon may Mori (1964) describes several variations in extensor
send a slip to brachioradialis (Macalister 1875). It may also carpi radialis longus anatomy. In 2.5% of cases, the inser-
have connections to abductor pollicis longus or to the inter- tion tendon of this muscle was fused with that of extensor
osseous muscles (Macalister 1875; Bergman et al. 1988; carpi radialis brevis, and this anatomy was associated with
Akita and Nimura 2016b). Extensor carpi radialis longus a two-headed fexor carpi radialis longus that was fused
may be trigastric and insert onto both the frst and sec- with fexor carpi radialis brevis. In another 2.5% of cases,
ond metacarpals (Bergman et al. 1988; Akita and Nimura the extensor carpi radialis longus arose from two heads
2016b). Yang et al. (2018) report a similar case in which the that both fused with extensor carpi radialis brevis to some
lateral and intermediate heads of a trigastric extensor carpi extent. The insertion tendon of extensor carpi radialis bre-
radialis longus muscle fused and inserted onto the second vis was split into two, and the smaller one fused with the
metacarpal while the medial head merged with extensor insertion tendon of extensor carpi radialis longus (Mori
carpi radialis brevis. 1964). In another 2.5% of cases, extensor carpi radialis
The radial extensor muscles are associated with two longus and extensor carpi radialis brevis were each divided
(potentially three) supernumerary muscles. Extensor carpi into two bellies that each had their own insertion tendons,
radialis intermedius originates from the lateral epicon- thus presenting as a group of four muscles that fused proxi-
dyle between extensor carpi radialis longus and brevis, or mally to form extensor carpi radialis intermedius (Mori
directly from one of these muscles, and inserts onto the sec- 1964). Lastly, 5.4% of the radial extensors showed overall
ond and/or third metacarpals (Albright and Linburg 1978; variation in the separation of the tendon of insertion (Mori
Bergman et al. 1988; Tountas and Bergman 1993; Akita 1964).
and Nimura 2016b; West et al. 2017). Nayak et al. (2007)
and Bharambe et al. (2017) referred to an additional radial Anomalies
extensor they term extensor carpi radialis tertius. Nayak
Description
et al. (2007) suggest that extensor carpi radialis tertius orig-
inates from the lateral epicondyle between extensor carpi Extensor carpi radialis longus had a duplicate tendon on
radialis longus and extensor digitorum and attaches to the the right side of one anencephalic male studied by Windle
second and third metacarpals via two tendons. Bharambe (1893). Barash et al. (1970) found that extensor carpi radia-
et al. (2017) use extensor carpi radialis tertius to refer to lis longus and extensor carpi radialis brevis were fused in
a muscle that originates from the posterolateral surface of a trisomy 18 infant. In a female neonate with trisomy 18
the radius deep to extensor carpi radialis brevis and inserts (Aziz 1979) and a female fetus with trisomy 18 (Alghamdi
onto the third metacarpal. Extensor carpi radialis tertius et al. 2018), the proximal ends of extensor carpi radialis
is likely synonymous with extensor carpi radialis interme- longus and brevis were fused bilaterally. In a fetus with tri-
dius. Extensor carpi radialis accessorius is the most radial somy 18 and cyclopia, extensor carpi radialis longus was
of these supernumerary muscles, and its insertion is more fused with extensor carpi radialis brevis and brachioradia-
closely associated with the thenar muscles (see the entry for lis on the left side of the body (Smith et al. 2015), and both
this muscle). radial extensors were fused to each other on the right side.
Extensor carpi radialis longus has been observed to share
Prevalence a common muscle belly with extensor carpi radialis brevis
Knott (1883a) found extensor carpi radialis longus fused in infants with Neu-Laxova syndrome (Shved et al. 1985).
with extensor carpi radialis brevis in 5 out of 34 sub- Bersu et al. (1976) describe a male infant with Hanhart syn-
jects (14.7%). Bojsen-Møller (1978) found that extensor drome. On the right side of this specimen, the wrist and
carpi radialis longus inserted onto both the frst and sec- hand were defcient. Extensor carpi radialis longus inserted
ond metacarpals in 2 out of 23 individuals (8.7%). In a onto the trapezium.
study of 173 upper limbs, Albright and Linburg (1978)
found that extensor carpi radialis longus and extensor Prevalence
carpi radialis brevis exhibited typical presentations (the In their literature review, Smith et al. (2015) found that the
former going to metacarpal two and the latter going to radial extensor muscles were fused in 1 out of 24 individu-
metacarpal three with no interconnections) in eight-six als with trisomy 13 (4.2%) and in 4 out of 26 individuals
limbs (49.7%). Interconnections between the two muscles with trisomy 18 (15.4%). Smith et al. (2015) also found that
were observed in 61 limbs (35.3%). A tendon originating extensor carpi radialis (both muscles) were variable (e.g.,
proximally from extensor carpi radialis longus inserted fusion of the two, variable origins and insertions, doubled
on the third metacarpal in front of the extensor carpi tendon or muscle bellies) in 3 out of 24 individuals with
radialis brevis in 45 limbs (26%) (Albright and Linburg trisomy 13 (12.5%) and in 10 out of 26 individuals with
trisomy 18 (38.5%). In a study of six upper limbs from three et al. 1988; Tountas and Bergman 1993; Akita and Nimura
infants with Neu-Laxova syndrome, Shved et al. (1985) 2016b; Standring 2016). Slips may pass from this muscle
found that extensor carpi radialis longus and extensor carpi to the second metacarpal, fourth metacarpal, trapezium, or
radialis brevis shared a common muscular belly in all six dorsal interossei (Macalister 1875; Knott 1883a; Albright
upper limbs (100%). and Linburg 1978; Bergman et al. 1988; Lewis 1989; Akita
and Nimura 2016b; Standring 2016). Reina-de la Torre et al.
Clinical Implications (1994) report a case in which extensor carpi radialis brevis
The presence of extra tendons of extensor carpi radialis lon- was attached via a tendinous accessory slip to the insertion
gus should be checked when planning for tendon transfer tendon of biceps brachii. Mitsuyasu et al. (2004) report a
surgery (Albright and Linburg 1978). case in which extensor carpi radialis brevis had a normal
insertion onto the base of the third metacarpal but origi-
nated from the fascia/tendon of extensor digitorum instead
extensOr carpi raDialis brevis (figure 3.15)
of the lateral epicondyle. Extensor carpi radialis brevis
See also: Extensor carpi radialis longus, Extensor carpi may be absent (Macalister 1875). It may also be doubled
radialis accessorius (Macalister 1875).
The radial extensor muscles are associated with two
Synonyms (potentially three) supernumerary muscles. See the entry
This muscle is also referred to as radialis externus brevior for extensor carpi radialis longus for details on extensor
or epicondylo sus-metacarpien (Todd 1839). carpi radialis intermedius and extensor carpi radialis ter-
tius. Extensor carpi radialis accessorius is the most radial
Typical Presentation of these supernumerary muscles, and its insertion is more
Description closely associated with the thenar muscles (see the entry for
Extensor carpi radialis brevis originates from the lateral this muscle).
epicondyle of the humerus and inserts via a tendon onto the Prevalence
base of the third metacarpal (Standring 2016).
According to Macalister (1875), Wood observed extensor
Innervation carpi radialis brevis sending tendons into both the second
Extensor carpi radialis brevis is innervated by the radial and third metacarpals in 4 out of 36 subjects (11%). Knott
nerve or the posterior interosseous nerve (Standring 2016). (1883a) found extensor carpi radialis longus fused with
extensor carpi radialis brevis in 5 out of 34 subjects (14.7%).
Comparative Anatomy In a study of 173 upper limbs, Albright and Linburg (1978)
found that extensor carpi radialis longus and extensor carpi
Extensor carpi radialis brevis has a similar typical pre-
radialis brevis exhibited typical presentations (the former
sentation in the apes, with an origin from the lateral epi-
going to metacarpal two and the latter going to metacarpal
condyle and frequent insertion onto both the second and
three with no interconnections) in eight-six limbs (49.7%).
third metacarpals (Kohlbrügge 1890–1892; Straus 1941;
Interconnections between the two muscles were observed
Raven 1950; Gibbs 1999; Michilsens et al. 2009; Diogo
in 61 limbs (35.3%). A tendon originating proximally from
et al. 2010, 2013, 2013a,b, 2017). There may be an origin
extensor carpi radialis longus inserted on the third metacar-
from extensor digitorum in gibbons (Michilsens et al. 2009;
pal in front of the extensor carpi radialis brevis in 45 limbs
Diogo et al. 2012). There may be an origin from the lateral
(26%) (Albright and Linburg 1978). A tendon originating
condyle of the humerus in gibbons and gorillas (Deniker
from extensor carpi radialis brevis and inserting onto the
1885; Kohlbrügge 1890–1892; Diogo et al. 2010; 2012). It
second metacarpal next to extensor carpi radialis longus
may be blended with extensor carpi radialis longus in com-
was present in 37 limbs (21.4%). Extensor carpi radialis
mon chimpanzees, gibbons, and orangutans (Deniker 1885;
intermedius was present in 42 limbs (24.3%).
Beddard 1893; Sonntag 1923; Gibbs 1999; Diogo et al.
Mori (1964) describes several variations in extensor
2012, 2013b).
carpi radialis brevis anatomy. In 2.5% of cases, the inser-
tion tendon of this muscle was fused with that of extensor
Variations carpi radialis longus, and this anatomy was associated with
Description a two-headed fexor carpi radialis longus that was fused
Extensor carpi radialis brevis shares a developmental ori- with fexor carpi radialis brevis. In another 2.5% of cases,
gin with extensor carpi radialis longus so sometimes these the extensor carpi radialis longus arose from two heads
muscles can be completely or partially fused (Macalister that both fused with extensor carpi radialis brevis to some
1875; Knott 1883a; Mori 1964; Albright and Linburg 1978; extent. The insertion tendon of extensor carpi radialis bre-
Bergman et al. 1988; Akita and Nimura 2016b; Standring vis was split into two, and the smaller one fused with the
2016). The muscle belly or tendon of extensor carpi insertion tendon of extensor carpi radialis longus (Mori
radialis brevis can split into multiple slips prior to inser- 1964). In another 2.5% of cases, extensor carpi radialis
tion (Macalister 1875; Knott 1883a; Mori 1964; Bergman longus and extensor carpi radialis brevis were each divided
into two bellies that each had their own insertion tendons, Synonyms
thus presenting as a group of four muscles that fused proxi- N/A
mally to form extensor carpi radialis intermedius (Mori
1964). Lastly, 5.4% of the radial extensors showed overall Typical Presentation
variation in the separation of the tendon of insertion (Mori This muscle is only present as a variation or anomaly.
1964).
Comparative Anatomy
Anomalies N/A
Description
Barash et al. (1970) found that extensor carpi radialis lon- Variations
gus and extensor carpi radialis brevis were fused in a tri-
Description
somy 18 infant. In a female neonate with trisomy 18 (Aziz
1979) and a female fetus with trisomy 18 (Alghamdi et al. Extensor carpi radialis accessorius (Wood) originates
2018), the proximal ends of extensor carpi radialis longus from the humerus with, or from, extensor carpi radialis
and brevis were fused bilaterally. In a trisomy 18 cyclopic longus and inserts onto the first metacarpal (Wood 1867b;
fetus, extensor carpi radialis brevis was fused with exten- Macalister 1867b, 1875; Knott 1883a; Bergman et al. 1988;
sor carpi radialis longus and brachioradialis on the left Hong and Hong 2005; Akita and Nimura 2016b; West et al.
side of the body (Smith et al. 2015), and both radial exten- 2017). Macalister (1875) notes that other anatomists, includ-
sors were fused to each other on the right side. Extensor ing Wood and Cruveilhier, observed that this muscle arose
carpi radialis brevis has been observed to share a com- from the fascia over the other radial extensors. West et al.
mon muscle belly with extensor carpi radialis longus in (2017) note an origin in one case from extensor carpi radia-
infants with Neu-Laxova syndrome (Shved et al. 1985). lis brevis.
Bersu et al. (1976) describe a male infant with Hanhart Extensor carpi radialis accessorius may insert onto, or
syndrome. On the right side of this specimen, the wrist give origin to, abductor pollicis brevis (Macalister 1875;
and hand were deficient. Extensor carpi radialis brevis Knott 1883a; Frohse and Frankel 1908; Khaledpour and
inserted onto the trapezoid. Schindelmeiser 1994; Hong and Hong 2005). It also may
be continuous with the origin of flexor pollicis brevis
Prevalence (Macalister 1875; Knott 1883a). Knott (1883a) also observed
In their literature review, Smith et al. (2015) found that the an insertion into the scaphoid. This muscle can sometimes
radial extensor muscles were fused in 1 out of 24 individu- be represented by a slip that arises from the extensor carpi
als with trisomy 13 (4.2%) and in 4 out of 26 individuals radialis longus tendon and inserts onto the first metacar-
with trisomy 18 (15.4%). Smith et al. (2015) also found that pal and the first dorsal interosseous muscle (Wood 1867b;
extensor carpi radialis (both muscles) were variable (e.g., Macalister 1875; Bergman et al. 1988).
fusion of the two, variable origins and insertions, doubled Hong and Hong (2005) suggest that extensor carpi
tendon or muscle bellies) in 3 out of 24 individuals with radialis intermedius is an accessory head of extensor carpi
trisomy 13 (12.5%) and in 10 out of 26 individuals with tri- radialis brevis, while extensor carpi radialis accessorius is
somy 18 (38.5%). Extensor carpi radialis brevis was absent an accessory head of extensor carpi radialis longus. These
in 2 out of 26 individuals with trisomy 18 (7.7%) (Smith authors also note that the tendon of extensor carpi radialis
et al. 2015). In a study of six upper limbs from three infants accessorius can pass through its own dorsal tunnel under
with Neu-Laxova syndrome, Shved et al. (1985) found that the extensor retinaculum as it enters the hand (Hong and
extensor carpi radialis longus and extensor carpi radialis Hong 2005), a feature that has been noted by others (Frohse
brevis shared a common muscular belly in all six upper and Frankel 1908; Claassen and Wree 2002; West et al.
limbs (100%). 2017).
Clinical Implications Innervation

The presence of extra tendons of extensor carpi radialis Extensor carpi radialis accessorius is innervated by the deep
brevis should be checked when planning for tendon trans- branch of the radial nerve (Khaledpour and Schindelmeiser
fer surgery (Albright and Linburg 1978). Mitsuyasu et al. 1994).
(2004) suggest that a variant origin of the extensor carpi
radialis brevis may affect the severity of, or ability to Prevalence
develop, lateral epicondylitis (tennis elbow). In a study of 82 forearms from 41 cadavers, West et al.
(2017) found that additional wrist extensors were present in
seven forearms (8.5%) from five individuals, with a preva-
Extensor carpi radialis accessorius (Figure 3.15) lence of four extensor carpi radialis intermedius muscles
See also: Extensor carpi radialis brevis, Extensor carpi (4.9%) and three extensor carpi radialis accessorius muscles
radialis longus (3.6%).
Anomalies minimi may originate from the tendon of extensor carpi

Description ulnaris (Macalister 1875). Extensor carpi ulnaris may be
doubled (Bergman et al. 1988; Akita and Nimura 2016b).
Smith et al. (2015) describe extensor carpi radialis acces-
The tendon of extensor carpi ulnaris may split distally close
sorius as superfcial to extensor carpi radialis.
to insertion (Erickson et al. 2019). Extensor carpi ulnaris is
Prevalence associated with an accessory slip, ulnaris digiti quinti (see
Smith et al. (2015) note that this muscle was present in 5 out the entry for this muscle).
of 26 individuals with trisomy 18 (19.2%). Prevalence
Clinical Implications In a study of 17 extensor carpi ulnaris tendons, Erickson
Extensor carpi radialis accessorius may contribute to lat- et al. (2019) found that 11 specimens demonstrated at least
eral epicondylitis (tennis elbow) or nerve entrapment (West one split in the distal tendon (64.7%), with seven specimens
et al. 2017). having one split, one specimen having two splits, and three
specimens having three splits.
extensOr carpi ulnaris (figure 3.15) Anomalies

See also: Ulnaris digiti quinti Description
In a fetus with trisomy 18 and cyclopia, the proximal end
Synonyms of extensor carpi ulnaris was fused with extensor digiti
This muscle is also referred to as ulnaris externus or cubito minimi and extensor digitorum on the left side (Smith
sus-metacarpien (Todd 1839). et al. 2015). Bersu et al. (1976) describe a male infant with
Hanhart syndrome. On the right side of this specimen, the
Typical Presentation wrist and hand were defcient. Extensor carpi ulnaris had a
Description normal insertion onto the base of the ffth metacarpal.
Extensor carpi ulnaris originates from the lateral epicon-
Prevalence
dyle of the humerus and the posterior margin of the ulna
(Standring 2016). It inserts via a tendon onto the base of the In their literature review, Smith et al. (2015) found that
ffth metacarpal (Standring 2016). extensor carpi ulnaris had a variable insertion (onto the
medial aspect of the ffth metacarpal distal to its base) in 1
Innervation out of 24 individuals with trisomy 13 (4.2%).
Extensor carpi ulnaris is innervated by the posterior inter-
osseous nerve (Standring 2016).
Erickson et al. (2019) note that distal splitting of the exten-
Comparative Anatomy sor carpi ulnaris tendon may mimic tears or tendinopathy
Extensor carpi ulnaris has a similar typical presenta- in MRI and ultrasound imaging of the wrist.
tion in the apes, extending from the lateral epicondyle
and the ulna to the ffth metacarpal (Gratiolet and Alix ulnaris Digiti quinti (figure 3.15)
1866; Champneys 1872; Kohlbrügge 1890–1892; Beddard
1893; Primrose 1899, 1900; Sonntag 1923, 1924; Sullivan See also: Extensor carpi ulnaris
and Osgood 1927; Straus 1941; Raven 1950; Miller 1952;
Synonyms
Swindler and Wood 1973; Gibbs 1999; Michilsens et al.
2009; Diogo et al. 2010, 2012, 2013a,b, 2017). The ulnar This muscle is also referred to as ulnaris digiti minimi
origin may be absent in orangutans (Kallner 1956; Diogo (Bergman et al. 1988) or ulnaris quinti (Macalister 1875).
et al. 2013b). The humeral origin may be absent in bonobos
(Diogo et al. 2017). Typical Presentation
Variations
The insertion tendon of extensor carpi ulnaris can also Macalister (1871) observed the presence of an ulnaris digiti
attach to the bases of metacarpal three or four, or the quinti tendon in a chimpanzee that inserted on the proximal
proximal phalanx of the ffth digit (Bergman et al. 1988; phalanx of digit fve (Diogo et al. 2013a).
Akita and Nimura 2016b). A second tendon of exten-
sor carpi ulnaris can replace the tendon of extensor digiti Variations
minimi (Macalister 1875; Bergman et al. 1988; Akita and Description
Nimura 2016b). A slip may connect extensor carpi ulna- Ulnaris digiti quinti originates from the dorsal surface of
ris to triceps brachii (Macalister 1875). Abductor digiti the ulna and inserts onto the base of the proximal phalanx
of the ffth digit (Bergman et al. 1988; Akita and Nimura Prevalence
2016b). It can be represented by a bundle from the belly In their literature review, Smith et al. (2015) noted that
of extensor carpi ulnaris, or more often by a slip from the ulnaris digiti quinti was only present in 1 out of 20 indi-
tendon of extensor carpi ulnaris, which can insert onto viduals with trisomy 13 (5%).
the metacarpal, proximal phalanx, or extensor tendon of
the ffth digit (Barfred and Adamsen 1986; Bergman et al. Clinical Implications
1988). In many cases, a slip can extend from the extensor The presence of ulnaris digiti quinti may lead to the impair-
carpi ulnaris tendon and insert into the fascia covering the ment of the wrist joint and ffth digit (Barfred and Adamsen
metacarpal, the capsule of the metacarpophalangeal joint, 1986; Pınar et al. 2012).
or the proximal phalanx of the ffth digit (Macalister 1875;
Barfred and Adamsen 1986; Bergman et al. 1988). This slip
can also be joined or replaced by a bundle originating from extensOr DigitOruM (figure 3.17)
the pisiform (Bergman et al. 1988). Barfred and Adamsen Synonyms
(1986) observe the presence of this muscle, described as a This muscle is also referred to as extensor digitorum com-
duplication of the extensor carpi ulnaris tendon, in three munis (Knott 1883a).
patients. Pınar et al. (2012) describe in three arms the pres-
ence of tendinous accessory slips that originate from the Typical Presentation
head of extensor carpi ulnaris and insert onto the extensor
Description
apparatus of the ffth digit.
Extensor digitorum originates from the lateral epicondyle
Innervation of the humerus and the neighboring intermuscular septum
(Standring 2016). It typically inserts via four tendons onto
N/A
the extensor expansions, and thus to the middle and dis-
Prevalence tal phalanges, of digits two through fve (Standring 2016).
Intertendinous connections between the terminal tendons
Macalister (1875) notes that Wood found “ulnaris quinti” in
are referred to as juncturae tendinae (Standring 2016; Akita
12% of subjects. Pınar et al. (2012) found three accessory
and Nimura 2016b).
slips that likely correspond to ulnaris digiti quinti in 3 out
of 54 arms (5.6%). Innervation
Extensor digitorum is innervated by the posterior interosse-
Anomalies ous nerve (Standring 2016).
Description
Aziz (1980) observed the bilateral presence of ulnaris digiti Comparative Anatomy
quinti in a neonate with trisomy 13. On the right side, it Extensor digitorum has a similar typical presentation in the
arose as a tendinous slip from extensor carpi ulnaris and apes with similar variations in tendon absence, duplication,
inserted onto the proximal phalanx of the ffth digit. On the and insertions (Owen 1868; Macalister 1873; Chapman
left side, it consisted of two slips. 1878; Kohlbrügge 1890–1892; Hepburn 1892; Sommer
FIGURE 3.17 Extensor muscles on the dorsal (posterior) aspect of the hand and wrist. Extensor medii digiti is illustrated on the left
side, and the other fnger extensors are illustrated on the right side.
1907; Sonntag 1923, 1924; Straus 1941; Raven 1950; Miller hands, tripled in 15% of left hands, and quadrupled in 4% of
1952; Preuschoft 1965; Kaneff 1979; Gibbs 1999; Diogo both right and left hands. The tendon to the fourth digit was
et al. 2010, 2012, 2013a,b, 2017). Additional origins from single in 17% of right hands and 19% of left hands, doubled
the radius and/or ulna have been observed in all apes except in 41% of right hands and 62% of left hands, tripled in 25%
bonobos (Deniker 1885; MacDowell 1910; Sonntag 1924; of right hands and 19% of left hands, and quadrupled in
Straus 1941; Michilsens et al. 2009; Diogo et al. 2010, 2012, 17% of right hands. The tendon to the ffth digit was absent
2013a,b). in 71% of right hands and 50% of left hands, single in 21%
of right hands and 42% of left hands, doubled in 4% of right
Variations hands and 8% of left hands, and tripled in 4% of right hands
(Zilber and Oberlin 2004).
Description
Zilber and Oberlin (2004) also provide a comprehensive
The extensor digitorum tendon to any digit may be dou- literature review of extensor tendon variation, summariz-
bled, tripled, or quadrupled (Wood 1868; Macalister 1875; ing results from a study of 150 hands by Mestdagh et al.
Knott 1883a; Mori 1964; Mestdagh et al. 1985; Godwin and (1985), a study of 50 hands by Godwin and Ellis (1992),
Ellis 1992; von Schroeder and Botte 1995; el-Badawi et al. a study of 43 hands by von Schroeder and Botte (1995), a
1995; Hirai et al. 2001; Zilber and Oberlin 2004; Akita study of 181 hands by el-Badawi et al. (1995), and a study of
and Nimura 2016b; Standring 2016; Bharambe et al. 2017). 548 hands by Hirai et al. (2001). Among these studies, the
Occasionally, the tendon to digit fve is missing, or there may prevalence ranges are as follows. The extensor digitorum
also be a tendon that acts on the frst digit (Macalister 1875; communis tendon for digit two was single in 92%–100%
Knott 1883a; Sato 1969; Kaneff 1979; Bergman et al. 1988; of hands and doubled in 2%–8% of hands. The extensor
Lewis 1989; Zilber and Oberlin 2004; Standring 2016). The digitorum communis tendon for digit three was single in
muscle belly itself can be split into up to four bellies, one 51%–92% of hands, doubled in 4%–39% of hands, tripled
for each target digit (Macalister 1875; Mori 1964; Bergman in 4%–19% of hands, and quadrupled in 5% of hands. The
et al. 1988; Akita and Nimura 2016b). Extensor digitorum extensor digitorum communis tendon for digit four was sin-
may be connected to extensor indicis, extensor digiti min- gle in 12%–96% of hands, doubled in 2%–63% of hands,
imi, or extensor pollicis longus (Macalister 1875; Bergman tripled in 1%–16% of hands, and quadrupled in 4%–9% of
et al. 1988; Akita and Nimura 2016b; Standring 2016). hands. The extensor digitorum communis tendon for digit
fve was absent in 1%–54% of hands, single in 2%–30% of
Prevalence hands, doubled in 10%–25% of hands, tripled in 1%–2% of
Mori (1964) found that the extensor muscle belly of exten- hands, and a common tendon for the fourth and ffth digit
sor digitorum had three separations in 90% of cases and was present in 41%–96% of hands.
four separations in 10% of cases. Doubling of the tendons In a study of 41 hands, Tanaka et al. (2007) found that
was reported as follows: tendon to digit three in 2.5% of the extensor digitorum tendon to digit fve was absent in
cases, tendon to digit four in 5% of cases, tendon to digit ten specimens (24.4%) and present as a single slip in 25
fve in 12.5% of cases, tendons to digits two and four in specimens (61%). The tendon going to the ffth digit was an
2.5% of cases, tendons to digits four and fve in 2.5% of independent tendon in 17 specimens (41.5%) and present as
cases, tendons to digits three, four, and fve in 15% of cases, a shared slip with the fourth digit in 14 specimens (34.1%).
and tendons to digits three and four in 2.5% of cases (Mori From dissections of 100 upper limbs, Suwannakhan
1964). The juncturae tendinae were absent between the ten- et al. (2016) found that the extensor digitorum tendon to
dons to digits two and three in 10% of cases, the tendons digit two was a single, invariable tendon in all cases (100%).
to digits three and four in 10% of cases, and the tendons The tendon to digit three was doubled at its origin in four
to digits four and fve in 2.5% of cases (Mori 1964). The hands (4%) and single at its origin in 96 hands (96%). In 42
tendon to digit fve was absent in 4% of cases. (42%) hands, the tendon to digit three split into two tendons
Sato (1969) found the tendon to digit fve missing in 69 distally, while the tendon split into three in six hands (6%)
out of 366 limbs (18.85%) from Kyushu-Japanese males and and into four tendons in two hands (2%) However, in 99
missing in 38 out of 242 limbs (15.7%) from females. A ten- hands (99%), the tendon to digit three inserted as a single
don to digit one was present in 99 out of 354 limbs (27.97%) tendon while in one hand (1%) it inserted as a double tendon
from males and present in 63 out of 232 limbs from females (Suwannakhan et al. 2016). The tendon to digit four origi-
(27.16%). Kaneff (1979) found that extensor digitorum sent nated as a single tendon in 94 hands (94%), a double tendon
tendons to digits two, three, and four in all 200 upper limbs in fve hands (5%), and a triple tendon in one hand (1%). The
examined by this author (100%), while the tendon to digit tendon to digit four inserted as a single tendon in 64 hands
fve was present in about 94% of cases. (64%), a double tendon in 30 hands (30%), a triple tendon in
In a study of 50 hands (26 left, 24 right), Zilber and fve hands (5%), and a quadruple tendon in one hand (1%).
Oberlin (2004) found that extensor digitorum sent a single The tendon to digit fve was absent in 50 hands (50%). In 49
tendon to the second digit in 100% of cases. The tendon to hands (49%), the tendon to digit fve originated as a single
the third digit was single in 67% of right hands and 62% of tendon and in one hand (1%) it originated as a double tendon
left hands, doubled in 29% of right hands and 19% of left (Suwannakhan et al. 2016).
In a study of 110 upper limbs, Bharambe et al. (2017) Clinical Implications

found that the tendon to digit fve was absent in 28% of Understanding variation in extensor digitorum tendons
cases. In 2% of cases, both the tendons to the third and is important for planning and performing tendon transfer
fourth digit were tripled. Duplication of the tendon to digit surgery (Zilber and Oberlin 2004).
two was seen in 1% of cases, the tendon to digit three in
1% of cases, the tendon to digit four in 12% of cases, and
the tendon to digit fve in 2% of cases. Triplication of the extensOr Digiti MiniMi (figure 3.17)
tendon to the third digit was seen in 1% of cases and of the Synonyms
tendon to the fourth digit in 5% of cases.
This muscle is also referred to as extensor digiti minimi
proprius, extensor digiti minimi manus (Bergman et al.
Anomalies 1988), or extensor digiti quinti proprius (Aziz 1979).
Description
On the right upper limb of a fetus with craniorachischisis,
Alghamdi et al. (2017) noted that a muscle that seemed to Description
correspond to extensor digitorum originated from the lat- Extensor digiti minimi originates from the common exten-
eral epicondyle and had three tendons, one that inserted on sor tendon and neighboring intermuscular septa (Standring
the ulnar styloid process, one that inserted onto digit three, 2016). It inserts via a bifurcated tendon that often joins
and one that inserted onto digit fve. On the left side, the with the extensor digitorum tendon to insert onto the dorsal
muscle had a more typical presentation, but the fourth ten- digital expansion of digit fve (Standring 2016).
don attached to both digits four and fve and the frst tendon
was joined with that of extensor indicis (Alghamdi et al. Innervation
2017). Extensor digiti minimi is innervated by the posterior
In a trisomy 18 cyclopic fetus dissected by Smith et al. interosseous nerve (Standring 2016).
(2015) extensor digitorum did not send a tendon to digit fve
in both upper limbs. On the left side, the proximal end of Comparative Anatomy
extensor digitorum was fused with extensor digiti minimi Extensor digiti minimi is similarly variable in the apes, with
and extensor carpi ulnaris (Smith et al. 2015). frequent connections to digit four (Barnard 1875; Chapman
In a female fetus with triploidy, Moen et al. (1984) found 1878, 1879, 1880, 1900; Kohlbrügge 1890–1892; Hepburn
that the extensor digitorum tendons were doubled bilat- 1892; Beddard 1893; Dwight 1895; Primrose 1899, 1900;
erally. Doubling of the extensor digitorum tendon to the Sonntag 1923, 1924; Sullivan and Osgood 1927; Straus
fourth fnger and the ffth fnger tendon arising from the 1941; Raven 1950; Miller 1952; Kallner 1956; Preuschoft
fourth has been observed in infants with Neu-Laxova syn- 1965; Kaneff 1980a; Lewis 1989; Gibbs 1999; Michilsens
drome (Shved et al. 1985). et al. 2009; Oishi et al. 2009; Diogo et al. 2010, 2012,
Mieden (1982) describes two male fetuses with cyclopia 2013a,b, 2017). Additional origins from the ulna have been
and alobar holoprosencephaly. On the right side of one spec- observed in gibbons, orangutans, and common chimpan-
imen, the tendon of extensor digitorum to the fourth digit zees (Deniker 1885; Kohlbrügge 1890–1892; MacDowell
on the right side was doubled. Bersu et al. (1976) describe 1910; Straus 1941; Michilsens et al. 2009; Gibbs 1999;
a male infant with Hanhart syndrome. On the right side of Diogo et al. 2012, 2013a,b).
this specimen, the wrist and hand were defcient. Extensor
digitorum had a normal attachment to the distal phalanx of Variations
the ffth digit, but the tendons to the other digits were fused Description
and attached to connective tissue distal to the carpus. The entire muscle may be absent (Macalister 1875; Knott
1883a; Kaneff 1980a; Bergman et al. 1988; Akita and
Prevalence Nimura 2016b; Standring 2016). When absent, it may be
Extensor digitorum was variable (e.g., extra slips/ten- replaced by a slip of extensor digitorum or extensor carpi
dons, medio-lateral displacement of the extensor hood) ulnaris (Macalister 1875; Bergman et al. 1988). The muscle
in 1 out of 24 individuals with trisomy 13 (4.2%) and in belly or tendon may be doubled (Macalister 1875; Knott
9 out of 26 individuals with trisomy 18 (34.6%). Extensor 1883a; Mori 1964; Bergman et al. 1988; Perkins and Hast
digitorum sent a tendon to the ffth digit in all individuals 1993; Zilber and Oberlin 2004; Akita and Nimura 2016b).
with trisomy 13, 18, and 21 (Smith et al. 2015). In a study The tendon may also be tripled (Macalister 1875; Mori
of six upper limbs from three infants with Neu-Laxova 1964; Perkins and Hast 1993; Zilber and Oberlin 2004).
syndrome, Shved et al. (1985) found that the tendon of Extensor digiti minimi often sends a slip to digit four
extensor digitorum to the fourth fnger was doubled in (Macalister 1875; Knott 1883a; Le Double 1897; Mori 1964;
two upper limbs (33%) and the tendon to the ffth fnger Kaneff 1980a; Bergman et al. 1988; Standring 2016). Le
arose from the tendon to the fourth fnger in three upper Double (1897) noted a case in which this muscle had four
limbs (50%). tendons and sent two to digit fve, one to digit four, and one
to digit three. When the muscle is divided into two, extensor was single in 2%–35% of hands, doubled in 63%–87%
digiti minimi accessorius (Krause) may be used to refer to of hands, tripled in 2%–8% of hands, and quadrupled in
the radial slip that inserts onto the dorsal carpal ligament 7%–10% of hands.
(Knott 1883a; Mori 1964; Akita and Nimura 2016b). In a study of 41 hands, Tanaka et al. (2007) found that
This muscle may be fused with extensor digitorum or extensor digiti minimi had a single tendon in one speci-
its tendon (Macalister 1875; Bergman et al. 1988; Akita men (2.4%), double tendon in 29 specimens (70.7%), and a
and Nimura 2016b; Standring 2016). There may be a slip triple tendon in nine specimens (21.9%). From dissections
to extensor digiti minimi from the lateral epicondyle of the of 100 upper limbs, Suwannakhan et al. (2016) found that
humerus or an additional origin from the posterior aspect the extensor digiti minimi was absent in one hand (1%).
of the ulna (Bergman et al. 1988; Akita and Nimura 2016b). The tendons split at insertion into two tendons in 75 hands
Another slip from the ulna may be present, which attaches (75%), remained single in 16 hands (16%), and inserted as
to the base of the ffth metacarpal (Bergman et al. 1988; a triple tendon in four hands (4%). In four hands (4%), the
Akita and Nimura 2016b). Sato (1969) notes that this mus- tendon originated as single, split into two, then rejoined to
cle can attach elsewhere without passing into the aponeuro- insert as one tendon (Suwannakhan et al. 2016). In a study
sis of the ffth digit. of 110 upper limbs, Bharambe et al. (2017) found that in
16% of cases the tendon to digit fve split into two and in 3%
Prevalence of cases three tendons were present, two going to digit fve
Macalister (1875) notes that Wood found that extensor digiti and one going to digit four.
minimi sent a tendon to both digits four and fve in 13 out
of 106 cases (12.3%). Macalister (1875) observed a tendon Anomalies
to both digits four and fve in 1 out of 14 Irish individuals
(7.1%). Le Double (1897) noted that extensor digiti minimi Description
sent a tendon to digit four in 12 out of 144 cases (8.3%). In a fetus with craniorachischisis, Alghamdi et al. (2017)
Mori (1964) found that the tendon of extensor digiti observed that the left extensor digiti minimi attached to
minimi was single in 6% of cases, doubled in 82% of cases, both digits four and fve. Similarly, in a trisomy 18 cyclopic
and tripled in 12% of cases. Extensor digiti minimi acces- fetus, Smith et al. (2015) found that the left extensor digiti
sorius (see description above) was present in 4% of cases minimi attached to both digits four and fve. “Extensor
(Mori 1964). Mori (1964) also noted a slip to the fourth digiti quinti proprius” was absent bilaterally in a neonate
digit (extensor digiti minimi et quarti) in 2% of cases. The with trisomy 18 (Aziz 1979). It was also absent bilaterally
attachment variation described by Sato (1969) was present in a neonate with trisomy 13 (Aziz 1980). Doubling of the
in 2 out of 344 limbs (0.87%) from Kyushu-Japanese males extensor digiti minimi tendon and an origin of this mus-
and present in 2 out of 236 limbs (0.85%) from females. An cle from extensor digitorum has been observed in infants
accessory head of extensor digiti minimi was present in 19 with Neu-Laxova syndrome (Shved et al. 1985). Bersu et al.
out of 312 limbs (6.09%) from males and present in 13 out (1976) describe a male infant with Hanhart syndrome. On
of 220 limbs (5.91%) from females. the right side of this infant, the wrist and hand were def-
Kaneff (1980a) examined 300 upper limbs and found cient. Extensor digiti minimi was absent.
that extensor digiti minimi was absent in 1% of cases, sent
a tendon to digit fve only in about 94% of cases, and sent Prevalence
tendons to both digits four and fve in about 5% of cases. In their literature review, Smith et al. (2015) found that
Based on dissections of 80 hands from 40 individuals, extensor digiti minimi was variable (e.g., gives off extra
Perkins and Hast (1993) found that the tendon of exten- tendons, tendon splits into thirds, medio-lateral displace-
sor digiti minimi split into two or three slips in 77 hands ment of tendons, tendons doubled) in 2 out of 24 individu-
(96%). In a study of 50 hands (26 left, 24 right), Zilber and als with trisomy 13 (8.3%), 12 out of 26 individuals with
Oberlin (2004) found that the insertion tendon of extensor trisomy 18 (46.2%), and one out of seven individuals with
digiti minimi was single in 25% of right hands and 31% trisomy 21 (14.3%). These authors also found that extensor
of left hands, doubled in 71% of right hands and 69% of digiti minimi was absent in 1 out of 24 individuals with tri-
left hands, and tripled in 4% of right hands. Extensor digiti somy 13 (4.2%) and in 2 out of 26 individuals with trisomy
minimi set a tendon to digit four in one hand (2%) of cases 18 (7.7%). In a study of six upper limbs from three infants
(Zilber and Oberlin 2004). with Neu-Laxova syndrome, Shved et al. (1985) found that
Zilber and Oberlin (2004) also provide a comprehensive the tendon of extensor digiti minimi was doubled in three
literature review of extensor tendon variation, summarizing upper limbs (50%), and this muscle took an origin from
results from a study of 150 hands by Mestdagh et al. (1985), extensor digitorum in all six upper limbs (100%).
a study of 50 hands by Godwin and Ellis (1992), a study of
43 hands by von Schroeder and Botte (1995), a study of 181 Clinical Implications
hands by el-Badawi et al. (1995), and a study of 548 hands Understanding variation in extensor digiti minimi tendons
by Hirai et al. (2001). Among these studies, the prevalence is important for planning and performing tendon transfer
ranges are as follows. The extensor digiti minimi tendon surgery (Zilber and Oberlin 2004; Tanaka et al. 2007).
extensOr inDicis (figures 3.16 anD 3.17) and third digits, and another tendon slip that inserts onto
the fourth and ffth digits (Suwannakhan et al 2020).
See also: Extensor medii digiti, Extensor digitorum brevis Kaneff (1980a) suggested that during human development,
manus extensor indicis can extend from digits two through four,
Synonyms and usually reduces to the region of digit two only in later
ontogenetic stages.
This muscle is also referred to as extensor indicis proprius
Accessory muscles and tendons associated with extensor
(Bergman et al. 1988) or the indicator (Todd 1839).
indicis have been designated as distinct muscles. Extensor
Typical Presentation pollicis-et-indicis has attachments to digits one and two
(Macalister 1875; Cauldwell et al. 1943). Extensor digiti
Description quarti sends a slip to digit four (Cauldwell et al. 1943).
Extensor indicis originates from the posterior aspect of Extensor indicis is also associated with extensor digitorum
the ulna and the neighboring interosseous membrane brevis manus (extensor indicis brevis) and extensor medii
(Standring 2016). It inserts via a tendon onto the extensor digiti (see the entries for these muscles).
expansion of digit two (Standring 2016).
Prevalence
Innervation Extensor indicis goes to digit two in 87%–95% of humans
Extensor indicis is innervated by the posterior interosseous according to Straus (1941). Wood (1865, 1866, 1867b, 1868)
nerve (Standring 2016). found that out of 276 upper limbs, 6.5% had duplicated
tendons to digit two, 6.9% had an abbreviated extensor
Comparative Anatomy indicis, 1.8% had attachments to both digits 1 and 2 (exten-
Extensor indicis has a similar typical presentation in the sor-pollicis-et-indicis), 7.2% had attachments to digit three
apes, and accessory insertions onto digits three, four, (extensor digiti medii), and 0% of cases exhibited a com-
and/or fve are common (Bischoff 1870; Macalister 1871; plete absence of this muscle. Cauldwell et al. (1943) found
Chapman 1880, 1900; Deniker 1885; Kohlbrügge 1890– that within 263 upper limbs, 2.7% had duplicated tendons
1892; Hepburn 1892; Beddard 1893; Primrose 1899, 1900; to digit two, 1.1% had an abbreviated extensor indicis, 0.7%
MacDowell 1910; Sullivan and Osgood 1927; Straus 1941; had an extensor pollicis-et-indicis, 6.5% had an extensor
Raven 1950; Kallner 1956; Jouffroy 1971; Kaneff 1980a,b; digiti medii, 0.38% had an attachment to digit four (exten-
Gibbs 1999; Michilsens et al. 2009; Oishi et al. 2009; Diogo sor digiti quarti), and 0% of cases exhibited a complete
et al. 2010, 2012, 2013a,b, 2017). Origins from the radius absence of extensor indicis. Kaneff (1980b) found that out
and/or humerus have been observed in all apes except of 300 upper limbs, extensor indicis was missing in 1% of
bonobos (Gratiolet and Alix 1866; Barnard 1875; Deniker cases, and an attachment to digit three was exhibited in
1885; Sonntag 1923; Diogo et al. 2010, 2012, 2013a,b). 12.97% of cases.
Mori (1964) observed that the distal end of the muscu-
Variations lar portion of extensor indicis was distal to the radiocarpal
Description joint in 48% of cases, was on the radiocarpal joint in 8%
Extensor indicis may instead originate from the radius, of cases, and did not reach the radiocarpal joint in 44% of
carpal bones, or the interosseous membrane (Wood 1868; cases. Mori (1964) also observed that the insertion tendon
Macalister 1875; Bergman et al. 1988; Akita and Nimura was doubled in 12% of cases and that the distal half the
2016b). Occasionally this muscle may be doubled or have muscle was doubled, and each portion had its own tendon
two distinct heads, and its tendon may also be dupli- in 8% of cases.
cated (Wood 1865, 1866, 1867b, 1868; Macalister 1875; In a study of 50 hands (26 left, 24 right), Zilber and
Cauldwell et al. 1943; Mori 1964; Bergman et al. 1988; Oberlin (2004) found that extensor indicis was absent in
Perkins and Hast 1993; Tountas and Bergman 1993; two hands (4%), the insertion tendon was single in 79%
Standring 2016). Its tendon may also be tripled (el-Badawi of right hands and 77% of left hands, and the tendon was
et al. 1995; von Schroeder and Botte 1995). It may receive doubled in 17% of right hands and 19% of left hands. These
a slip from extensor digitorum or join with extensor medii authors also provide a comprehensive literature review
digiti (Wood 1868; Macalister 1875). Extensor indicis may of extensor tendon variation, summarizing results from a
be absent (Macalister 1875; Mestdagh et al. 1985; Bergman study of 150 hands by Mestdagh et al. (1985), a study of 50
et al. 1988; Tountas and Bergman 1993; el-Badawi et al. hands by Godwin and Ellis (1992), a study of 43 hands by
1995; Zilber and Oberlin 2004; Akita and Nimura 2016b; von Schroeder and Botte (1995), a study of 181 hands by
Fernandes et al. 2016; Bharambe et al. 2017). el-Badawi et al. (1995), and a study of 548 hands by Hirai
Suwannakhan et al. (2020) report the presence of what et al. (2001). Among these studies, the prevalence ranges
they term an extensor digitorum profundus complex with are as follows. The tendon of extensor indicis was single
two bellies originating from the distal ulna in a cadaver. in 77%–93% of hands, doubled in 5%–16% of hands, and
The frst belly resembled extensor indicis, and the second tripled in 4%–7% of hands. Extensor indicis was absent in
belly gave off one tendon slip that inserts onto the second 1% of hands.
Based on dissections of 80 hands from 40 individuals, trisomy 18 (46.2%), and 2 out of 7 individuals with trisomy
Perkins and Hast (1993) found that the tendon of exten- 21 (28.6%). Extensor pollicis-et-indicis was present in one
sor indicis split into two slips in 30 hands (37.5%). From out of seven individuals with trisomy 21 (14.3%). Extensor
a study of 34 upper limbs from 17 cadavers, extensor indi- indicis was absent in 4 out of 24 individuals with trisomy
cis was absent in two upper limbs (5.9%) from the same 13 (16.7%).
cadaver, and extensor indicis brevis was present on the
right side (Fernandes et al. 2016). From dissections of 100 Clinical Implications
upper limbs, Suwannakhan et al. (2016) found that extensor Understanding variations in the extensor indicis tendons
pollicis et indicis was present in six hands (6%) from four is important for planning and performing tendon transfer
cadavers (7.7%), and in four cases (4%), extensor indicis was surgery (Zilber and Oberlin 2004; Fernandes et al. 2016;
absent and replaced by either extensor pollicis et indicis or Matsumae et al. 2018).
extensor indicis et medii communis. In a study of 110 upper
limbs, Bharambe et al. (2017) found extensor indicis pro-
prius was duplicated in 1% of cases and absent in 2% of extensOr MeDii Digiti (figure 3.17)
cases.
Based on a study of 176 upper limbs, Georgiev et al. See also: Extensor indicis
(2018b) found that extensor indicis exhibited the typical
presentation in 147 cases (83.5%) In the 29 other limbs Synonyms
(16%), variations included the replacement of extensor indi- This muscle is also referred to as extensor digiti medii
cis with extensor indicis brevis, the coexistence of those (Bergman et al. 1988) or extensor medii proprius (Bharambe
two muscles, and the presence of accessory tendons. et al. 2017).

Description This muscle is only present as a variation or anomaly.
Mieden (1982) describes two male fetuses with cyclopia and
Comparative Anatomy
alobar holoprosencephaly. On the left side of one specimen,
the tendon of extensor indicis was doubled. On the right Extensor medii digiti is usually not present as a distinct
side of a male infant with Hanhart syndrome, the wrist and muscle in the apes (Diogo et al. 2010, 2012, 2013a,b, 2017).
hand were defcient and extensor indicis was absent (Bersu
et al. 1976). In a study of specimens with triploidy, exten- Variations
sor indicis had an extra tendon to the extensor retinaculum Description
bilaterally in a male fetus and was absent bilaterally in a Extensor medii digiti is considered a variation of extensor
female fetus (Moen et al. 1984). indicis (Macalister 1875). It originates from the ulna distal
In a male neonate with Meckel syndrome, Pettersen to the origin of extensor indicis and sends a tendon to digit
(1984) observed that extensor indicis blended into fascia on three, or sometimes to both digits two and three (Macalister
the dorsum of the right hand where it partly attached to a 1875; Bergman et al. 1988; Carlos et al. 2011; Akita and
small feshy mass, from which a tendon arose that attached Nimura 2016b). It may originate from the radius or the
normally onto digit two. On the left side, extensor indicis carpus (Macalister 1875). It can insert onto the metacarpo-
was represented by a diminutive slip on the dorsum of the phalangeal joint (Lee et al. 2019b) or base of the proximal
wrist and had no insertion tendon. In a trisomy 18 cyclopic phalanx (Campos et al. 2011; Carlos et al. 2011) of digit
fetus, the extensor indicis tendon was doubled and went to three. Extensor medii digiti can sometimes be fused with
both digits two and three (Smith et al. 2015). In a fetus with extensor indicis (Macalister 1875; Bergman et al. 1988).
craniorachischisis dissected by Alghamdi et al. (2017), the Extensor indicis et medii communis is a similar muscle that
left extensor indicis joined the tendon of extensor digito- arises from the ulna and inserts onto digits two and three
rum to attach onto digit two. On the right side, extensor (von Schroeder and Botte 1995; Yammine 2015a; Akita and
indicis only sent a tendon to digit four, not digit two. Nimura 2016b).
Vaida et al. (2021) report a case in which both exten-
Prevalence sor medii proprius and extensor indicis et medii commu-
In their literature review, Smith et al. (2015) found that nis were present in the same forearm. The tendon of the
extensor indicis was variable (e.g., variable origin, diminu- former coursed through the second extensor compartment
tive muscle, doubled muscle or tendons) in 7 out of 24 indi- between the tendons of extensor carpi radialis longus
viduals with trisomy 13 (29.2%), 12 out of 26 individuals and brevis. The latter had three tendons, two to digit two
with trisomy 18 (46.2%), and 2 out of 7 individuals with tri- and one to digit three. A third muscle was present with
somy 21 (28.6%). The extensor indicis tendon was doubled a common origin from extensor carpi radialis brevis and
and went to both digits two and three in one neonate out an insertion into the tendon of extensor indicis et medii
of 24 with trisomy 13 (4.2%), 12 out of 26 individuals with communis.
Innervation extensOr DigitOruM brevis Manus (figure 3.17)

Extensor medii digiti is innervated by the posterior interos- See also: Extensor indicis
seous nerve (Carlos et al. 2011).
Synonyms
Prevalence This muscle is also referred to as extensor brevis digitorum
Le Double (1897) stated that extensor medii digiti occurs manus (Knott 1883a), extensor anomalus (Sauser 1935), le
in 10% of humans. Wood (1865, 1866, 1867b, 1868) found muscle manieux (Le Double 1897); extensor indicis brevis
that within 276 upper limbs, 7.2% of extensor indicis (Gahhos and Ariyan 1983), extensor indicis brevis manus
muscles had attachments to digit three. Macalister (1875) (Standring 2016), extensor brevis digiti indicis vel medii
states that Wood found extensor medii digiti in 11 out of (Albinus 1758), indicator biceps (Gantzer), and indicator
102 subjects (10.8%). Cauldwell et al. (1943) found that anomalus brevis (Otto) (Macalister 1875).
within 263 upper limbs, 6.5% had an extensor medii
digiti. Based on dissections of 80 hands from 40 individ- Typical Presentation
uals, Perkins and Hast (1993) found that extensor digiti This muscle is only present as a variation or anomaly.
medii originated with extensor indicis from the ulna and
sent a tendon to the dorsal expansion of digit three in 17 Comparative Anatomy
hands (21%). Carlos et al. (2011) observed the presence of Extensor digitorum brevis manus is generally absent in
this muscle in 7 out of 94 limbs (7.4%), from 6 out of 47 the apes (Diogo et al. 2010, 2012, 2013a,b, 2017). The only
cadavers (12.8%). Extensor indicis et medii communis has cases of its presence were reported by Lewis (1989). Lewis
a prevalence of 1.6% (Yammine 2015a). From dissections (1989) observed this muscle in an orangutan, and it was
of 100 upper limbs from 52 cadavers, Suwannakhan et al. associated with the tendon of extensor indicis to the third
(2016) found that extensor indicis et medii communis was digit (Diogo et al. 2013b). Lewis (1989) also reported the
present in six hands (6%) from four cadavers (7.7%) and presence of this muscle in a gorilla where it originated from
“extensor medii proprius” was present in eight hands (8%) the extensor indicis tendon to digit two and inserted onto
from fve cadavers (9.6%). In a study of 110 upper limbs, digit three (Diogo et al. 2010).
Bharambe et al. (2017) found extensor medii proprius in
4% of cases. Variations
Description
Anomalies Extensor digitorum brevis manus is considered a variation
Description of extensor indicis, and it is often joined with this muscle,
particularly upon insertion (Ogura et al. 1987; Yammine
Windle (1893) reported the presence of extensor medii digiti
2015b). The presence of extensor digitorum brevis manus
in one fetus with anencephaly. This muscle was present
may signify the failure of a proximal migration of the ulno-
bilaterally in a neonate with trisomy 18 (Aziz 1979). Dunlap
carpal parts of the antebrachial muscle mass (Sauser 1935;
et al. (1986) also found extensor medii digiti in individuals
Ross and Troy 1969). It may also originate from the deep
with trisomy 18. In two cases, it sent an additional tendon
portion of the forearm extensor precursor muscle, which is
to digit two. In another case, the entire extensor medii digiti
prone to variations (Yammine 2015b).
muscle was doubled. In another case, an “extensor medii
Extensor digitorum brevis manus originates from the
digiti et quarti” tendon was present. It branched from the
ulnar portion of the carpus and the ligaments that connect
extensor medii digiti tendon over the metacarpal of digit
the bones (capitate, hamate, lunate, triquetrum) on this side
three and inserted on the metacarpophalangeal joint of digit
of the carpus (Knott 1883a; Cauldwell et al. 1943; Bergman
four (Dunlap et al. 1986).
et al. 1988; Akita and Nimura 2016b). It may also origi-
nate from the distal radius or ulna (Macalister 1875; Knott
Prevalence 1883a; Ogura et al. 1987; Paraskevas et al. 2002; Gonzalez
In a sample of 20 limbs from ten individuals with trisomy and Netscher 2016). It can be comprised of between one
18, Dunlap et al. (1986) found that extensor medii digiti was to four bundles and is most commonly represented by a
present in ten limbs (50%). Smith et al. (2015) note in their bundle going to digit two, and sometimes a bundle that
literature review that extensor medii digiti is present in 5 goes to digit three, four, and/or fve (Macalister 1875; Knott
out of 26 individuals with trisomy 18 (19.2%). 1883a; Ross and Troy 1969; Ogura et al. 1987; Bergman
et al. 1988; Paraskevas et al. 2002; Doyle and Botte 2003;
Clinical Implications Ranade et al. 2008; Akita and Nimura 2016b; Lee et al.
Campos et al. (2011) argue that understanding the presence 2019b). The tendon of extensor digitorum brevis manus
and anatomy of supernumerary extensor muscles, includ- inserts onto the extensor expansion or the metacarpals of
ing the extensor medii digiti they describe (but do not refer the target digit (Knott 1883a; Ogura et al. 1987; Bergman
to by name), is important for diagnosing carpal tunnel et al. 1988; Rodríguez-Niedenführ et al. 2002; Akita and
syndromes. Nimura 2016b).
Gahhos and Ariyan (1983) report an “extensor indicis Prevalence

brevis” originating from the joint capsule and ligaments In a sample of ten limbs from fve individuals with trisomy
of the scaphoid and lunate that attached to the extensor 13, Dunlap et al. (1986) found that extensor digitorum bre-
expansion of the second digit. Fernandes et al. (2016) vis manus was present in three limbs (30%).
report the presence of extensor indicis brevis on the
right side of one cadaver that was associated with bilat- Clinical Implications
eral absence of extensor indicis proprius. Suwannakhan Extensor digitorum brevis manus can be mistaken for a
et al. (2016) report a case in which extensor digitorum dorsal wrist ganglion, tendon sheath cyst, infectious mass,
brevis manus was present with extensor medii proprius bone spur (exostosis), synovitis, hemangioma, rheumatoid
and the tendon of the latter inserted onto the tendon of tenosynovitis, or a soft tissue tumor (Gahhos and Ariyan
the former. 1983; Rodríguez-Niedenführ et al. 2002; Ammendolia
2008; Ranade et al. 2008; Gonzalez and Netscher 2016).
Innervation Though largely asymptomatic, extensor digitorum brevis
Extensor digitorum brevis manus is innervated by the manus may present as a tender, painful, and/or swollen
posterior interosseous nerve (Rodríguez-Niedenführ et al. mass (Gahhos and Ariyan 1983), particularly if it is present
2002). on the dominant hand, potentially due to compression of
the hypertrophied muscle in the fourth dorsal compartment
Prevalence (Ross and Troy 1969; Patel et al. 1989).
Macalister (1875) found extensor digitorum brevis manus in
1 out of 15 subjects (6.7%) and states that Wood found the
muscle in 1 out of 36 (2.8%). Smith (1896) reports a high
extensOr pOllicis lOngus (figures 3.16 anD 3.17)
prevalence (70%) for this muscle, as this author claims to Synonyms
have found it in 35 out of 50 hands. Cauldwell et al. (1943) This muscle is also referred to as extensor pollicis
found that within 263 upper limbs, three cases (1.1%) had major, extensor secondi internodii pollicis, or cubito sus-
an abbreviated extensor indicis, and one of these muscles phalangettien du pouce (Todd 1839; Macalister 1875;
was designated as an extensor digitorum brevis manus as it Bergman et al. 2008).
was completed separated from the ulna.
Gama (1983) found extensor digitorum brevis manus Typical Presentation
in 38 out of 3404 adults (1.1%). Ogura et al. (1987) Description
observed this muscle in 17 out of 559 hands from 286
Extensor pollicis longus originates from the middle third
cadavers (3%). Bergman et al. (1988) provide a prevalence
of the posterior ulna and from the neighboring interosseous
of 9%. Perkins and Hast (1993) found extensor digiti
membrane (Standring 2016). It inserts via a tendon onto the
brevis manus in 1 out of 80 hands (1.25%). Rodríguez-
base of the distal phalanx of the frst digit (Standring 2016).
Niedenführ et al. (2002) observed the presence of exten-
sor digitorum brevis manus in 3 out of 128 cadavers Innervation
(2.3%). Ranade et al. (2008) found this muscle in 3 out of
Extensor pollicis longus is innervated by the posterior inter-
72 upper limbs (4.2%).
osseous nerve (Standring 2016).
Yammine (2015b) conducted a meta-analysis of 26
studies and found an overall crude cadaveric prevalence Comparative Anatomy
of 4% and an overall true cadaveric prevalence of 2.5%.
Extensor pollicis longus has a similar typical presentation
Akita and Nimura (2016b) list a prevalence of 1.6%–3.2%,
in the apes, with occasional insertions onto the proximal
citing Kosugi et al. (1984) and Yoshida et al. (1984).
phalanx, metacarpal, or metacarpophalangeal joint of the
Gonzalez and Netscher (2016) list a prevalence of 2%–3%
frst digit (Champneys 1872; Deniker 1885; Sutton 1883;
and state that its presence is found more often in males.
Kohlbrügge 1890–1892; Hepburn 1892; Chapman 1900;
From dissections of 100 upper limbs, Suwannakhan et al.
Sonntag 1924; Straus 1941; Raven 1950; Kallner 1956;
(2016) found that extensor digitorum brevis manus was
Ziegler 1964; Gibbs 1999; Michilsens et al. 2009; Diogo
present in one cadaver (1%). In a study of 110 upper limbs,
et al. 2010, 2012, 2013a,b, 2017). A slip to digit two has been
Bharambe et al. (2017) found extensor digitorum brevis
observed in gorillas and common chimpanzees (Hepburn
manus in 2% of cases.
1892; Kaneff 1980a; Gibbs 1999; Diogo et al. 2010, 2013a).
Description
Dunlap et al. (1986) state that individuals with trisomy 13 Description
exhibit a diminution of the extensor digitorum profundus Extensor pollicis longus may be doubled (Macalister 1875;
complex, which may lead to the presence of extensor digi- Bergman et al. 1988; Akita and Nimura 2016b). Its ten-
torum brevis manus. don may also be split or doubled (Wood 1868; Macalister
1875; Mori 1964; Palatty et al. 2018). Accessory tendons of with the second digit) in 2 out of 24 individuals with tri-
this muscle may pass through the frst, second, fourth, or somy 13 (8.3%), 13 out of 26 individuals with trisomy 18
an additional separate dorsal compartment (as opposed to (50%), and 1 out of 7 individuals with trisomy 21 (14.3%).
the typical third dorsal compartment) (Sevivas et al. 2009; In a study of six upper limbs from three infants with Neu-
Türker et al. 2010; Rubin et al. 2011; Kim et al. 2016). It Laxova syndrome, Shved et al. (1985) found that extensor
may be connected via slips to the proximal phalanx of digit pollicis longus shared a common muscle belly with exten-
one, extensor digitorum, extensor pollicis brevis, or extensor indicis in fve upper limbs (83%).
sor indicis (Wood 1868; Macalister 1875; Knott 1883a;
Mori 1964; Bergman et al. 1988; Akita and Nimura 2016b). Clinical Implications
An accessory extensor (sometimes termed extensor polli- An accessory extensor pollicis longus may cause pain at
cis tertius) between extensor pollicis longus and extensor the wrist (Beatty et al. 2000). Doubling and/or variable
indicis can be present as a rare variation (Le Double 1897; insertion of its tendon may limit the ability to extend the
Bergman et al. 1988; Bluth et al. 2011; Akita and Nimura frst digit (Masada et al. 2003; Sawaizumi et al. 2003;
2016b). Extensor pollicis longus may be absent (Bergman Türker et al. 2010), cause hyperextension of the frst digit
et al. 1988; Akita and Nimura 2016b). (Alsharif et al. 2017), or be mistaken for a ruptured tendon
(Masada et al. 2003). Variation in the course of the exten-
Prevalence sor pollicis longus tendon can also cause tenosynovitis that
Mori (1964) reports that in 5% of cases the insertion tendon may mimic intersection syndrome or de-Quervain’s disease
of extensor pollicis longus was doubled (misprinted as e. (Rubin et al. 2011).
p. brevis). The tendon inserted onto the distal phalanx of
digit one in 80% of cases, fused with the tendon of exten-
extensOr pOllicis brevis (figure 3.16)
sor pollicis brevis and inserted into the distal phalanx in
20% of cases, and was doubled with one part inserting onto Synonyms
the proximal phalanx and the other onto the distal phalanx This muscle is also referred to as extensor primi minor,
in 15% of cases (Mori 1964). Palatty et al. (2018) studied extensor primi internodii pollicis, extensor minor pollicis
variations in the extrinsic thumb muscles in 30 adult and manus, or cubito sus-phalangettien du pouce (Todd 1839;
20 fetal hands. The tendon of extensor pollicis longus was Bergman et al. 2008).
single in 43 hands (86%) and doubled in seven hands (14%).
Anomalies Description
Description Extensor pollicis brevis originates from the posterior aspect
On the left side of one male fetus with anencephaly, Windle of the distal radius and from the neighboring interosseous
(1983) observed that extensor pollicis longus was doubled, membrane (Standring 2016). It inserts via a tendon onto the
with the accessory muscle originating with extensor indicis. base of the proximal phalanx of digit one (Standring 2016).
On the left upper limb of a fetus with craniorachischisis,
Alghamdi et al. (2017) found that the tendon of exten- Innervation
sor pollicis longus split at the base of the frst metacarpal Extensor pollicis brevis is innervated by the posterior inter-
and then rejoined at the metacarpophalangeal joint before osseous nerve (Standring 2016).
inserting onto the base of the distal phalanx of digit one.
This morphology was also observed on both sides of the Comparative Anatomy
trisomy 18 cyclopic fetus studied by Smith et al. (2015). Among the apes, extensor pollicis brevis is present as a dis-
Extensor pollicis longus has been observed to share a tinct muscle belly only in gibbons, and it is partially blended
common muscular belly with extensor indicis in infants with abductor pollicis longus (Bischoff 1870; Kohlbrügge
with Neu-Laxova syndrome (Shved et al. 1985). On the left 1890–1892; Michilsens et al. 2009; Diogo et al. 2010, 2012,
arm of a male neonate with Meckel syndrome, Pettersen 2013a,b, 2017).
(1984) observed that extensor pollicis brevis joined and
Variations
inserted with extensor pollicis longus. In a male infant with
Hanhart syndrome, the wrist and hand were defcient on Description
the right side and extensor pollicis longus inserted into the The tendon of extensor pollicis brevis may have an addi-
capsule of the radiocarpal joint (Bersu et al. 1976). tional attachment onto the base of the distal phalanx of the
frst digit, which commonly presents as a slip that join the
Prevalence tendon of extensor pollicis longus (Wood 1868; Macalister
In their literature review, Smith et al. (2015) found that 1875; Knott 1883a; Mori 1964; Bergman et al. 1988; Nayak
extensor pollicis brevis and longus were variant (e.g., fusion et al. 2009b; Akita and Nimura 2016b; Standring 2016). The
of the two muscles, doubling of the tendons, muscles split tendon can also bifurcate near its insertion, with the sec-
into more than one part, variable attachments, connections ond insertion attaching to the base of the frst metacarpal
(Macalister 1875; Dawson and Barton 1986; Bergman et al. Anomalies

1988; Akita and Nimura 2016b). The tendon may also be Description
tripled (Nayak et al. 2009b).
Extensor pollicis brevis may be fused with abductor
Alghamdi et al. (2017) noticed that the muscle belly of
pollicis longus (Wood 1867a, 1868; Macalister 1875; Knott
extensor pollicis brevis was fused with abductor pollicis
1883a; Inoue 1934; Mori 1964; Dawson and Barton 1986;
longus. This muscle narrowed into three tendons which
Bergman et al. 1988; Akita and Nimura 2016b; Standring
attached to connective tissue on the palmar surface of the
2016). It may also be completely absent (Wood 1868;
first metacarpal to form a u-shaped structure with the ten-
Macalister 1875; Knott 1883a; Stein 1951; Mori 1964;
don of flexor pollicis longus.
Dawson and Barton 1986; Bergman et al. 1988; Akita and
In both a female neonate (Aziz 1979) and a female
Nimura 2016b; Standring 2016).
fetus with trisomy 18 (Alghamdi et al. 2018), the proxi-
Prevalence mal ends of extensor pollicis brevis and abductor pollicis
longus were fused on both sides of the body. The extensor
Wood (1867a) noted that this muscle was absent in 1 out
pollicis brevis tendon was also bifurcated bilaterally in
of 36 cases (2.8%) and later noted that it was absent in 3
the neonate, with both tendons inserting onto the distal
out of 36 cases (8.3%) (Wood 1868; Macalister 1875). In a
phalanx (Aziz 1979). On the left side of a trisomy 18 cyc-
study of 84 wrists, Stein (1951) found that extensor pollicis
lopic fetus, Smith et al. (2015) noted that extensor pollicis
brevis was absent in 7% of wrists, and an accessory ten-
brevis was diminutive and was also fused with abductor
don of extensor pollicis brevis was present in 4% of wrists.
pollicis longus. On the right side of this specimen, the
Mori (1964) found that extensor pollicis brevis was absent
muscle was absent.
in 2% of cases. It was fused with abductor pollicis longus in
In a male fetus with triploidy, extensor pollicis brevis was
10% of cases and extensor pollicis longus in 50% of cases.
absent bilaterally (Moen et al. 1984). Absence or “anoma-
The insertion tendon was split in 10% of cases. The tendon
lous structure” of extensor pollicis brevis has been observed
inserted onto the proximal phalanx of digit one in 58.3% of
in infants with Neu-Laxova syndrome (Shved et al. 1985).
cases, the distal phalanx in 21% of cases, and both phalan-
In a male neonate with Meckel syndrome, extensor pol-
ges of digit one in 20.8% of cases.
licis brevis inserted with abductor pollicis longus into the
Dawson and Barton (1986) dissected 16 arms from eight
first metacarpal on the right hand (Pettersen 1984). On the
cadavers and found that the muscle belly for extensor pol-
left arm, extensor pollicis brevis joined and inserted with
licis brevis was present in 13 arms (81.25%) and absent in
extensor pollicis longus. Bersu et al. (1976) describe a male
three (18.75%). The tendon of extensor pollicis brevis had
infant with Hanhart syndrome. On the right side of this
the typical presentation in ten arms (62.5%), had a branch
infant, the wrist and hand were deficient. Extensor pollicis
joining the tendon of abductor pollicis longus in two arms
brevis was absent.
(12.5%), and was attached to the base of the first metacarpal
in one arm (6.25%). In the three specimens where the mus-
cle belly was missing, the tendon was absent and replaced Prevalence
by an accessory tendon of abductor pollicis longus in one
arm (6.25%) and arose from the fat at the base of the first
extensor pollicis brevis and longus were variant (e.g.,
metacarpal in two arms (12.5%). Regarding insertion, the
fusion of the two muscles, doubling of the tendons, mus-
tendon inserted partially to the base of the proximal pha-
cles split into more than one part, variable attachments,
lanx and partially to the extensor hood of digit one in nine
connections with the second digit) in 2 out of 24 indi-
arms (56.25%), entirely to the base of the proximal phalanx
viduals with trisomy 13 (8.3%), 13 out of 26 individuals
in four arms (25%), and entirely to the extensor hood in
with trisomy 18 (50%), and 1 out of 7 individuals with
three arms (18.75%) (Dawson and Barton 1986).
trisomy 21 (14.3%). Extensor pollicis brevis was absent
In a study of 156 upper limbs, Nayak et al. (2009b)
in 4 out of 26 individuals with trisomy 18 (15.4%). In a
found that extensor pollicis brevis had a single tendon in
study of six upper limbs from three infants with Neu-
133 limbs (85.3%), double tendon in 17 limbs (10.9%), and
Laxova syndrome, Shved et al. (1985) found that exten-
triple tendon in 6 limbs (3.8%). In 11 limbs (7%), the acces-
sor pollicis brevis was aplasic in three upper limbs (50%)
sory tendon inserted onto the distal phalanx of digit one. In
and had an “anomalous structure” in three upper limbs
two limbs (1.3%), extensor pollicis brevis originated from
(50%).
the abductor pollicis longus tendon. In five limbs (3.2%),
the extensor pollicis brevis tendon attached to the base of
the first metacarpal bone (Nayak et al. 2009b). Palatty et al. Clinical Implications
(2018) studied variations in the extrinsic thumb muscles in Variation of the course of accessory tendons of extensor
30 adult and 20 fetal hands. The tendon of extensor pollicis pollicis brevis and compression of these tendons can con-
brevis was single in 47 hands (94%) and doubled in three tribute to de Quervain’s tenosynovitis (Giles 1960; Nayak
hands (6%) (Palatty et al. 2018). et al. 2009b).
abDuctOr pOllicis lOngus (figure 3.16) et al. 2015; Akita and Nimura 2016b; Standring 2016). It
is often fused with extensor pollicis brevis (Macalister
Synonyms 1875; Inoue 1934; Mori 1964; Standring 2016). Opponens
This muscle is also referred to as extensor ossis meta- pollicis may originate from the tendon of abductor pol-
carpi pollicis or cubito-radi sus-metacarpien (Todd 1839; licis longus (Bergman et al. 1988; Akita and Nimura
Macalister 1875). 2016b). Abductor pollicis longus may be reduced, only
having a radial origin (Bergman et al. 1988; Akita and
Typical Presentation Nimura 2016b). Palatty et al. (2018) found it absent in a
Description fetus.
Abductor pollicis longus originates from the posterior Paul and Das (2007) describe a case in which the abduc-
surface of the ulnar shaft, the neighboring interosseous tor pollicis longus muscle had a normal origin on the ulna
membrane, and the middle third of the posterior surface and radius, but after its tendon emerged from the extensor
of the radius (Standring 2016). It inserts via a tendon that retinaculum it continued as a small muscle belly that then
typically splits into two slips, with the frst attaching to the had a tendinous insertion onto the proximal phalanx of digit
base of the frst metacarpal and the second to the trapezium one. Ranade et al. (2017) also describe a similar case in
(Standring 2016). which abductor pollicis longus had a normal origin, but its
tendon split proximal to the wrist and one tendon inserted
Innervation onto the frst metacarpal, while the other tendon converted
Abductor pollicis longus is innervated by the posterior into a small muscle belly that inserted onto the proximal
interosseous nerve (Standring 2016). phalanx.
Abductor pollicis longus is associated with one rare
Comparative Anatomy supernumerary muscle, abductor pollicis tertius, which is
Abductor pollicis longus has a similar typical presentation also referred to as extensor atque abductor pollicis acces-
in the apes, extending from the radius and ulna to the frst sorius (Bergman et al. 1988). This muscle originates from
metacarpal and trapezium (Macalister 1871; Champneys the posterior surface of the radius along with abductor pol-
1872; Beddard 1893; Chapman 1900; Straus 1941; Miller licis longus, fuses with abductor pollicis brevis, and inserts
1952; Gibbs 1999; Michilsens et al. 2009; Diogo et al. 2010, into the frst metacarpal (Bergman et al. 1988; Akita and
2012, 2013a,b, 2017). An insertion into the sesamoid bone Nimura 2016b).
adjacent to the trapezium has been observed in all apes
Prevalence
except bonobos (Macalister 1873; Kohlbrügge 1890–1892;
Hepburn 1892; Dwight 1895; Sonntag 1923, 1924; Primrose According to Macalister (1875), Wood found that abduc-
1899, 1900; Straus 1941; Ziegler 1964; Diogo et al. 2010, tor pollicis longus sent a slip to the origin of abductor
2012, 2013a,b). An insertion into the scaphoid has been pollicis brevis in 7 out of 36 subjects (19.4%). In a study
observed in orangutans and common chimpanzees (Kallner of 84 wrists, Stein (1951) found that an accessory ten-
1956; Ziegler 1964; Diogo et al. 2013a,b). An insertion into don of abductor pollicis longus was present in 68% of
the proximal phalanx of the frst digit has been observed in wrists. In a study of 134 wrists, Baba (1954) found that
gibbons and gorillas (Owen 1868; Chapman 1878; Bischoff 132 wrists (98.5%) had accessory tendons of abductor
1880; Deniker 1885; Hepburn 1892; Pira 1913; Straus 1941; pollicis longus. Based on dissections of 80 hands from 40
Raven 1950; Preuschoft 1965; Kaneff 1980a,b; Diogo et al. individuals, Perkins and Hast (1993) found that abductor
2010, 2012). pollicis longus inserted via two or three tendons in 73
hands (91%).
Mori (1964) found that abductor pollicis longus was
Variations
fused with extensor pollicis brevis in 10% of cases. The
Description insertion tendon was doubled in 35% of cases, tripled in
The entire muscle and/or the tendon may be divided or 46% of cases, quadrupled in 14% of cases, and quintupled
doubled (Macalister 1875; Knott 1883a; Stein 1951; Baba in 4% of cases (Mori 1964). The tendon inserted onto the
1954; Bergman et al. 1988; Akita and Nimura 2016b; frst metacarpal only in 2% of cases, onto the frst meta-
Standring 2016; Jain et al. 2018). There may be three, four, carpal and the tendon of abductor pollicis brevis in 24% of
fve, or six tendons (Macalister 1875; Knott 1883a; Mori cases, and onto the frst metacarpal, the tendon of abductor
1964; Perkins and Hast 1993; El-Beshbishy and Abdel- pollicis brevis, and the trapezium in 40% of cases (Mori
Hamid 2013; Tewari et al. 2015). Slips from the tendon of 1964).
abductor pollicis longus may pass to the abductor pollicis El-Beshbishy and Abdel-Hamid (2013) examined varia-
brevis, opponens pollicis, the fexor retinaculum, thenar tions in abductor pollicis longus in 50 upper limbs. The tendon
fascia, scaphoid, trapezium, or proximal phalanx of digit was not found to be single in any case (0%). It had two tendons
one (Macalister 1875; Knott 1883a; Mori 1964; Bergman in 20 cases (40%), three tendons in 17 cases (34%), four ten-
et al. 1988; El-Beshbishy and Abdel-Hamid 2013; Tewari dons in nine cases (18%), fve tendons in two cases (4%), and
six tendons in two cases (4%). The lateral tendons in all cases hand were defcient. Abductor pollicis longus inserted into
inserted onto the base of the frst metacarpal bone (100%). the trapezium.
The insertion of the medial tendons varied from the antero-
lateral surface of the frst metacarpal (80%), lateral surface of Prevalence
the frst metacarpal (20%), the trapezium (80%), abductor pol- In their literature review, Smith et al. (2015) found that
licis brevis (60%), thenar fascia (40%), the carpometacarpal abductor pollicis longus was variant (e.g., doubling of the
joint of digit one (30%), and opponens pollicis (20%). muscle, variable attachments) in 4 out of 24 individuals
Tewari et al. (2015) examined variations in abductor pol- with trisomy 13 (16.7%) and in 9 out of 26 individuals
licis longus in 50 upper wrists. The tendon was single in two with trisomy 18 (34.6%), but was normal in seven indi-
cases (4%), doubled in 31 cases (62%), tripled in eight cases viduals with trisomy 21. Abductor pollicis longus was
(16%), quadrupled in eight cases (16%), and six tendons were absent in 1 out of 26 individuals with trisomy 18 (3.8%).
present in one hand (2%). At least one attachment onto the In a study of six upper limbs from three infants with Neu-
frst metacarpal was seen in all cases (100%), and an acces- Laxova syndrome, Shved et al. (1985) found that abduc-
sory insertion was observed onto the trapezium in 46 cases tor pollicis longus shared a common muscular belly with
(92%). Of all 126 tendons observed, 68 (53.9%) inserted onto extensor pollicis brevis in two upper limbs (33%).
the base of the frst metacarpal, 52 (41.3%) inserted onto the
trapezium, 4 (3.2%) inserted onto the opponens pollicis, and Clinical Implications
2 (1.6%) inserted onto the thenar fascia. Variation of the course of accessory tendons of abductor
Jain et al. (2018) examined variations in abductor pol- pollicis longus, and compression of these tendons, can con-
licis longus in 40 forearms from 20 cadavers. The tendon tribute to de Quervain’s tenosynovitis (Giles 1960).
of abductor pollicis longus was single in two cases (5%),
doubled in 30 cases (75%), and tripled in eight cases (20%).
It inserted onto the frst metacarpal in all hands (100%) and MUSCLES OF THE HAND
had a secondary insertion onto the trapezium in 20 hands
cOntrahentes DigitOruM Manus (figure 3.18)
(50%) (Jain et al. 2018). Palatty et al. (2018) studied varia-
tions in the extrinsic thumb muscles in 30 adult and 20 fetal Entry adapted by permission from Springer Nature
hands. The tendon of abductor pollicis longus was single in Customer Service Centre GmbH: Springer Current
38 hands (76%) and doubled or tripled in 11 hands (22%). Molecular Biology Reports, Muscles Lost in Our Adult
The muscle was absent in one of the fetal hands (2%). In one Primate Ancestors Still Imprint in Us: on Muscle Evolution,
adult male cadaver, the abductor pollicis longus had three Development, Variations, and Pathologies. E. Boyle, V.
slips of origin on the right side and two slips of origin on the Mahon, R. Diogo, 2020.
left side (Palatty et al. 2018).
Synonyms
This muscle may also be referred to as transversus manus
Anomalies (Hepburn 1892). Contrahens refers to a singular muscle, and
Description contrahentes refer to multiple muscles (Cihak 1972).
Alghamdi et al. (2017) observed that the tendon of abductor Typical Presentation
pollicis longus pierced the second tendon of brachioradialis This muscle is only present as a variation or anomaly.
and attached to the palmar fascia. It was also fused with
extensor pollicis brevis. In both a female neonate (Aziz 1979) Comparative Anatomy
and a female fetus with trisomy 18 (Alghamdi et al. 2018), Contrahentes digitorum are sometimes present as adductors
the proximal ends of extensor pollicis brevis and abductor of the fngers in nonhuman apes (Diogo and Wood 2011,
pollicis longus were fused on both sides of the body. The 2012a). These muscles are present in gibbons in about 60%
abductor pollicis longus tendon was also doubled bilaterally of cases, and in most cases, they extend from the contrahens
in the neonate, one attaching to the frst metacarpal and the fascia to the base of the proximal phalanges and extensor
other attaching to the trapezium (Aziz 1979). On the left side expansions on the ulnar side of digit two (contrahens 1), the
of a trisomy 18 cyclopic fetus, Smith et al. (2015) observed radial side of digit 4 (contrahens 2), and the radial side of digit
that abductor pollicis longus was fused with a diminutive fve (contrahens 3) (Deniker 1885; Kohlbrügge 1890–1892;
extensor pollicis brevis. The tendon of this muscle was tri- Hepburn 1892; Lewis 1989; Diogo et al. 2012). Contrahentes
furcated, with a usual insertion onto the frst metacarpal and digitorum are absent in gorillas (Deniker 1885; Macalister
sent two tendons to an abnormal thenar mass of muscles. 1873; Hepburn 1892; Sommer 1907; Pira 1913; Raven 1950;
Abductor pollicis longus sharing a common muscle Preuschoft 1965; Diogo et al. 2010) and rarely present in
belly with extensor pollicis brevis has been observed in orangutans (Diogo et al. 2013b). Contrahentes to digits four
infants with Neu-Laxova syndrome (Shved et al. 1985). and fve are occasionally present in common chimpanzees
Bersu et al. (1976) describe a male infant with Hanhart and bonobos (Hepburn 1892; Miller 1952; Swindler and
syndrome. On the right side of this infant, the wrist and Wood 1973; Lewis 1989; Diogo et al. 2013a, 2017).
FIGURE 3.18 Interossei and contrahentes muscles of the hand in palmar view.
Variations Anomalies
Contrahentes of the hand (contrahentes 3, 4, 5) are typically Dunlap et al. (1986) dissected individuals with trisomy 13,
only present in the early stages of human development and 18, and 21 to determine the common forelimb anomalies
then diffuse and/or fuse with the interossei (Cihak 1972; common to these syndromes. They found that in individu-
Diogo et al 2019). Adductor pollicis develops from con- als with trisomy 18, contrahentes muscles extending to
trahentes 1 and 2 and is present as a differentiated muscle digits two, four, and/or fve were sometimes present. These
at CR20 mm (crown-rump length of 20 mm) (Cihak 1972; muscles originated from the carpal bones, their associated
Diogo et al. 2019). Contrahentes 3, 4, and 5 persist in the ligaments, and from a median raphe extending from the
embryo until CR36 mm (crown-rump length of 36 mm), at ventral shaft of the third metacarpal.
which point they become undistinguishable (Diogo et al.
2019). Contrahentes are very rare in karyotypically nor- Prevalence
mal humans. Remnants of contrahentes digitorum may In their literature review, Smith et al. (2015) found that out
present as more proximal and medial attachments of the of 26 individuals with trisomy 18, contrahens to digit two
oblique head of adductor pollicis (Yamamoto et al. 1988; was present in four cases (15.4%), contrahens to digit four
Tubbs et al. 2005b). When present as distinct muscles, con- was present in one case (3.8%), and contrahens to digit
trahentes originate from the carpals, or the bases of the fve was present in two cases (7.7%).
metacarpals, and can have separate distal insertions into
the metacarpals and phalanges of digits four and fve (Stark Clinical Implications
et al. 1979; Bergman et al. 1988; Tubbs et al. 2005b). Stark The presence of contrahentes has been associated with prob-
et al. (1979) described the presence of contrahentes in three lems including cramping of the hand (Stark et al. 1979) or
children, and in each case, the muscle was present unilater- potentially compression of the median nerve (Tubbs et al.
ally and was associated with an accessory palmaris brevis. 2005b).
Innervation
palMaris brevis (figure 3.19)
Contrahentes digitorum are innervated by the deep branch
of the ulnar nerve (Tubbs et al. 2005b). Synonyms
This muscle is also referred to as palmaris cutaneus, cavo
Prevalence quadrata manus, or Peaucier de la main (Cruveilhier)
N/A (Bergman et al. 2008).
Typical Presentation obliquely from the pisiform to the fexor retinaculum.

Description Palmaris brevis was absent bilaterally in two neonates with
trisomy 18 and three neonates with trisomy 13 (Aziz 1979,
Palmaris brevis extends from the fexor retinaculum and the
1980, 1981). It was also absent bilaterally in a fetus with
medial (ulnar) border of the palmar aponeurosis to the skin
trisomy 18 and cyclopia (Smith et al. 2015). The absence
on the medial border of the palm (Standring 2016).
of palmaris brevis has been observed in infants with Neu-
Innervation Laxova syndrome (Shved et al. 1985).
Palmaris brevis is innervated by the ulnar nerve (Standring Prevalence
2016).
According to Smith et al. (2015), palmaris brevis was absent
Comparative Anatomy in 20 out of 24 individuals with trisomy 13 (83.3%), 15 out of
Palmaris brevis has a similar typical presentation in the 26 individuals with trisomy 18 (57.7%), and in 1 out of 7 indi-
apes, extending from the fexor retinaculum and/or the pisi- viduals with trisomy 21 (14.3%). This muscle had an abnor-
form to the skin on the medial border of the palm (Raven mal presentation in 1 out of 24 individuals with trisomy 13
1950; Miller 1952; Swindler and Wood 1973; Preuschoft (4.2%). In a study of six upper limbs from three infants with
1965; Gibbs 1999; Diogo et al. 2010, 2012, 2013a,b, 2017). Neu-Laxova syndrome, Shved et al. (1985) found that pal-
Occasional absence has been observed in all apes (Bischoff maris brevis was absent in all six upper limbs (100%).
1880; Deniker 1885; Kohlbrügge 1890–1892; Hepburn
1892; Pira 1913; Loth 1931; Kallner 1956; Gibbs 1999; Clinical Implications
Diogo et al. 2010, 2012, 2013a,b, 2017). An accessory palmaris brevis can be mistaken for a soft-
tissue tumor of the hand (Bergman et al. 1988; Tountas
Variations and Bergman 1993). A hypertrophic palmaris brevis or the
Description presence of an accessory palmaris brevis can compress the
Palmaris brevis can be variable in its size, length, and orien- ulnar nerve (Tonkin and Lister 1985; Tountas and Bergman
tation (Macalister 1875; Bergman et al. 1988). It may present 1993; Mackinnon and Dellon 1998; Gonzalez and Netscher
as several fasciculi or one continuous sheet (Macalister 1875). 2016).
Palmaris brevis may have a more radial origin via slips from
the trapezium or scaphoid (Gonzalez and Netscher 2016). luMbricales (luMbricals) (figure 3.19)
Its insertion may extend to the thenar eminence (Macalister
1875). It may join with fexor digiti minimi brevis or attach Synonyms
to the pisiform (Macalister 1875; Mori 1964; Bergman et al. A singular lumbrical muscle may be referred to as lumbri-
1988; Tountas and Bergman 1993; Nayak and Krishnamurthy calis, and multiple lumbrical muscles may be referred to as
2007; Gonzalez and Netscher 2016). It may receive fbers lumbricales (Bergman et al. 1988; Smith et al. 2015).
from the fexor carpi ulnaris tendon (Macalister 1875). This
muscle may present deeper in the hand without an attach- Typical Presentation
ment to the skin (Das and Paul 2006; Gonzalez and Netscher Description
2016). Palmaris brevis may be absent (Macalister 1875; The lumbricals originate from the tendons of fexor digito-
Bergman et al. 1988; Tountas and Bergman 1993; Gonzalez rum profundus and attach to the radial sides of the extensor
and Netscher 2016). Palmaris brevis may also be doubled, expansions of digits two through fve (Standring 2016). The
and the extra muscle can be referred to as palmaris brevis muscles can be either unipennate (single origin) or bipen-
profundus (Bergman et al. 1988; Tountas and Bergman 1993; nate (dual origin), and when bipennate, the two heads of
Gonzalez and Netscher 2016). the muscle arise from two tendons of fexor digitorum pro-
fundus (Standring 2016). The confguration of the lumbri-
Prevalence
cals is variable, but the most common origins are that the
Macalister (1875) found palmaris brevis absent in about 1 frst lumbrical arises from the tendon of fexor digitorum
out of every 45 subjects (2.2%). Mori (1964) did not observe profundus to digit two, the second lumbrical from the ten-
the absence of palmaris brevis in any cases. Mori (1964) don of fexor digitorum profundus to digit three, the third
did fnd that palmaris brevis was well-developed in 10% of lumbrical from the tendons of fexor digitorum profundus
cases, not well-developed in 32% of cases, and had a slip to to digits three and four, and the fourth lumbrical from the
the pisiform bone in 16.2% of cases. tendons of fexor digitorum profundus to digits four and fve
(Eladoumikdachi et al. 2002b; Standring 2016; Gonzalez
Anomalies
and Netscher 2016).
Description
In the literature review conducted by Smith et al. (2015), Innervation
they noted that in one individual with trisomy 13, palmaris The frst and second lumbricals are innervated by the median
brevis ran deep to the ulnar artery and nerve and extended nerve, and the third and fourth lumbricals are innervated
by the ulnar nerve (Standring 2016). Occasionally, the frst 1875; Bergman et al. 1988; Tountas and Bergman 1993;
and second lumbricals are supplied by the ulnar nerve, and Gonzalez and Netscher 2016).
the third lumbrical may be supplied by the median nerve Silawal et al. (2018) describe an unusual case where the
(Standring 2016). frst lumbrical had two muscle bellies connected by a long
(16.2 cm) tendon. The frst belly was located in the proximal
Comparative Anatomy forearm and originated near the medial epicondyle of the
Lumbricals have a similar typical presentation in the apes humerus with the tendon of fexor digitorum superfcialis,
and can have similarly variable origins and insertions (Gibbs and the second belly presented as a typical frst lumbrical
1999; Diogo et al. 2010, 2012, 2013a,b, 2017). The fourth muscle in the hand (Silawal et al. 2018). The tendon con-
lumbrical (to digit fve) in gibbons, gorillas, and bonobos necting the bellies ran through the carpal tunnel (Silawal
can be reduced or absent (Raven 1950; Gibbs 1999; Diogo et al. 2018).
et al. 2010, 2012, 2017).
Prevalence
Variations Macalister (1875) found that the lumbricals were variable in
50 out of 400 subjects (12.5%). In a sample of 34 subjects,
Description
Knott (1883a) found that the frst lumbrical was absent in
Variations on the typical origins and insertions are fre- two cases (5.9%), and both the frst and second lumbrical
quent (Bergman et al. 1988; Standring 2016). The frst or were absent in one case (2.9%). The third lumbrical bifur-
second lumbrical may be bipennate, and the third or fourth cates distally and inserts into two digits in 40% of individu-
lumbrical may be unipennate (Wood 1866; Macalister als (Bergman et al. 1988).
1875; Bergman et al. 1988; Tountas and Bergman 1993; Mori (1964) found that the frst lumbrical had a typical
Eladoumikdachi et al. 2002b; Gonzalez and Netscher 2016; origin in 100% of cases. The second lumbrical had a typi-
Standring 2016). Nation et al. (2019) report a tripinnate frst cal origin in 78% of cases and was bipennate with origins
lumbrical in a male cadaver. The frst lumbrical may have on digits two and three in 22% of cases (Mori 1964). The
a second origin from the tendon of fexor pollicis longus third lumbrical had a typical origin in 94% of cases and was
(Macalister 1875; Tountas and Bergman 1993; Gonzalez unipennate with an origin from digit three only in 6% of
and Netscher 2016; Standring 2016). The third and fourth cases (Mori 1964). The fourth lumbrical had a typical ori-
lumbricals appear to vary more in origin than the frst two gin in 98% of cases and was unipennate with an origin from
(Eladoumikdachi et al. 2002b; Gonzalez and Netscher digit four only in 2% of cases. Mori (1964) also found that
2016; but see description of studies from Mori 1964, below). the tendons of the frst two lumbricals were attached along
The lumbricals may also attach to the proximal phalanges the side of the proximal phalanx to the radial border of the
of the target digits (Mori 1964; Bergman et al. 1988; Doyle extensor digitorum tendon for digits two and three in 10%
and Botte 2003; Gonzalez and Netscher 2016). of cases. The third lumbrical bifurcated into two slips, with
In some cases, only two or three muscles may be pres- one inserting onto the ulnar surface of extensor digitorum
ent (Macalister 1875; Knott 1883a; Bergman et al. 1988). In for digit four in 20% of cases (Mori 1964). The tendon for
rare cases, all four muscles may be absent, or there could the third lumbrical inserted onto the radial surface of exten-
be additional lumbricals due to the presence of accessory sor digitorum for digit four in 80% of cases (Mori 1964).
slips (Macalister 1875; Bergman et al. 1988; Tountas and The fourth lumbrical bifurcated into two slips that inserted
Bergman 1993; Gonzalez and Netscher 2016). Sometimes, onto the ulnar surface of extensor digitorum for digits four
the lumbricals can bifurcate distally and insert into two and fve in 24% of cases (Mori 1964). The fourth lumbri-
digits (Wood 1866, 1868; Bergman et al. 1988; Tountas cal inserted onto the ulnar surface of extensor digitorum
and Bergman 1993; Doyle and Botte 2003; Gonzalez and for digit four in 4% of cases and onto the radial surface of
Netscher 2016). In the region of the frst lumbrical, an extensor digitorum for digit fve in 96% of cases.
accessory slip may join with the tendons of fexor pollicis Based on dissections of 80 hands from 40 individuals,
longus or fexor digitorum superfcialis, opponens polli- Perkins and Hast (1993) found that 40 hands (50%) did not
cis, the muscle bellies of fexor digitorum superfcialis or have the typical presentation of lumbricals. Twenty-seven
fexor digitorum profundus, the frst metacarpal, or the (34%) third lumbricals and four (5%) fourth lumbricals
palmar carpal ligament (Macalister 1875; Bergman et al. bifurcated upon insertion (Perkins and Hast 1993). Four
1988; Tountas and Bergman 1993; Gonzalez and Netscher third lumbricals (5%) had a single insertion onto the third
2016; Standring 2016). The origins of the lumbrical muscles digit, and four fourth lumbricals (5%) had a single insertion
may be displaced proximally and can arise from the fexor onto the fourth digit.
pollicis longus tendon (frst lumbrical), from the fexor
retinaculum, or an accessory tendon from fexor digitorum
superfcialis or fexor digitorum profundus (Macalister 1875; Anomalies
Bergman et al. 1988; Tountas and Bergman 1993; Gonzalez Description
and Netscher 2016). The fourth lumbrical may replace the Camptodactyly, a congenital fexion deformity typically
fourth tendon of fexor digitorum superfcialis (Macalister of the ffth digit, that can be present alone or as part of a
suite of malformations (including trisomies), may be caused abnormal fusions) in 3 out of 26 individuals with trisomy
by variations in lumbrical size, origin, and/or insertions 18 (11.5%). The fourth lumbrical was variable (variable
(Eladoumikdachi et al. 2002b; Gonzalez and Netscher insertions) in 1 out of 26 individuals with trisomy 18 (3.8%).
2016; Favril et al. 2019). Overall, the lumbricals were variable (e.g., variable origins
In a fetus with craniorachischisis, Alghamdi et al. (2017) and insertions, dual insertions, accessory slips) in 5 out of
observed that, on the right side of the body, fexor digito- 24 individuals with trisomy 13 (20.8%), 9 out of 26 indi-
rum profundus only sent tendons to digits four and fve, viduals with trisomy 13 (34.6%), and 1 out of 7 individuals
and thus only two lumbricals were present. On the left side, with trisomy 21 (14.3%) (Smith et al. 2015). These authors
the lumbricals had a normal presentation. On both sides of also found that the frst lumbrical was absent in 3 out of 24
a trisomy 18 cyclopic fetus, Smith et al. (2015) observed individuals with trisomy 13 (12.5%) and 4 out of 26 indi-
that the frst lumbrical was absent, and the tendon of fexor viduals with trisomy 18 (15.4%). The second lumbrical was
digitorum profundus to digit fve did not contribute to the absent in 2 out of 26 individuals with trisomy 18 (7.7%). The
lumbrical attaching to the ffth digit. In the female fetus third lumbrical was absent in 2 out of 26 individuals with
with trisomy 18, all four lumbricals were underdeveloped trisomy 18 (7.7%). The fourth lumbrical was absent in 1 out
in both hands (Alghamdi et al. 2018). On the right side of a of 24 individuals with trisomy 13 (4.2%) and 2 out of 26
child with trisomy 21, the lumbrical muscles were doubled individuals with trisomy 18 (7.7%).
between the third and fourth digits, and between the fourth McFarlane et al. (1992) found that the fourth lumbrical
and ffth digits (Bersu 1980). muscle had a variant insertion onto digit fve in 74 patients
Mieden (1982) describes an infant with median cleft lip, (100%) undergoing surgery for camptodactyly. In a study of
hypotelorism, and alobar holoprosencephaly. The fourth six upper limbs from three infants with Neu-Laxova syn-
lumbrical muscle was absent on the left side (Mieden drome, Shved et al. (1985) found that the frst, second, and
1982). The author does not specify if it was a hand or foot fourth lumbricals had anomalous insertions in two upper
lumbrical. Absence, doubling, and anomalous insertions limbs (33%), the fourth lumbrical was absent in two upper
of one or more of the lumbricals have been observed in limbs (33%), and the fourth lumbrical was doubled in two
infants with Neu-Laxova syndrome (Shved et al. 1985). upper limbs (33%).
On the right hand of a male neonate with Meckel syn-
drome, Pettersen (1984) observed that the fourth lumbri- Clinical Implications
cal attached to the ulnar aspect of the fourth digit, and this Variations in the origins and insertions of lumbrical mus-
muscle also had an attachment to the sixth digit. On the cles may compress the median nerve and cause carpal tun-
left hand, the third lumbrical attached to the ulnar side of nel syndrome (Tountas and Bergman 1993; Gonzalez and
the fourth digit. Netscher 2016).
Prevalence aDDuctOr pOllicis (figure 3.19)

In their literature review, Smith et al. (2015) found that 5
out of 26 individuals with trisomy 18 had an extra lumbri- Synonyms
cal between digits one and two (19.2%). They also found Bergman et al. (1988) refer to this muscle as adductor
that the frst lumbrical was variant (e.g., variable origin, pollicis brevis.
FIGURE 3.19 Intrinsic muscles of the hand and palmaris brevis in palmar view.
Typical Presentation 17 specimens (85%), the second metacarpal in all 20 speci-

Description mens (100%), the third metacarpal in 15 specimens (75%),
the fourth metacarpal in 17 specimens (85%), the hamate in
The oblique head of adductor pollicis originates from the
one specimen (5%), and from the trapezium, trapezoid, and
capitate and the bases of the second and third metacarpals.
capitate in all 20 specimens (100%). The transverse head
The transverse head originates from the third metacarpal
had origins from the third metacarpal in all 20 specimens
(Standring 2016). Both heads end in tendons that converge
(100%) with an additional origin from the second metacar-
to attach together to the base of the proximal phalanx of the
pal in one specimen (5%).
frst digit (Standring 2016).
Anomalies
Innervation
Adductor pollicis is innervated by the deep branch of the Description
ulnar nerve (Standring 2016). On the left side of a fetus with craniorachischisis, Alghamdi
et al. (2017) observed that the transverse head of adductor
Comparative Anatomy pollicis originated from metacarpals three and four, and the
Adductor pollicis has a similar typical presentation in the oblique head originated from the bases of metacarpals three
apes with similar variation in origins and insertions (Gibbs and four and from the capitate. Both had a normal insertion.
1999; Diogo et al. 2010, 2012, 2013a,b, 2017). Its insertion On the left side of a fetus with trisomy 18 and cyclopia, the
may extend to the distal phalanx of the frst digit (Owen oblique head of adductor pollicis was separated into distinct
1868; Macalister 1873; Raven 1950; Preuschoft 1965; Diogo bundles (Smith et al. 2015). Lack of separation between the
et al. 2010, 2012, 2013a,b, 2017). There may be an extensive heads of adductor pollicis and the absence of the oblique
attachment to the frst metacarpal in gibbons, and partial head have been observed in infants with Neu-Laxova syn-
attachments to this bone in the other apes (Gratiolet and drome (Shved et al. 1985).
Alix 1866; Michaelis 1903; Raven 1950; Kallner 1956;
Preuschoft 1965; Diogo et al. 2010, 2012, 2013a,b, 2017). Prevalence
In 2 out of 24 individuals with trisomy 13, the transverse
Variations head of adductor pollicis was hypoplastic and originated
Description from the fourth metacarpal (8.3%). The transverse head of
The sizes and extent of the connection between the oblique adductor pollicis was absent in 1 out of 24 individuals with
head and the transverse head are variable (Mori 1964; trisomy 13 (4.2%) and in 2 out of 26 individuals with tri-
Bergman et al. 1988; Doyle and Botte 2003; Gonzalez and somy 18 (7.7%). In a study of six upper limbs from three
Netscher 2016; Standring 2016). The oblique head may infants with Neu-Laxova syndrome, Shved et al. (1985)
have connections to the frst metacarpal, fourth metacar- found absence of the oblique head in two upper limbs (33%)
pal, trapezium, trapezoid, capitate, and hamate (Chang and lack of separation between the heads in three upper
and Blair 1985). The transverse head may have an origin limbs (50%).
from the second metacarpal or the fourth metacarpal (Mori Clinical Implications
1964; Chang and Blair 1985; Gonzalez and Netscher 2016).
N/A
Adductor pollicis may be divided into multiple bellies or
fascicles (Knott 1883a; Tountas and Bergman 1993; Jan and
Rooze 1994; Gonzalez and Netscher 2016). An accessory interOssei palMaris (palMar interOssei) (figure 3.18)
head may originate near the oblique head from the base of
the second metacarpal and insert onto the dorsal aponeuro- Synonyms
sis of the frst digit (Doyle and Botte 2003; Gonzalez and The frst palmar interosseus muscle is also referred to as
Netscher 2016). Priyadharshini et al. (2019) report a case in interosseous volaris primus of Henle or adductor pollicis
which adductor pollicis and opponens pollicis lacked feshy accessorius (Macalister 1875; Bello-Hellegouarch et al.
muscle fbers and were replaced with fbrous tissue. 2013; Smith et al. 2015).
Prevalence Typical Presentation

Mori (1964) found that the transverse head of adductor pol- Description
licis originated from the third and fourth metacarpal bones There are four palmar interosseous muscles, though many
in 76% of cases and from the third metacarpal bone only textbooks and anatomy resources often do not consider the
in 24% of cases. The two heads were completely separated frst palmar interosseous muscle to be part of the typical pre-
in 82% of cases and partially separated in 16% of cases, sentation of this muscle group (e.g., Gonzalez and Netscher
and the transverse head was divided into two layers in 2% 2016) (Bello-Hellegouarch et al. 2013). The frst originates
of cases (Mori 1964). Based on dissections of 20 cadaveric from the ulnar side of the frst metacarpal and inserts onto
hands, Chang and Blair (1985) found that the oblique head the ulnar sesamoid bone and proximal phalanx of the
of adductor pollicis had origins from the frst metacarpal in frst digit and eventually into the dorsal digital expansion
(Standring 2016). The second palmar interosseous muscle Bello-Hellegouarch et al. (2013) review literature on the
originates from the ulnar side of the second metacarpal and prevalence of the frst palmar interosseous muscle and list a
inserts into the dorsal digital expansion of the second digit range of prevalence from 50% to 100% based on seven stud-
(Standring 2016). The third palmar interosseous muscle ies. Out of the 72 hands dissected by Bello-Hellegouarch
originates from the radial side of the fourth metacarpal and et al. (2013), the frst palmar interosseous muscle was
inserts into the dorsal digital expansion of the fourth digit present in 67 hands (93%). The origin and insertion of the
with the third lumbrical (Standring 2016). The fourth pal- muscle were observed in 66 hands. It originated from the
mar interosseous muscle originates from the radial side of frst metacarpal only in 25 hands (37.9%); from the frst
the ffth metacarpal and inserts into the proximal phalanx metacarpal and a common origin with the dorsal interosse-
of the ffth digit and into the dorsal digital expansion with ous in seven hands (10.6%); from the wrist bones/ligaments
the fourth lumbrical (Standring 2016). only in seven hands (10.6%); from the frst metacarpal and
the wrist bones/ligaments in seven hands (10.6%); from a
Innervation thin membrane shared with the dorsal interosseous in four
The palmar interossei are innervated by the deep branch of hands (6.1%); from the dorsal interosseous in four hands
the ulnar nerve (Standring 2016). (6.1%); from the frst metacarpal and a common origin
with the oblique head of adductor pollicis in four hands
Comparative Anatomy (6.1%); from the frst metacarpal and the thin membrane
In all apes except common chimpanzees, the palmar inter- shared with the dorsal interosseous in three hands (4.5%);
ossei have a similar confguration as those in humans from the frst and second metacarpals in two hands (3%);
(Diogo et al. 2010, 2012, 2013a,b, 2017). “Adductor pollicis from the frst metacarpal, wrist bones/ligaments, and the
accessorius” is occasionally present in gorillas, common thin membrane shared with the dorsal interosseous in one
chimpanzees, and bonobos extending from the ulnar side hand (1.5%); from the wrist bones/ligaments and the thin
of the base of the frst metacarpal to the base of the frst membrane shared with the dorsal interosseous in one hand
proximal phalanx (Gratiolet and Alix 1966; Champneys (1.5%); and from the frst and second metacarpals and the
1872; Diogo et al. 2012, 2013a, 2017). thin membrane in one hand (1.5%) (Bello-Hellegouarch
et al. 2013).
Variations This frst palmar interosseous inserted onto the proxi-
Description mal phalanx of the frst digit only in 34 hands (51.5%), the
One or more of the interosseous muscles may be doubled proximal phalanx of the frst digit and a common inser-
or absent (Macalister 1875; Kaplan 1965; Bergman et al. tion with the oblique head of adductor pollicis in 18 hands
1988; Perkins and Hast 1993; Doyle and Botte 2003; Bello- (27.3%), the frst metacarpal and the proximal phalanx
Hellegouarch et al. 2013; Gonzalez and Netscher 2016; of the frst digit in fve hands (7.6%), the frst metacarpal
Standring 2016). The palmar interossei may also have one only in three hands (4.5%), the frst metacarpal and a com-
or more bellies (Eladoumikdachi et al. 2002a, b; Bello- mon insertion with the oblique head of adductor pollicis in
Hellegouarch et al. 2013; Gonzalez and Netscher 2016). three hands (4.5%), fusion and common insertion with the
The frst palmar interosseous muscle is often diminutive oblique head of adductor pollicis in two hands (3%), and
(Standring 2016) and is sometimes considered to be a deep onto the frst metacarpal, proximal phalanx of digit one,
head of fexor pollicis brevis (Kaplan 1965). It may receive and a common insertion with the oblique head of adduc-
a slip from extensor carpi radialis longus (Macalister 1875). tor pollicis in one hand (1.5%) (Bello-Hellegouarch et al.
Eladoumikdachi et al. (2002a) state that the frst three 2013). Bello-Hellegouarch et al. (2013) also note that out
palmar interossei are often variable in their origins and of 66 hands, the muscle presented as a single muscular
insertions. In their literature review on the frst palmar bundle in 38 hands (57.6%), two muscular bundles in ten
interosseous muscle, Bello-Hellegouarch et al. (2013) note hands (15.2%), more than two muscular bundles in ten
that this muscle can originate from the capitate, trapezoid, hands (15.2%), a single thin tendon with no muscle fbers
trapezium, frst, metacarpal, second metacarpal, and/or in fve hands (7.6%), and both a muscular and tendinous
third metacarpal, and have insertions onto the ulnar sesa- bundle in three hands (4.5%).
moid, proximal phalanx of digit one, and/or extensor expan-
sion. The second and third palmar interosseous muscle may
Anomalies
receive fbers from the third metacarpal (Eladoumikdachi
et al. 2002a; Gonzalez and Netscher 2016). Description
On the right side of a fetus with craniorachischisis,
Prevalence Alghamdi et al. (2017) observed that there were only
According to Macalister (1875), Wood found the frst pal- two palmar interossei, which attached to the radial sides
mar interosseous muscle in 3 out of 36 cases (8.3%). Based of digits four and fve. On both sides of a fetus with tri-
on dissections of 80 hands from 40 individuals, Perkins and somy 18 and cyclopia, “adductor pollicis accessorius” was
Hast (1993) found that the frst palmar interosseous muscle present and arose from the proximal portion of the frst
was present in 68 hands (85%) and 36 (90%) individuals. metacarpal and inserted onto the base of the frst proximal
phalanx (Smith et al. 2015). Its insertion was blended with median nerves (Kaplan 1965; Bergman et al. 1988; Tountas
the insertion of adductor pollicis on both sides (Smith and Bergman 1993; Doyle and Botte 2003; Gonzalez and
et al. 2015). Netscher 2016).
In individuals with camptodactyly, the fourth pal-
mar interosseous muscle can insert onto the fourth digit Comparative Anatomy
(Tountas and Bergman 1993). Anomalous insertion of the In all apes except common chimpanzees, the fexores breves
frst palmar interosseous muscle has been observed in profundi 3, 5, 6, and 8 are fused with the intermetacarpales
infants with Neu-Laxova syndrome (Shved et al. 1985). In 1, 2, 3, and 4, forming the interossei dorsales 1, 2, 3, and 4,
a male neonate with Meckel syndrome, Pettersen (1984) respectively (Diogo et al. 2010, 2012, 2013a,b, 2017). As in
found that on the right hand, the third palmar interosseous humans, the dorsal interossei in the apes extend from the
muscle attached to the sixth digit. sides of adjacent metacarpals to insert onto the proximal
phalanges and extensor expansions of digits two, three, and
Prevalence four (Diogo et al. 2010, 2012, 2013a,b, 2017). Accessory
In their literature review, Smith et al. (2015) found that the interosseus muscles to digit two are common in gibbons
fourth palmar interosseous muscle originates from oppo- (Susman et al. 1982; Diogo et al. 2012).
nens digiti minimi in 1 out of 24 individuals with trisomy
13 (4.2%). They also found that the frst palmar and dorsal Variations
interossei were absent in 2 out of 26 individuals with tri- Description
somy 18 (7.7%). “Adductor pollicis accessorius” was present
One or more of the dorsal interossei may be divided,
in 5 out of 24 individuals with trisomy 13 (20.8%), 10 out
doubled, reduced, or absent (Wood 1868; Macalister
of 26 individuals with trisomy 18 (38.5%), and two out of
1875; Kaplan 1965; Bergman et al. 1988; Doyle and Botte
seven individuals with trisomy 21 (28.6%). In a study of
2003; Gonzalez and Netscher 2016). They may have addi-
six upper limbs from three infants with Neu-Laxova syn-
tional heads, with the second and third having up to three
drome, Shved et al. (1985) found that frst palmar interosse-
heads (Macalister 1875; Bergman et al. 1988; Tountas and
ous muscle had an anomalous insertion in two upper limbs
Bergman 1993; Eladoumikdachi et al. 2002a; Bharambe
(33%).
et al. 2013; Nayak et al. 2016). The frst dorsal interosse-
ous muscle may receive a slip from extensor carpi radialis
longus, and the third may receive a slip from extensor carpi
N/A radialis brevis (Macalister 1875). Slips may pass to exten-
sor digitorum brevis manus (Macalister 1875). The dorsal
interOssei DOrsalis (DOrsal interOssei) (figure 3.18) interossei may also be associated with accessory muscles
(Nayak et al. 2016).
Synonyms
The frst dorsal interosseous muscle may also be referred to Prevalence
as abductor indicis (Standring 2016). Macalister (1875) notes that a split frst dorsal interosseous
muscle, with one part attaching to the thumb and the other
Typical Presentation to the index fnger, occurs in about 1 out of 120 subjects
Description (0.8%). In dissections of 25 hands, Bharambe et al. (2013)
There are four dorsal interosseous muscles (Standring found an additional head of the second dorsal interosseous
2016). The frst originates from sides of the frst and second muscle in three upper limbs (12%) and additional heads of
metacarpals and attaches to the proximal phalanx and the the second, third, and fourth dorsal interossei in one upper
metacarpophalangeal joint capsule of digit two (Standring limb (4%). In dissections of 30 hands, Nayak et al. (2016)
2016). The second originates from the sides of metacarpals found supernumerary muscles that originated from the
two and three and attaches onto the radial side of the third third metacarpal in three hands (10%), and the third dorsal
digit into the proximal phalanx and dorsal digital expan- interosseous muscle had three heads in one hand (3%).
sion (Standring 2016). The third originates from the sides of
metacarpals three and four and attaches to the ulnar side of Anomalies
the third digit, into the dorsal digital expansion (Standring Description
2016). The fourth originates from the sides of metacarpals On the right side of a fetus with craniorachischisis,
four and fve and attaches to the proximal phalanx and digi- Alghamdi et al. (2017) observed that there were only three
tal expansion of digit four (Standring 2016). dorsal interossei.
The dorsal interossei are innervated by the deep branch of In their literature review, Smith et al. (2015) found that the
the ulnar nerve (Standring 2016). Innervation may vary, with frst palmar and dorsal interossei were absent in 2 out of 26
potential innervation by the radial, musculocutaneous, or individuals with trisomy 18 (7.7%).
Clinical Implications pollicis. The deep head in many cases showed additional
Bharambe et al. (2013) and Nayak et al. (2016) suggest origins from the base of the second, third, and/or fourth
that supernumerary muscles or additional heads of the metacarpals, or from a fbrous extension of the carpal
dorsal interossei may be largely asymptomatic but could tunnel wall (Dunlap et al. 2017). This muscle, often the
potentially lead to chronic compartment syndrome due to superfcial head, can be partially or fully blended with
increased intracompartmental pressure. opponens pollicis, abductor pollicis brevis, or adductor
pollicis (Macalister 1875; Bergman et al. 1988; Tountas
and Bergman 1993; Doyle and Botte 2003; Gonzalez and
flexOr pOllicis brevis (figure 3.19) Netscher 2016; Standring 2016).
Synonyms
The deep head of this muscle can be referred to as the deep Prevalence
head of Cruveilhier (Dunlap et al. 2017). Caetano et al. (2017) studied this muscle in 60 hands and
found that the superfcial head was supplied by the median
Typical Presentation nerve in 42 hands (70%) and was supplied by both the
Description median and ulnar nerves in 18 hands (30%). The deep head
The superfcial head of fexor pollicis brevis originates was absent in nine hands (15%) and of the remaining 51
from the fexor retinaculum and trapezium and inserts via hands, one hand was innervated by the median nerve only
a tendon onto the radial side of the base of the proximal (1.96%), 10 were supplied by the ulnar nerve only (19.6%),
phalanx of the frst digit (Standring 2016). The deep head and 40 had dual innervation (78.4%) (Caetano et al. 2017).
originates from the trapezoid and capitate and joins with Dunlap et al. (2017) studied the innervation of both heads
the superfcial head upon its insertion into the base of the in 11 hands and found that the superfcial head was inner-
proximal phalanx of the frst digit (Standring 2016). vated by the median nerve in all (100%) hands while the
deep head was innervated by the median nerve only in
Innervation three hands (27.3%) and both the ulnar and median nerves
The superfcial head is typically innervated by the median in eight hands (72.7%).
nerve but may also be innervated by the ulnar nerve, or both In dissections of 80 hands, Dunlap et al. (2017) note that
nerves (Bergman et al. 1988; Tountas and Bergman 1993; all superfcial heads (100%) originated from the fexor reti-
Gonzalez and Netscher 2016; Standring 2016; Caetano naculum and the trapezium and inserted onto the proximal
et al. 2017). The deep head is primarily innervated by the phalanx of the frst digit. Additional insertions were present
ulnar nerve but is sometimes innervated by the median in 36 hands (45%) and include attachments to the shaft of
nerve or by both nerves (Bergman et al. 1988; Tountas and the frst metacarpal in 12 hands (15%), with the opponens
Bergman 1993; Gonzalez and Netscher 2016; Standring pollicis in nine hands (11.3%), or both of these insertions
2016; Caetano et al. 2017; Dunlap et al. 2017). in 15 hands (18.8%). The deep heads of the muscle origi-
nated from the trapezoid, capitate, and ligamentum carpi
Comparative Anatomy radiatum in all but one hand (1.3%), which lacked an ori-
Flexor pollicis brevis has a similar typical presentation in gin from the trapezoid. Additional origins of the deep head
the apes, and the deep head (corresponding to fexor bre- include from the base of the second metacarpal in one hand
vis profundus 2 of lower mammals) is occasionally absent (1.3%), the base of the third metacarpal in 20 hands (25%),
(Gibbs 1999; Diogo et al. 2010, 2012, 2013a,b, 2017). This the bases of both the third and fourth metacarpals in four
muscle may also be partially or fully fused with neighbor- hands (5%), or from a fbrous extension of the carpal tunnel
ing muscles, including opponens pollicis or abductor pol- wall in 43 hands (53.8%). The deep head had two heads in
licis brevis (Gibbs 1999; Diogo et al. 2010, 2012, 2013a,b, 37 hands (46.3%).
2017).
Anomalies
Variations Description
Description On the left side of a fetus with craniorachischisis, the deep
The entire fexor pollicis brevis muscle, or sometimes head had an uncommon insertion onto the distal end of the
just the deep head, can be absent or doubled (Macalister proximal phalanx of digit one, while the superfcial head
1875; Kaplan 1965; Tountas and Bergman 1993; Bergman inserted abnormally onto the proximal end of the distal
et al. 1988; Gonzalez and Netscher 2016; Standring 2016; phalanx of digit one (Alghamdi et al. 2017). In a fetus with
Caetano et al. 2017). The superfcial or deep heads may be trisomy 18 and cyclopia, fexor pollicis brevis was absent
divided, or there can be an additional third head (Bergman on the right side (Smith et al. 2015). On the left side of
et al. 1988; Doyle and Botte 2003; Dunlap et al. 2017). this specimen, fexor pollicis brevis was fused with abduc-
Dunlap et al. (2017) note an origin of the superfcial head tor pollicis brevis and opponens pollicis. The absence of
from the trapezoid in one case, and additional insertions fexor pollicis brevis has been observed in infants with Neu-
onto the shaft of the frst metacarpal or with opponens Laxova syndrome (Shved et al. 1985).
Prevalence pollicis lacked feshy muscle fbers and were replaced with
In their literature review, Smith et al. (2015) found that fbrous tissue.
fexor pollicis brevis was variant (e.g., fused with opponens
pollicis, diminutive muscle, variable origin) in 2 out of 24 Prevalence
individuals with trisomy 13 (8.3%) and in 10 out of 26 indi- N/A
viduals with trisomy 18 (38.5%). These authors also noted
that fexor pollicis brevis was absent in 1 out of 24 indi- Anomalies
viduals with trisomy 13 (4.2%) and in 8 out of 26 individu- Description
als with trisomy 18 (30.8%). In a study of six upper limbs On the left side of a fetus with craniorachischisis, oppo-
from three infants with Neu-Laxova syndrome, Shved et al. nens pollicis abnormally originated from the second tendon
(1985) found that fexor pollicis brevis was absent in all six of brachioradialis at the base of the frst metacarpal and
upper limbs (100%). inserted onto the proximal end of the distal phalanx of digit
one (Alghamdi et al. 2017). In a fetus with trisomy 18 and
Clinical Implications cyclopia, opponens pollicis was absent on the right side and
N/A fused with abductor pollicis brevis and fexor pollicis brevis
on the left side (Smith et al. 2015). The absence of opponens
pollicis has been observed in infants with Neu-Laxova syn-
OppOnens pOllicis (figure 3.19)
drome (Shved et al. 1985).
Synonyms
Prevalence
N/A
Typical Presentation opponens pollicis was diminutive in 1 out of 24 individu-
Description als with trisomy 13 (4.2%) and in 3 out of 26 individuals
with trisomy 18 (11.5%). Opponens pollicis was absent in
Opponens pollicis originates from the trapezium and fexor
1 out of 24 individuals with trisomy 13 (4.2%) and in 6 out
retinaculum and inserts onto the lateral (radial) half of the
of 26 individuals with trisomy 18 (23.1%). In a study of six
palmar surface of the frst metacarpal (Standring 2016).
upper limbs from three infants with Neu-Laxova syndrome,
Innervation Shved et al. (1985) found that opponens pollicis was absent
in all six upper limbs (100%).
Opponens pollicis is typically innervated by the median
nerve, and sometimes by the ulnar nerve (Standring 2016).
Comparative Anatomy N/A
Opponens pollicis has a similar typical presentation in the
apes, often with an additional origin from the radial sesa- flexOr Digiti MiniMi brevis (figure 3.19)
moid bone and/or the scaphoid (Brooks 1887; Miller 1952;
Gibbs 1999; Diogo et al. 2010, 2012, 2013a,b, 2017). It may Synonyms
have an origin from the tendon of abductor pollicis brevis This muscle may also be referred to as fexor digiti minimi
in orangutans (Brooks 1887; Primrose 1899, 1900; Diogo (Gonzalez and Netscher 2016), abductor digiti quinti brevis
et al. 2013b). Its insertion may extend to the proximal and/ (Sullivan and Osgood 1927), or fexor brevis minimi digiti
or distal phalanx of the frst digit in gibbons (Kohlbrügge (Macalister 1875).
1890–1892; Hepburn 1892; Diogo et al. 2012).
Description Flexor digiti minimi brevis originates from the hook of
Opponens pollicis may be divided into two parts, doubled, the hamate and the fexor retinaculum (Standring 2016). It
or in some cases, it may be absent entirely (Bergman et al. inserts onto the ulnar side of the base of the proximal pha-
1988; Doyle and Botte 2003; Gonzalez and Netscher 2016). lanx of the ffth digit (Standring 2016).
It may also be divided into multiple bellies or fascicles
(Jan and Rooze 1994). It may be partially or fully fused Innervation
with fexor pollicis brevis (Bergman et al. 1988; Tountas Flexor digiti minimi brevis is innervated by the deep branch
and Bergman 1993; Doyle and Botte 2003; Gonzalez and of the ulnar nerve (Standring 2016).
Netscher 2016; Standring 2016). It may give origin to a third
head of abductor pollicis brevis (Wood 1868; Macalister Comparative Anatomy
1875; Bergman et al. 1988; Tountas and Bergman 1993; Flexor digiti minimi brevis has a similar typical pre-
Gonzalez and Netscher 2016). Priyadharshini et al. (2019) sentation in the apes, extending from the hamate and/or
report a case in which opponens pollicis and adductor fexor retinaculum to the proximal phalanx of digit fve
(Gratiolet and Alix 1866; Macalister 1873; Deniker 1885; Anomalies

Hepburn 1892; Primrose 1899, 1900; Sonntag 1923, 1924; Description
Raven 1950; Miller 1952; Kallner 1956; Swindler and
On the right side of a fetus with craniorachischisis, Alghamdi
Wood 1973; Diogo et al. 2010, 2012, 2013a,b, 2017). It may
et al. (2017) observed that the fexor digiti minimi brevis
have an origin from the pisiform in gibbons and gorillas
potentially originated from the trapezoid and inserted onto
(Kohlbrügge 1890–1892; Preuschoft 1965; Diogo et al.
the proximal phalanx of the fourth digit. Flexor digiti min-
2010, 2012). It was completely fused with abductor digiti
imi brevis was normal on the left side of this specimen.
minimi in one bonobo (Diogo et al. 2017).
Flexor digiti minimi brevis has been observed to have an
Variations anomalous insertion in infants with Neu-Laxova syndrome
(Shved et al. 1985).
Description
Flexor digiti minimi brevis may be doubled, reduced, or Prevalence
absent (Macalister 1875; Le Double 1897; Bergman et al. In a study of six upper limbs from three infants with Neu-
1988; Doyle and Botte 2003; Wingerter et al. 2003; Murata Laxova syndrome, Shved et al. (1985) found that fexor
et al. 2004; Gonzalez and Netscher 2016; Standring 2016, digiti minimi brevis had an anomalous insertion in three
May 2020). It may be fused with or replaced by adjacent upper limbs (50%).
muscles, including abductor digiti minimi or opponens
digiti minimi (Macalister 1875; Le Double 1897; Bergman Clinical Implications
et al. 1988; Tountas and Bergman 1993; Gonzalez and Variant origins, insertions, or accessory slips associated
Netscher 2016; Standring 2016). It could also be replaced by with fexor digiti minimi brevis can lead to ulnar nerve
a tendon that originates from fexor carpi ulnaris (Bergman compression (Spinner et al. 1996; Madhavi and Holla 2003;
et al. 1988; Doyle and Botte 2003; Gonzalez and Netscher Wingerter et al. 2003; Georgiev and Jelev 2007) or poten-
2016). Flexor digiti minimi brevis may have an origin from tially ulnar artery thrombosis (Pribyl and Moneim 1994).
the distal antebrachial fascia (Wingerter et al. 2003). It
may also originate from the tendon of fexor carpi radialis
(Georgiev and Jelev 2011). Georgiev and Jelev (2007) note OppOnens Digiti MiniMi (figure 3.19)
a case with a triple origin from the hook of the hamate, Synonyms
the fexor retinaculum, and from the fbers forming Guyon’s
This muscle may also be referred to as opponens digiti
canal. Flexor digiti minimi brevis may be connected via
quinti (Sullivan and Osgood 1927) or opponens minimi
accessory slips to the shaft or distal end of the ffth meta-
digiti (Macalister 1875).
carpal, the hook of the hamate, the palmaris longus ten-
don, or fexor carpi radialis (Macalister 1875; Kaplan 1965; Typical Presentation
Tountas and Bergman 1993; Pribyl and Moneim 1994;
Description
Spinner et al. 1996; De Smet 2002; Gonzalez and Netscher
2016; Standring 2016). Opponens digiti minimi originates from the hook of the
Madhavi and Holla (2003) note a case in which the mus- hamate and the fexor retinaculum and inserts along the
cle was shifted proximally with an origin from the super- ulnar border and palmar surface of the ffth metacarpal
fcial transverse septum in the distal forearm, extended (Standring 2016). It is frequently divided into two layers
through Guyon’s canal above the ulnar nerve and vessels, by the deep branches of the ulnar artery and ulnar nerve
fused with abductor digiti minimi, and had a typical inser- (Doyle and Botte 2003; Standring 2016).
tion onto the proximal phalanx of the ffth digit. Nation
Innervation
et al. (2019) describe a muscle with a similar presentation
and term it “fexor digiti minimi longus.” Flexor digiti min- Opponens digiti minimi is innervated by the deep branch of
imi brevis is also associated with tensor capsularis articula- the ulnar nerve (Standring 2016).
tionis metacarpophalangei digiti minimi, a supernumerary
Comparative Anatomy
muscle that originates from the ligaments connecting the
hamate and pisiform and inserts onto the metacarpophalan- Opponens digiti minimi generally has a similar typical
geal joint of digit fve (Bergman et al. 1988). presentation in the apes, except the division into a super-
fcial and deep layer may be poorly differentiated in some
Prevalence specimens, and the origin is often blended with fexor digiti
In a study of 35 hands, Murata et al. (2004) found that fexor minimi brevis (Lewis 1989; Gibbs 1999; Diogo et al. 2010,
digiti minimi brevis was absent in eight hands (22.9%), had 2012, 2013a,b, 2017).
one belly in 24 hands (68.6%), and had two bellies in three
hands (8.6%). In a study of 38 individuals, May (2020) Variations
found that this muscle had one belly in 22 cases (58%), two Description
bellies in eight cases (21%), and was absent in eight cases Opponens digiti minimi may be absent (Macalister 1875).
(21%). Opponens digiti minimi may be partially blended with the
adjacent muscles, including the fexor digiti minimi bre- attachment may extend to the distal end of the frst meta-
vis or abductor digiti minimi (Macalister 1875; Tountas carpal or distal phalanx of the frst digit in gibbons, goril-
and Bergman 1993; Doyle and Botte 2003; Gonzalez and las, and common chimpanzees (Kohlbrügge 1890–1892;
Netscher 2016; Standring 2016). This muscle may be func- Hepburn 1892; Dwight 1895; Raven 1950; Preuschoft 1965;
tionally related to ulnaris externis brevis, which originates Diogo et al. 2010, 2012, 2013a).
near the distal end of the ulna and inserts onto the fourth
and ffth metacarpals (Bergman et al. 1988). Variations
Description
Prevalence Abductor pollicis brevis can be divided into an outer and
N/A inner part (Bergman et al. 1988) or into multiple fascicles
(Jan and Rooze 1994). The entire muscle may be absent or
Anomalies doubled (Macalister 1875; Bergman et al. 1988; Tountas
Description and Bergman 1993; Doyle and Botte 2003; Gonzalez and
Opponens digiti minimi had a generally normal presenta- Netscher 2016). It may have a third head that originates from
tion on both upper limbs of a fetus with craniorachischisis opponens pollicis (Wood 1868; Macalister 1875; Bergman
dissected by Alghamdi et al. (2017), but the right opponens et al. 1988; Tountas and Bergman 1993). Abductor polli-
digiti minimi had a connection to the pisiform cartilage cis brevis may receive slips from extensor pollicis brevis,
in addition to the hamate. The absence of opponens digiti extensor pollicis longus, opponens pollicis, scaphoid, or
minimi has been observed in infants with Neu-Laxova syn- the styloid process of the radius (Macalister 1875; Bergman
drome (Shved et al. 1985). et al. 1988; Standring 2016; Gonzalez and Netscher 2016).
It can also be connected to abductor pollicis longus, adduc-
Prevalence tor pollicis, extensor carpi radialis longus, fexor pollicis
brevis, palmaris longus, or the skin over the thenar emi-
In a study of six upper limbs from three infants with Neu-
nence (Macalister 1875; Bergman et al. 1988; Gonzalez and
Laxova syndrome, Shved et al. (1985) found that opponens
Netscher 2016).
digiti minimi was absent in three upper limbs (50%).

An accessory opponens digiti minimi can be mistaken for a N/A
soft-tissue tumor of the hand (Tountas and Bergman 1993;
Gonzalez and Netscher 2016). Anomalies
Description
abDuctOr pOllicis brevis (figure 3.19) On the left side of a fetus with craniorachischisis, Alghamdi
et al. (2017) found that abductor pollicis brevis formed a
Synonyms u-shaped structure with the tendons of extensor pollicis bre-
N/A vis and fexor pollicis longus. Abductor pollicis brevis was
absent on the right side of a fetus with trisomy 18 and cyclo-
Typical Presentation pia studied by Smith et al. (2015). On the left side of this
Description specimen, abductor pollicis brevis was blended with some
Abductor pollicis brevis originates primarily from the tendons of abductor pollicis longus. Furthermore, abductor
fexor retinaculum, with additional fbers arising from the pollicis brevis, opponens pollicis, and fexor pollicis brevis
scaphoid, trapezium, and abductor pollicis longus tendon were all fused (Smith et al. 2015). Absence and anomalous
(Standring 2016). It inserts via a tendon onto the radial insertion of abductor pollicis brevis has been observed in
side of the base of the proximal phalanx and dorsal digital infants with Neu-Laxova syndrome (Shved et al. 1985).
expansion of the frst digit (Standring 2016).
Prevalence
Innervation In their literature review, Smith et al. (2015) found that
Abductor pollicis brevis is innervated by the recurrent abductor pollicis brevis was diminutive in 5 out of 26
branch of the median nerve (Standring 2016). individuals with trisomy 18 (19.2%). They also found that
abductor pollicis brevis was absent in 1 out of 24 individ-
Comparative Anatomy uals with trisomy 13 (4.2%) and in 9 out of 26 individu-
Abductor pollicis brevis has a similar typical presentation in als with trisomy 18 (34.6%). In a study of six upper limbs
the apes, arising from the fexor retinaculum and sometimes from three infants with Neu-Laxova syndrome, Shved et al.
the trapezium, adjacent sesamoid bone, and the scaphoid (1985) found absence of abductor pollicis brevis in two
and inserting onto the proximal phalanx of digit one (Gibbs upper limbs (33%) and anomalous insertion in two upper
1999; Diogo et al. 2010, 2012, 2013a, 2017). The muscle limbs (33%).
Clinical Implications 2016; Standring 2016). It may have a connection to the

An accessory abductor pollicis brevis may be mistaken for ffth metacarpal via a slip from the pisiform (Standring
a soft-tissue tumor of the hand or lead to compression of the 2016). Macalister (1875) reports an origin entirely from the
recurrent motor branch of the median nerve (Tountas and pisiform.
Bergman 1993; Gonzalez and Netscher 2016). Bergman et al. (1988), Tountas and Bergman (1993), and
many others have described the presentation of an acces-
sory head of this muscle, termed accessorius ad abductorem
abDuctOr Digiti MiniMi (figure 3.19) digiti minimi manus, otherwise known as an accessory
abductor digiti minimi muscle or abductor digiti minimi
Synonyms accessorius. When present, it can originate from a variety of
This muscle may also be referred to as abductor minimi structures including from the fexor retinaculum, the tendon
digiti (Macalister 1875). of fexor carpi ulnaris, the tendon of palmaris longus, the
tendon of pronator quadratus, the antebrachial fascia, the
radius, the ulna, or from the intermuscular fascia beneath
fexor carpi ulnaris or fexor carpi radialis (Bergman et al.
Description
1988; Tountas and Bergman 1993; Curry and Kuz 2000;
Abductor digiti minimi originates from the pisiform, fexor Claassen et al. 2013; Gonzalez and Netscher 2016). This
carpi ulnaris tendon, and the ligament that connects the accessory head can insert into the main belly of abductor
pisiform and hamate (Standring 2016). It ends in a bifur- digiti minimi or fexor digiti minimi brevis (Bergman et al.
cated tendon that inserts onto the base of the proximal 1988; Gonzalez and Netscher 2016). It can also have inser-
phalanx and the extensor expansion of digit fve (Standring tions into the ffth metacarpal, the proximal phalanx of the
2016). ffth digit, or into the palmar ligament of the metacarpopha-
langeal joint of the ffth digit (Bergman et al. 1988; Sañudo
Innervation
et al. 1993; Curry and Kuz 2000).
Abductor digiti minimi is innervated by the deep branch of Abductor digiti minimi is associated with three other
the ulnar nerve (Standring 2016). supernumerary slips. Pisimetacarpeus may extend from
the pisiform to the ffth metacarpal (Bergman et al. 1988;
Comparative Anatomy
Tountas and Bergman 1993; Gonzalez and Netscher
Abductor digiti minimi has a similar typical presentation 2016). Pisiuncinatus has been termed to describe a muscle
in the apes, arising from the pisiform and sometimes the extending between the pisiform and hamate (Bergman et al.
fexor retinaculum and/or hamate and inserting onto the 1988; Tountas and Bergman 1993; Gonzalez and Netscher
proximal phalanx of digit fve (Kohlbrügge 1890–1892; 2016). Pisiannularis extends from the pisiform to the fexor
Hepburn 1892; Swindler and Wood 1973; Gibbs 1999; retinaculum (Bergman et al. 1988; Tountas and Bergman
Diogo et al. 2010, 2012, 2013a,b, 2017). The insertion 1993; Gonzalez and Netscher 2016).
may be blended with fexor digiti minimi brevis (Hepburn
1892; Gibbs 1999; Diogo et al. 2012). Preuschoft (1965) Prevalence
observed an additional origin from the triquetrum in one Bergman et al. (1988) state that pisiuncinatus has a preva-
gorilla. There may be an additional insertion onto the dis- lence of between 2% and 5%. Based on dissections of 80
tal end of the ffth metacarpal in gibbons (Deniker 1885; hands from 40 individuals, Perkins and Hast (1993) found
Diogo et al. 2012). that abductor digiti minimi had extra slips of origin from
various structures (e.g., belly or tendon of palmaris lon-
Variations
gus, fascia of the forearm, fexor retinaculum) in 8 out of
Description 80 hands (10%). Sañudo et al. (1993) note the presence of
Abductor digiti minimi may be absent (Macalister 1875; accessory bundles of abductor digiti minimi in 2 out of
Bergman et al. 1988; Tountas and Bergman 1993; Gonzalez 62 forearms (3.2%). In a study of 35 hands, Murata et al.
and Netscher 2016). Abductor digiti minimi may be (2004) found that abductor digiti minimi had one belly in
divided into two or three slips (Macalister 1875; Knott six hands (17.1%), two bellies in 28 hands (80%), and three
1883a; Bergman et al. 1988; Murata et al. 2004; Standring bellies in one hand (2.9%). May (2020) found accessory
2016). It may also be fused with fexor digiti minimi bre- muscles associated with abductor digiti minimi in 2 out of
vis (Macalister 1875; Kaplan 1965; Bergman et al. 1988; 76 hands (2.6%).
Tountas and Bergman 1993; Gonzalez and Netscher 2016;
Standring 2016). Additional fbers of abductor digiti min-
imi may originate from the fexor retinaculum, extensor Anomalies
retinaculum, antebrachial fascia, the palmaris longus ten- Description
don, the fexor carpi ulnaris tendon, or the extensor carpi Abductor digiti minimi with a high insertion has been
ulnaris tendon (Wood 1866, 1868; Macalister 1875; Knott observed in infants with Neu-Laxova syndrome (Shved
1883a; Perkins and Hast 1993; Gonzalez and Netscher et al. 1985).
Prevalence 1993; Bergman et al. 1988; Gonzalez and Netscher 2016) or

According to their literature review, Smith et al. (2015) the median nerve (Soldado-Carrera et al. 2000). Accessory
found that abductor digiti minimi had extra slips and ten- heads or muscle bellies of abductor digiti minimi may also
dons in 1 out of 24 individuals with trisomy 13 (4.2%). In compress the ulnar artery or lead to thrombosis or fbrosis
a study of six upper limbs from three infants with Neu- of the ulnar artery (Tountas and Bergman 1993; Soldado-
Laxova syndrome, Shved et al. (1985) found that abductor Carrera et al. 2000; Gonzalez and Netscher 2016). These
digiti minimi had a high insertion in two upper limbs (33%). accessory tissues may also be mistaken for soft tissue
tumors (Soldado-Carrera et al. 2000).
An accessory abductor digiti minimi may compress the deep
palmar branch of the ulnar nerve (Tountas and Bergman
4 Trunk Muscles
Eve K. Boyle
Vondel S. E. Mahon
Rui Diogo
Rowan Sherwood
University of Michigan
CONTENTS
Muscles of the Abdominal Region..................................................................................................................................... 213
Obliquus externus abdominis (External oblique).......................................................................................................... 213
Synonyms ................................................................................................................................................................. 213
Variations.................................................................................................................................................................. 214
Anomalies................................................................................................................................................................. 214
Obliquus abdominis externus profundus....................................................................................................................... 214
Synonyms ................................................................................................................................................................. 214
Variations.................................................................................................................................................................. 214
Anomalies................................................................................................................................................................. 215
Obliquus internus abdominis (Internal oblique) ........................................................................................................... 215
Synonyms ................................................................................................................................................................. 215
Variations.................................................................................................................................................................. 215
Anomalies................................................................................................................................................................. 215
Transversus abdominis .................................................................................................................................................. 216
Synonyms ................................................................................................................................................................. 216
Variations.................................................................................................................................................................. 216
Anomalies................................................................................................................................................................. 216
Rectus abdominis .......................................................................................................................................................... 216
Synonyms ................................................................................................................................................................. 216
Variations.................................................................................................................................................................. 217
Anomalies................................................................................................................................................................. 218
DOI: 10.1201/9781003083535-4 207

Rectus abdominis lateralis............................................................................................................................................. 218

Synonyms ................................................................................................................................................................. 218
Variations.................................................................................................................................................................. 218
Anomalies................................................................................................................................................................. 218
Pyramidalis.................................................................................................................................................................... 218
Synonyms ................................................................................................................................................................. 218
Variations.................................................................................................................................................................. 218
Anomalies................................................................................................................................................................. 219
Cremaster ...................................................................................................................................................................... 219
Synonyms ................................................................................................................................................................. 219
Variations.................................................................................................................................................................. 219
Anomalies................................................................................................................................................................. 220
Quadratus lumborum..................................................................................................................................................... 220
Synonyms ................................................................................................................................................................. 220
Variations.................................................................................................................................................................. 220
Anomalies................................................................................................................................................................. 221
Iliacus ............................................................................................................................................................................ 221
Synonyms ................................................................................................................................................................. 221
Variations.................................................................................................................................................................. 221
Anomalies................................................................................................................................................................. 222
Iliacus minor.................................................................................................................................................................. 222
Synonyms ................................................................................................................................................................. 222
Variations.................................................................................................................................................................. 222
Anomalies................................................................................................................................................................. 222
Psoas major ................................................................................................................................................................... 223
Synonyms ................................................................................................................................................................. 223
Variations.................................................................................................................................................................. 223
Anomalies.................................................................................................................................................................224
Psoas minor ...................................................................................................................................................................224
Synonyms .................................................................................................................................................................224
Variations..................................................................................................................................................................224
Anomalies................................................................................................................................................................. 225
Perineal, Coccygeal, and Anal Musculature ...................................................................................................................... 225
Trunk Muscles 209
Coccygeus ..................................................................................................................................................................... 225

Synonyms ................................................................................................................................................................. 225
Variations.................................................................................................................................................................. 225
Anomalies................................................................................................................................................................. 226
Sacrococcygeus anterior................................................................................................................................................ 226
Synonyms ................................................................................................................................................................. 226
Variations.................................................................................................................................................................. 226
Anomalies................................................................................................................................................................. 227
Levator ani..................................................................................................................................................................... 227
Synonyms ................................................................................................................................................................. 227
Variations.................................................................................................................................................................. 227
Anomalies................................................................................................................................................................. 228
Sphincter ani externus ................................................................................................................................................... 228
Synonyms ................................................................................................................................................................. 228
Variations.................................................................................................................................................................. 228
Anomalies................................................................................................................................................................. 229
Transversus perinei profundus (Deep transverse perinei)............................................................................................. 229
Synonyms ................................................................................................................................................................. 229
Variations.................................................................................................................................................................. 229
Anomalies................................................................................................................................................................. 230
Transversus perinei superfcialis (Superfcial transverse perinei)................................................................................. 230
Synonyms ................................................................................................................................................................. 230
Variations.................................................................................................................................................................. 230
Anomalies................................................................................................................................................................. 230
Sphincter urethrae ......................................................................................................................................................... 230
Synonyms ................................................................................................................................................................. 230
Variations.................................................................................................................................................................. 231
Anomalies................................................................................................................................................................. 231
Ischiopubicus (Not Illustrated)...................................................................................................................................... 231
Synonyms ................................................................................................................................................................. 231
Variations.................................................................................................................................................................. 231
Anomalies................................................................................................................................................................. 231
Ischiocavernosus ........................................................................................................................................................... 231
Synonyms ................................................................................................................................................................. 231

Variations.................................................................................................................................................................. 231
Anomalies................................................................................................................................................................. 231
Bulbospongiosus ........................................................................................................................................................... 232
Synonyms ................................................................................................................................................................. 232
Variations.................................................................................................................................................................. 232
Anomalies................................................................................................................................................................. 232
Thorax, Spine, and Back Muscles...................................................................................................................................... 232
Diaphragm..................................................................................................................................................................... 232
Synonyms ................................................................................................................................................................. 232
Variations.................................................................................................................................................................. 233
Anomalies................................................................................................................................................................. 234
Hepatodiaphragmaticus (Not Illustrated)...................................................................................................................... 234
Synonyms ................................................................................................................................................................. 234
Variations.................................................................................................................................................................. 234
Anomalies................................................................................................................................................................. 234
Transversus thoracis ...................................................................................................................................................... 234
Synonyms ................................................................................................................................................................. 234
Variations.................................................................................................................................................................. 235
Anomalies................................................................................................................................................................. 235
Intercostales externi (External intercostals) .................................................................................................................. 235
Synonyms ................................................................................................................................................................. 235
Variations.................................................................................................................................................................. 236
Anomalies................................................................................................................................................................. 236
Intercostales interni (Internal intercostals).................................................................................................................... 236
Synonyms ................................................................................................................................................................. 236
Variations.................................................................................................................................................................. 236
Anomalies................................................................................................................................................................. 236
Intercostales intimi (Innermost intercostals)................................................................................................................. 237
Synonyms ................................................................................................................................................................. 237
Variations.................................................................................................................................................................. 237
Anomalies................................................................................................................................................................. 237
Subcostales (Subcostal Muscles) .................................................................................................................................. 237
Trunk Muscles 211
Synonyms ................................................................................................................................................................. 237

Variations.................................................................................................................................................................. 237
Anomalies................................................................................................................................................................. 238
Supracostalis anterior .................................................................................................................................................... 238
Synonyms ................................................................................................................................................................. 238
Variations.................................................................................................................................................................. 238
Anomalies................................................................................................................................................................. 238
Supracostalis posterior .................................................................................................................................................. 238
Synonyms ................................................................................................................................................................. 238
Variations.................................................................................................................................................................. 238
Anomalies................................................................................................................................................................. 239
Serratus posterior superior ............................................................................................................................................ 239
Synonyms ................................................................................................................................................................. 239
Variations..................................................................................................................................................................240
Anomalies.................................................................................................................................................................240
Serratus posterior inferior .............................................................................................................................................240
Synonyms .................................................................................................................................................................240
Variations.................................................................................................................................................................. 241
Anomalies................................................................................................................................................................. 241
Levatores costarum breves and longi ............................................................................................................................ 241
Synonyms ................................................................................................................................................................. 241
Variations.................................................................................................................................................................. 242
Anomalies................................................................................................................................................................. 242
Interspinales .................................................................................................................................................................. 242
Synonyms ................................................................................................................................................................. 242
Variations.................................................................................................................................................................. 242
Anomalies................................................................................................................................................................. 243
Sacrococcygeus posterior..............................................................................................................................................244
Synonyms .................................................................................................................................................................244
Variations..................................................................................................................................................................244
Anomalies.................................................................................................................................................................244
Intertransversarii ...........................................................................................................................................................244
Synonyms .................................................................................................................................................................244

Variations..................................................................................................................................................................244
Anomalies.................................................................................................................................................................244
Rotatores longus and brevis .......................................................................................................................................... 245
Synonyms ................................................................................................................................................................. 245
Variations.................................................................................................................................................................. 245
Anomalies................................................................................................................................................................. 245
Multifdus ...................................................................................................................................................................... 245
Synonyms ................................................................................................................................................................. 245
Variations.................................................................................................................................................................. 245
Anomalies.................................................................................................................................................................246
Semispinalis ..................................................................................................................................................................246
Synonyms .................................................................................................................................................................246
Variations..................................................................................................................................................................246
Anomalies................................................................................................................................................................. 247
Splenius capitis and Splenius cervicis........................................................................................................................... 247
Synonyms ................................................................................................................................................................. 247
Variations.................................................................................................................................................................. 247
Anomalies................................................................................................................................................................. 247
Rhomboatloideus........................................................................................................................................................... 247
Synonyms ................................................................................................................................................................. 247
Variations..................................................................................................................................................................248
Anomalies.................................................................................................................................................................248
Atlantomastoideus......................................................................................................................................................... 249
Synonyms ................................................................................................................................................................. 249
Variations.................................................................................................................................................................. 249
Anomalies................................................................................................................................................................. 249
Spinalis.......................................................................................................................................................................... 249
Synonyms ................................................................................................................................................................. 249
Variations.................................................................................................................................................................. 249
Anomalies................................................................................................................................................................. 251
Longissimus .................................................................................................................................................................. 251
Synonyms ................................................................................................................................................................. 251
Trunk Muscles 213

Variations.................................................................................................................................................................. 251
Anomalies................................................................................................................................................................. 251
Iliocostalis ..................................................................................................................................................................... 252
Synonyms ................................................................................................................................................................. 252
Variations.................................................................................................................................................................. 252
Anomalies................................................................................................................................................................. 252
MUSCLES OF THE ABDOMINAL REGION attach inferiorly to the iliac crest while the fbers from the
middle and upper ribs end in the external oblique aponeurosis,
Obliquus externus abDOMinis which forms the inguinal ligament and the anterior layer of
(external Oblique) (figure 4.1) the rectus sheath (Saga and Takahashi 2016; Standring 2016).
See also: Obliquus abdominis externus profundus Innervation
External oblique is innervated by the lower fve intercostal
Synonyms nerves and the subcostal nerve (Standring 2016).
N/A
Comparative Anatomy
Typical Presentation External oblique has a similar typical presentation in the
Description apes, extending from the lower ribs to the pelvic region, and
External oblique is typically attached superiorly to ribs 5 may vary in which rib provides the superiormost attach-
through 12 (Standring 2016). The fbers from the lowest ribs ment (Diogo et al. 2010, 2012, 2013a,b, 2017). The external
FIGURE 4.1 Lateral abdominal muscles in anterior view.

oblique of bonobos is innervated by the lower intercostal Clinical Implications

nerves and also by iliohypogastric and ilioinguinal nerves Dsouza et al. (2017) observe that the bilateral absence of
(Miller 1952). the external oblique and its aponeurosis can increase the
risk for hernia.
Variations
Description
Obliquus abDOMinis externus prOfunDus (figure 4.1)
The external oblique can vary in the number of costal
attachments (Macalister 1875; Mori 1964; Rickenbacher See also: Obliquus externus abdominis
et al. 1985; Bergman et al. 1988; Standring 2016; Saga
and Takahashi 2016). Some attachments, or even the Synonyms
entire muscle, may be doubled (Knott 1883b; Macalister This muscle is also referred to as obliquus externus secun-
1875; Bergman et al. 1988; Standring 2016). The superi- dus s. accessorius (Gruber 1875a), obliquus externus profun-
ormost and inferiormost attachments, or even the entire dus s. minor (Knott 1880), and obliquus externus abdominis
muscle, can also be absent (Knott 1883b; Macalister 1875; minor s. secundus (Knott 1883b) (Saga and Takahashi 2016).
Standring 2016; Dsouza et al. 2017). External oblique can
have connections to, or may be continuous with, pectora- Typical Presentation
lis major, serratus anterior, latissimus dorsi, the external This muscle is only present as a variation.
intercostals, or serratus posterior inferior (Knott 1883b;
Macalister 1875; Bergman et al. 1988; Standring 2016). Comparative Anatomy
Additional slips may originate from the lower ribs, the N/A
transverse process of the frst lumbar vertebra, the lumbar
Variations
fascia below rib 12, or the fascia over the ffth or sixth
intercostal space (Macalister 1875; Rickenbacher et al. Description
1985; Bergman et al. 1988). Obliquus abdominis externus profundus refers to distinct
The external oblique is associated with three variable muscle bellies that are variably associated with the external
supernumerary muscles. The saphenous muscle refers to a oblique. It may present as a deep accessory bundle that origi-
slip of muscle that is attached to the inguinal ligament and nates from the middle/lower ribs or the internal oblique fas-
loops under the saphenofemoral junction, blending with sar- cia to attach to the anterior superior iliac spine, iliac crest,
torius and the adductor longus (Tyrie 1894; du Plessis and inguinal ligament, or rectus sheath (Macalister 1875; Knott
Loukas 2016; Saga and Takahashi 2016). The interfoveolar 1883a,b; Nakayama and Okuda, 1952; Bergman et al. 1988;
muscle refers to muscle fbers within the interfoveolar liga- Saga and Takahashi 2016). This muscle can have two heads
ment (Kudo and Otobe 1952; Saga and Takahashi 2016). (Nakayama and Okuda 1952; Miyauchi et al. 1986). Knott
Obliquus externus abdominis profundus refers to variable (1883b) records an origin from ribs nine and ten. Nakayama
bundles that are occasionally associated with the external and Okuda (1952) report an origin from the eleventh rib.
oblique (see the entry for this muscle). Miyauchi et al. (1986) record origins from ribs fve, six,
or seven. It can either be fused with the external oblique
Prevalence or separated from it via a layer of fascia (Macalister 1875;
Based on a sample of 166 Japanese adults (332 sides), Mori Miyauchi et al. 1986; Bergman et al. 1988).
(1964) found that the superiormost attachment of the exter- Miyauchi et al. (1986) note the confusion in the literature
nal oblique was from the fourth rib in only 12% of cases on of the many variant bundles that have been given the name
the left side (0% on the right side), the ffth rib in 84.3% of obliquus abdominis externus profundus, and these authors
cases on the left side and 81.9% of cases on the right side, as well as Kodama (1986) argue for classifcation of these
the sixth rib in 14.4% of cases on the left side and 18.1% of bundles based on innervation (Saga and Takahashi 2016).
cases on the right side, and never from the seventh rib (0%).
Kudo and Otobe (1952) recorded the presence of the inter- Innervation
foveolar muscle in 46% of their study population. Obliquus abdominis externus profundus is supplied by the
ramus muscularis externus from the intercostal nerve of the
Anomalies corresponding segment (Miyauchi et al. 1986).
Description
Prevalence
Poland syndrome can lead to the absence of the external
oblique (Cingel et al. 2013). Prune belly syndrome is also Earlier studies typically report a prevalence for this muscle
associated with the partial reduction or complete absence <10%. Loth (1912, 1931) reports a prevalence of 3.8% in
of this muscle (Adebonojo 1973). Black individuals and 7% in Europeans. Kudo and Otobe
(1952) report a prevalence of 3% in a Chinese population.
Prevalence Nakayama and Okuda (1952) report a prevalence of 1.9%.
N/A More recent studies provide higher prevalence rates. Kodama
Trunk Muscles 215
(1986) reports a prevalence of 36.5%. This muscle was pres- opposite the eighth or twelfth rib (Macalister 1875; Knott
ent in 11 out of 50 sides (22%), from 8 out of the 25 Japanese 1883b; Bergman et al. 1988). The internal oblique muscle
cadavers studied by Miyauchi et al. (1986). Miyauchi et al. fascia may also be connected to the fascia of the external
(1986) found that the muscle originated via one slip from the oblique (Tekelioglu et al. 2015).
ffth rib in two cases, from the sixth rib in six cases, and An additional slip can connect to the eighth costal car-
from the seventh rib in one case. It originated via two slips tilage or the transverse process of the second lumbar ver-
from the ffth and sixth ribs in two cases. Obliquus abdomi- tebra (Bergman et al. 1988). Internal oblique can be fused
nis externus profundus was fused with the external oblique in with transversus abdominis (Macalister 1875; Knott 1883b;
four cases (8%) (Miyauchi et al. 1986). Standring 2016). The portion of the muscle that arises from
the iliac crest may be separated into an anterior and pos-
Anomalies terior part, the latter of which is referred to as accessory
N/A internal oblique (Chouke 1935; Bergman et al. 1988). There
are variations in how the aponeurosis of internal oblique
Clinical Implications contributes to the rectus sheath (see prevalence informa-
N/A tion, below). The rectus sheath can also vary in the degree
to which it is aponeurotic, as in some cases the sheath can
be more muscular (Monkhouse and Khalique 1986).
Obliquus internus abDOMinis (internal Oblique)
(figures 4.1 anD 4.2)
Prevalence
Synonyms Loth (1912) found that the superiormost attachment of the
N/A internal oblique was onto rib 11 in 31% of cases, rib 10 in
66.5% of cases, rib 9 in 1% of cases, and rib 8 in 1.5% of
Typical Presentation cases. In a sample of 200 individuals, Mori (1964) also
Description observed that the most common superior costal attachments
Internal oblique is deep to external oblique (Standring were to rib 10 (56% of right sides, 64% of left sides) and
2016). It originates from the iliopectineal arch, iliac crest, rib 11 (34% of right sides, 36% of left sides). According to
and the thoracolumbar fascia (Standring 2016). The fbers Monkhouse and Khalique (1986), Chouke (1935) reported
originating from the iliopectineal arch fuse with the apo- an accessory internal oblique with high incidence in a pop-
neurosis of transversus abdominis and attach to the pubis ulation of 136 cadavers, in which the accessory muscle was
and pectineal line to form the conjoint tendon (Standring only absent four times (97% prevalence).
2016). The fbers that originate from the iliac crest end in In a sample of 40 cadavers, Monkhouse and Khalique
the anterior aponeurosis that forms the rectus sheath, and (1986) found that in 24 cases (60%) the aponeurosis of
the posterior fbers insert onto the cartilage and inferior internal oblique split to enclose rectus abdominis, with the
borders of ribs 8/9 to rib 12 (Standring 2016). aponeurosis of external oblique remaining anterior and the
aponeurosis of transversus abdominis remaining posterior.
Innervation In 11 cases (27.5%), the aponeurosis of internal oblique
Internal oblique is innervated by the lower fve intercostal passed entirely anterior to rectus abdominis, while the
nerves, the subcostal nerve, the iliohypogastric nerve, and aponeurosis of transversus abdominis remained posterior
the ilioinguinal nerve (Standring 2016). to rectus abdominis. In fve cases (12.5%), the aponeurosis
of transversus abdominis split to enclose rectus abdomi-
Comparative Anatomy nis, and the anterior layer of this aponeurosis fused with
Internal oblique has a similar typical presentation in the apes, the aponeuroses of both the external and internal obliques.
extending from the iliac crest, thoracolumbar fascia, and Saga and Takahashi (2016) compare this study with that of
inguinal ligament to join the rectus sheath, lower four ribs, McVay and Anson (1940) and note that the latter authors
and the pubis via the conjoint tendon (Sonntag 1923; Miller found variations corresponding to the third presentation
1952; Gibbs 1999; Diogo et al. 2010, 2012, 2013a,b, 2017). described by Monkhouse and Khalique (1986) in 43 out of
56 cases (76.8%) and two cases (3.6%) that corresponded to
Variations the frst presentation described by these authors.
Description
Internal oblique can vary in its costal attachments, and Anomalies
the inguinal or anterior superior part of this muscle may Description
be absent (Macalister 1875; Mori 1964; Rickenbacher et al. Prune belly syndrome is associated with the partial reduc-
1985; Bergman et al. 1988). A fbrous band that is a tendi- tion or complete absence of this muscle (Adebonojo 1973).
nous connection between internal oblique and the internal
intercostal muscles often “interrupts” the muscle, typically Prevalence
opposite the tenth or eleventh rib, but it can also be present N/A
Clinical Implications A supernumerary muscle termed tensor laminae posteri-

Fibrous adhesions between the fascia of the internal oris vaginae musculi rectus or puboperitonealis may be pres-
oblique and the external oblique and other fbrous bands ent (Gruber 1873a,b; Macalister 1875; Bergman et al. 1988;
within this muscle can prevent a successful transverse Saga and Takahashi 2016). This muscle is comprised of slips
abdominis plane (TAP) block procedure (Tekelioglu et al. that originate inferiorly from the iliopectineal line and trans-
2015; Lew and Gray 2012). versus fascia and interdigitate with transversus abdominis
(Gruber 1873a,b; Macalister 1875; Saga and Takahashi 2016).
transversus abDOMinis (figure 4.1) Prevalence

Synonyms Knott (1883b) found that in 29 out of 36 cases (80.6%) trans-
versus abdominis had six costal attachments, in four cases
This muscle may also be referred to as transversalis
there were seven costal attachments (11.1%), and in three
(Macalister 1875) or transversalis abdominis (Knott 1883b).
cases there were fve costal attachments (8.3%). Anson and
Typical Presentation McVay (1938) report that transversus abdominis arises from
the whole of the inguinal ligament in only 3% of cases.
Description
Transversus abdominis is deep to both the internal and Anomalies
external oblique (Standring 2016). It originates from the Description
iliopectineal arch, iliac crest, thoracolumbar fascia, and the
Macalister (1875) notes that Charvet found the absence of
lower six costal cartilages (Standring 2016). The muscle
transversus abdominis in ectopia vesicae. Prune belly syn-
ends in an aponeurosis that inserts mostly into the rectus
drome is also associated with the partial reduction or com-
sheath (Standring 2016; Saga and Takahashi 2016). Its
plete absence of this muscle (Adebonojo 1973).
lower fbers fuse with the aponeurosis of internal oblique
to form the conjoint tendon and insert onto the pubis and
pectineal line (Standring 2016). Prevalence
N/A
Innervation
Transversus abdominis is supplied by the lower fve inter- Clinical Implications
costal nerves, the subcostal nerve, the iliohypogastric nerve, N/A
and the ilioinguinal nerve (Standring 2016).
Comparative Anatomy rectus abDOMinis (figure 4.2)

Transversus abdominis has a similar typical presentation Synonyms
in the apes, with attachments to the lower costal cartilages, N/A
thoracolumbar fascia, and iliac crest and contributions to
the conjoint tendon and posterior layer of the rectus sheath Typical Presentation
(Champneys 1872; Sonntag 1923; Miller 1952; Gibbs 1999; Description
Diogo et al. 2010, 2012, 2013a,b, 2017). Rectus abdominis spans the length of the anterior abdomi-
nal wall (Standring 2016). It originates by a lateral tendon
Variations
that is attached to the pubic crest and a medial tendon that
Description is attached closer to the pubic symphysis (Standring 2016).
Transversus abdominis may be absent in rare cases Rectus abdominis typically attaches superiorly to the ffth
(Macalister 1875; Rickenbacher et al. 1985; Bergman through seventh costal cartilages and to the xiphoid process
et al. 1988; Saga and Takahashi 2016; Standring 2016). It (Standring 2016). Rectus abdominis is interrupted by three
may also be doubled (Macalister 1875). The attachments transverse tendinous intersections that typically intersect
of transversus abdominis to the ribs can vary in number, the muscle at (1) the level of umbilicus, (2) the level of the
generally ranging from fve attachments to seven attach- end of the xiphoid process, and (3) the area approximately
ments (Macalister 1875; Knott 1883b; Bergman et al. 1988). midway between those two (Standring 2016).
Transversus abdominis can be fused with internal oblique
(Macalister 1875; Knott 1883b; Standring 2016). A tendi- Innervation
nous intersection may be present (Macalister 1875). The Rectus abdominis is innervated by the lower six or seven
spermatic cord may travel through the inferior portion of thoracic spinal nerves and sometimes the ilioinguinal nerve
transversus abdominis (Macalister 1875; Knott 1883b; Saga (Standring 2016).
and Takahashi 2016). There are variations in how the apo-
neurosis of transversus abdominis contributes to the rectus Comparative Anatomy
sheath (see the entry for obliquus internus abdominis for Rectus abdominis has not been described in gibbons (Diogo
prevalence information). et al. 2012). In gorillas, it extends from the xiphisternum and
Trunk Muscles 217
FIGURE 4.2 Medial abdominal muscles and rectus abdominis lateralis in anterior view.
costal cartilages fve through nine to the ventral spine of the 1883b; Bergman et al. 1988; Saga and Takahashi 2016;
pubis and symphyseal ligament (Gibbs 1999; Diogo et al. Standring 2016). Muscular slips may be found between
2010). In common chimpanzees and bonobos, it extends the rectus abdominis and the deep inguinal ring (Bergman
from the costal cartilages fve through seven to the pubic et al. 1988; Saga and Takahashi 2016). There are variations
crest and has four tendinous intersections (Champneys in how the aponeuroses of the muscles that surround rectus
1872; Sonntag 1923; Miller 1952; Gibbs 1999; Diogo et al. abdominis contribute to the rectus sheath (see the entry for
2013a, 2017). Similarly, in orangutans, it extends between obliquus internus abdominis for prevalence information).
the lower costal cartilages and pubic crest and has four ten-
dinous intersections (Diogo et al. 2013b). Prevalence
Knott (1883b) studied the variation in the tendinous inscrip-
Variations tions of rectus abdominis in 60 cases. In two cases (3.3%),
Description the tendinous inscription at the level of the xiphoid was
Rectus abdominis may be reduced, absent, or doubled absent unilaterally, and in two other cases, it was absent
(Macalister 1875; Knott 1883b; Bergman et al. 1988; Saga bilaterally (3.3%). In one case (1.7%), the inscription inter-
and Takahashi 2016). The tendinous intersections can mediate between that at the level of the xiphoid and that at
vary in number and may only extend halfway across the the level of umbilicus was absent. In fve cases (8.3%), an
muscle (Macalister 1875; Knott 1883b; Mori 1964; Sato inscription below the level of umbilicus was present.
1968c; Saga and Takahashi 2016; Standring 2016). One or In a study of 400 sides from 200 cadavers by Mori
two incomplete intersections may be found below the level (1964), he found that on 313 sides (78.25%), rectus abdomi-
of umbilicus (Figure 4.2) (Macalister 1875; Knott 1883b; nis attached to ribs fve, six, and seven, which corresponds
Standring 2016). The costal attachments can also vary in to Loth’s (1912) “normal type.” Three presentations corre-
number (Macalister 1875; Mori 1964; Bergman et al. 1988; spond to Loth’s (1912) “primitive type” and made up 8.1%
Saga and Takahashi 2016; Standring 2016). The slip to rib of cases, including origin from ribs 4 and 5 on 8 sides (2%),
fve may be absent (Bergman et al. 1988; Standring 2016). origin from ribs 4, 5, and 6 on 12 sides (3%), and origin
The superiormost attachment may extend to ribs two, three, from ribs 5 and 6 on 13 sides (3.1%). Two presentations cor-
or four, the sternum, or the clavicle (Macalister 1875; Knott respond to Loth’s (1912) “progressive type” made up 13%
of cases and include origin from ribs 6 and 7 in 32 cases
(8%) and origin from ribs 6, 7, and 8 in 20 cases (5%). Mori longitudinally to insert onto the iliac crest (Macalister
(1964) also found that tendinous intersections into the mus- 1875; Sato 1968c; Saga and Takahashi 2016). Sato (1968c)
cle varied among two (1% of cases), three (36% of cases), reported an origin from ribs 10 through 12.
four (61% of cases), and fve intersections (2% of cases).
Innervation
Sato (1968c) reported that in Kyushu-Japanese males,
three tendinous intersections were present in 250 out of 416 N/A
sides (60%), 4 in 125 sides (30%), 2 in 31 sides (7.45%), and
Prevalence
5 in 10 sides (2.4%). In females, three tendinous intersec-
tions were present in 153 out of 268 sides (57.1%), 4 in 83 Sato (1968c) found that rectus abdominis lateralis was
sides (31%), 2 in 28 sides (10.4%), and 5 in 4 sides (1.5%). present in 18 out of 212 sides (8.49%) in Kyushu-Japanese
females and in 32 out of 324 sides (9.88%) in males.
Anomalies
Anomalies
Description
N/A
Macalister (1875) notes that Charvet found the absence of
rectus abdominis in ectopia vesicae. Prune belly syndrome Clinical Implications
is associated with the partial reduction or complete absence N/A
of this muscle (Adebonojo 1973). Bersu and Ramirez-Castro
(1977) and Aziz (1979) note that in individuals with trisomy
18, there may be poor development of the tendinous inter- pyraMiDalis (figure 4.2)
sections of rectus abdominis. Diastasis recti can also occur Synonyms
in individuals with trisomy 18 (Aziz 1979; Roberts et al.
This muscle is also referred to as pyramidalis abdominis
2016) and was observed in a female individual with XO/
(Knott 1883b).
XY mosaicism and partial trisomy 9p (Klasen et al. 1981).
Mieden (1982) describes an infant with median cleft lip, Typical Presentation
hypotelorism, and alobar holoprosencephaly. On the left
side, the intermediate tendinous intersection was 1 cm lower Description
than that of the one on the right side (Mieden 1982). Itoh Pyramidalis is a triangular muscle situated anterior to the
et al. (1991) describe fetal akinesia/hypokinesia sequence lower portion of rectus abdominis within the rectus sheath
in one male and one female infant, each with a suite of (Standring 2016). It originates from the pubis and in an apex
anatomical anomalies. In the male, the rectus abdominis that is attached to the linea alba about halfway between the
muscles showed focal atrophy. pubis and umbilicus (Standring 2016).
Roberts et al. (2016) state that diastasis rectus abdominis, Pyramidalis is typically innervated by the subcostal nerve
among other abdominal wall defects, is present in over 50% (Standring 2016). It is occasionally supplied by the ilio-
of individuals with trisomy 18. hypogastric or ilioinuginal nerves (Bergman et al. 1988;
Standring 2016).
N/A Comparative Anatomy
Pyramidalis has not been described in orangutans (Ashley-
rectus abDOMinis lateralis (figure 4.2) Montagu 1939; Gibbs 1999; Diogo et al. 2013b). It is often
absent in the other apes, though its presence has been
Synonyms described in some dissection descriptions for gorillas, gib-
N/A bons, bonobos, and common chimpanzees (Ashley-Montagu
1939; Miller 1952; Diogo et al. 2010, 2012, 2013a, 2017).
Typical Presentation Ashley-Montagu (1939) reports that it is present in about 36%
This muscle is only present as a variation. of gibbons, 33% of gorillas, and about 60% of chimpanzees.
Comparative Anatomy Variations
N/A Description
Variations Pyramidalis varies substantially in size between individ-
uals (Bergman et al. 1988; Standring 2016). The average
Description length is about 6–7 cm, and the average width is about
Rectus abdominis lateralis (Kelch) is a supernumerary 1.5–2 cm (Anson et al. 1938; Sinha and Kumar 1985; Saga
muscle that is situated between the external and internal and Takahashi 2016). Pyramidalis may extend to umbili-
obliques (Macalister 1875; Saga and Takahashi 2016). It cus (Macalister 1875) or send a tendinous slip to umbili-
originates from the lower border of rib ten and descends cus (Knott 1883b) (Figure 4.2). This muscle may also be
Trunk Muscles 219
interrupted by a tendinous intersection (Macalister 1875; study of 31 Indian male cadavers, Sinha and Kumar (1985)
Knott 1883b). Pyramidalis can be doubled unilaterally found that pyramidalis was absent bilaterally in 5.88% and
or bilaterally (Winslow 1732; Cruveilhier 1837; Hallett absent unilaterally in 2.94% of cadavers. Pyramidalis was
1848; Macalister 1875; Le Double 1897; Chudzinski present on only 4 out of 112 sides (3.6%) of the 56 cadav-
1898; Anson et al. 1938; Saga and Takahashi 2016; ers examined Monkhouse and Khalique (1986). Dickson
Standring 2016). (1999) found that pyramidalis was absent in 42 out of 60
Pyramidalis is often asymmetric and may be absent northern European female subjects (70%). Didia et al.
unilaterally or bilaterally (Macalister 1875; Knott 1883b; (2009) observed that this muscle was absent in 2 out of 24
Chouke 1935; Anson et al. 1938; Beaton and Anson 1939; Nigerian male cadavers (8.33%). In a study of 96 Greek
Mori 1964; Sato 1968c; Sinha and Kumar 1985; Monkhouse cadavers (50 male, 46 female), Natsis et al. (2016) found
and Khalique 1986; Bergman et al. 1988; Dickson 1999; that pyramidalis was absent in 6.2% of specimens, was
Didia et al. 2009; Natsis et al. 2016; Saga and Takahashi more often present bilaterally, and more often present in
2016; Standring 2016; Das et al. 2017). When it is absent, the females (91.3% of specimens) than in males (68% of speci-
size of the lower portion of the rectus abdominis (Bergman mens). In a study of 25 Indian cadavers (17 male, 8 female),
et al. 1988) or the internal oblique (Macalister 1875) may be Das et al. (2017) found that pyramidalis was absent in 8%
increased. of cases, was more often present bilaterally, and was more
often present in males (94.11%) than in females (87.5%).
Prevalence
Saga and Takahashi (2016) report that it is absent in 10% Anomalies
of cases. Knott (1883b) found that this muscle was absent
N/A
bilaterally in 9 out of 60 subjects (15%) and absent uni-
laterally in fve subjects (8.3%). In six subjects (10%), the Clinical Implications
left muscle was smaller than the right. A tendinous slip Understanding variation in the presentation of pyramidalis
extending to umbilicus was present in one subject (1.7%), is important during imaging, so as to not mistake it for a
and a horizontal tendinous inscription was present bilat- mass (Natsis et al. 2016).
erally in another (1.7%) (Knott 1883b). Based on a study
of 123 cadavers, Chouke (1935) reported that pyramida-
lis was absent in over 23% of cases. Anson et al. (1938) creMaster (figure 4.2)
reported that this muscle was absent in 10.8% of 165
Synonyms
subjects. Beaton and Anson (1939) reported unilateral or
bilateral absence in 20.3% of a White population and in N/A
12.5% of a Black population.
Summarizing the work of other Japanese research-
ers, Mori (1964) notes that pyramidalis is absent in about Description
5%–6% of Japanese individuals. Mori (1964) also found Cremaster covers the spermatic cord and extends to the
that the most common presentation (21.7% of cases) of this testis (Standring 2016). It originates from the inferomedial
muscle was the apex of right pyramidalis being higher than border of the internal oblique and transversus abdominis
the left, and the origin of the left pyramidalis elongated to and attaches medially to the pubis (Standring 2016). In
the surface of the pubis. Other presentations of this muscle females, cremaster is represented by fbers on the round lig-
include: the apex of the left pyramidalis is higher than the ament of the uterus (Bergman et al. 1988; Standring 2016).
right, and the origin of the right muscle is elongated to the
surface of the pubis (19.3%); the apex of the right pyramida- Innervation
lis is higher than the left, and the origin of the right muscle Cremaster is innervated by the genital branch of the genito-
is elongated to the surface of the pubis (15.7%); the apex of femoral nerve (Standring 2016).
the left pyramidalis is higher than the right and its origin is
elongated to the pubis (13.3%); the apex of the muscle on Comparative Anatomy
both sides is at the same level and the right origin is elon- Cremaster has a similar typical presentation in the apes,
gated to the anterior surface of the pubis (9.6%); the apex of extending from the internal oblique and transversus abdom-
the right pyramidalis is higher than the left, and the origin of inis towards the “inguinal ligament” (Miller 1952; Gibbs
the muscle on both sides is at the same level (6%); the apex 1999; Diogo et al. 2010, 2012, 2013a,b, 2017). As Gibbs
of the left pyramidalis is higher than the right, and the ori- (1999) notes, apes do not have a true inguinal ligament but
gin of both sides is at the same level (3.6%), the apex of the a series of tendinous arches over the femoral vessels and
muscle on both sides is at the same level, and the left origin nerves in the inguinal region.
is elongated to the anterior surface of the pubis (2.4%); and
the pyramidalis is divided into two parts (1.1%). Variations
Sato (1968c) reported that pyramidalis is absent in 47 Description
out of 424 sides (11.08%) from Kyushu-Japanese males Cremaster is thickest in younger males (Standring 2016).
and absent in 30 out of 274 sides (10.95%) in females. In a The medial portion of the muscle that attaches to the pubis
may be absent (Standring 2016). Cremaster may origi- four lumbar vertebrae, and superiorly to the twelfth rib and
nate from the transversalis fascia or fuse with transversus twelfth thoracic vertebra (Standring 2016).
abdominis (Macalister 1875; Knott 1883b; Bergman et al.
1988; Saga and Takahashi 2016). It may also have two Innervation
heads (Macalister 1875; Bergman et al. 1988). Quadratus lumborum is innervated by the twelfth thoracic
nerve and upper three/four lumbar spinal nerves (Standring
Prevalence 2016).
N/A
Comparative Anatomy
Anomalies Quadratus lumborum has a similar typical presentation in
N/A the apes, extending from the iliac crest to the upper lumbar
vertebrae and last rib (Gibbs 1999; Diogo et al. 2010, 2012,
Clinical Implications 2013a,b, 2017). In bonobos, the muscle may extend to the
N/A penultimate rib (Miller 1952; Diogo et al. 2017).
Variations
quaDratus luMbOruM (figure 4.3)
Description
Synonyms The bundles that comprise quadratus lumborum may vary in
This muscle may also be referred to as ileolumbalis size and number (Macalister 1875; Knott 1883b; Rickenbacher
(Meyer), scalenus lumborum (Meyer), or rectus abdominis et al. 1985; Standring 2016). The extent of the develop-
posticus (Luschka) (Knott 1883b; Macalister 1875; Bakkum ment of the attachments to the vertebrae and the number of
and Miller 2016). these attachments may vary (Macalister 1875; Knott 1883b;
Rickenbacher et al. 1985; Bergman et al. 1988; Bakkum and
Typical Presentation Miller 2016). The superior attachment of this muscle may
Description extend to rib eleven, and the superomedial attachment may
Quadratus lumborum is part of the posterior abdominal extend to the tenth and/or eleventh thoracic vertebrae (Knott
wall. It attaches inferiorly to the iliac crest and iliolumbar 1883b; Macalister 1875; Rickenbacher et al. 1985; Bergman et
ligament, medially to the transverses processes of the upper al. 1988; Bakkum and Miller 2016; Saga and Takahashi 2016).
FIGURE 4.3 Posterior abdominal wall muscles in anterior view. Iliacus minimus is illustrated on the right side, and the other muscles
are illustrated on the left side.
Trunk Muscles 221
Knott (1883b) found that the attachment to the body of the Description
twelfth thoracic vertebra was present in 8 out of 30 speci- Iliacus may present as a completely separate muscle from
mens (26.7%). This author also found that two specimens psoas major (Macalister 1875; Bergman et al. 1988) or be
exhibited a slip to the eleventh thoracic vertebra, one speci- completely fused with it (Aleksandrova et al. 2013; Saga
men had a slip to the eleventh rib, and three specimens had and Takahashi 2016). The origin of iliacus may be fused
slips to both the eleventh and twelfth thoracic vertebrae. It with fbers from quadratus lumborum (Macalister 1875).
is not clear if these six cases are part of the 30 subjects or The iliopsoas tendon may be partially or completely
another unspecifed sample size. divided into one, two, or three tendons (Tatu et al. 2002;
Polster et al. 2008; Shu and Safran 2011; Crompton et al.
Anomalies
2014; Philippon et al. 2014; Saga and Takahashi 2016).
Description Macalister (1875) found the muscle itself divided in two.
Prune belly syndrome may be associated with the par- One or more accessory slips/muscle bundles may be pres-
tial reduction or complete absence of this muscle (King ent medial or lateral to iliacus (Macalister 1875; Bergman
et al. 1961). Bersu et al. (1976) describe a male infant with et al. 1988; Spratt et al. 1996; Jelev et al. 2005; Vázquez
Hanhart syndrome. On both sides of this infant, there was et al. 2007; Astik and Dave 2011; Saga and Takahashi
an extensive fusion between the lumbar muscular com- 2016). Aleksandrova et al. (2013) categorize the varia-
ponents of the diaphragm, psoas major, and quadratus tions of iliacus into ten types including partial agenesis,
lumborum. complete separation from psoas major, complete fusion
with psoas major, the presence of accessory slips/mus-
Prevalence cle bundles, the presence of iliacus minor, the presence
N/A of iliacus minimus, and the division of the muscle into
superfcial and deep fbers.
Clinical Implications Iliacus is associated with several named accessory slips
N/A and muscles (see the entry for iliacus minor). Also see the
entry for psoas major for information on psoas quartus and
psoas tertius. Jelev et al. (2005) describe an accessory ilio-
iliacus (figure 4.3)
psoas muscle that resulted from the connection between
See also: Iliacus minor an accessory iliacus muscle and an accessory psoas major.
Iliacus minimus (Figure 4.3) refers to a slip that pierces the
Synonyms femoral nerve and inserts onto the lesser trochanter or joins
N/A the iliopsoas muscle or its tendon (Spratt et al. 1996; Tubbs
and Salter 2006a; Aristotle et al. 2013; Aleksandrova et al.
Typical Presentation 2013; Saga and Takahashi 2016). It may originate from the
iliolumbar ligament (Spratt et al. 1996; Tubbs and Salter
Description 2006a) or from the iliac crest (Aristotle et al. 2013). In the
Iliacus originates from the upper two-thirds of the iliac case described by Spratt et al. (1996), the muscle divided
fossa, the iliac crest, and the lateral aspect of the sacrum into two tendons that inserted onto the lesser trochanter and
(Standring 2016). Iliacus forms the iliopsoas muscle with the medial thigh.
psoas major as the two insert together via a tendon onto the
lesser trochanter of the femur (Standring 2016). Some fbers Prevalence
attach to the femur below the lesser trochanter (Standring Tatu et al. (2002) found that the iliopsoas tendon com-
2016). pletely split into two bundles in 2 out of 24 cases (8.3%)
and partially split into two bundles in two other cases
Innervation (8.3%). Crompton et al. (2014) studied the iliopsoas tendon
Iliacus is innervated by branches of the femoral nerve via MRI images in 50 children and found that at least one
(Standring 2016). bifd tendon was present in 13 children (26%), and 5 out of
the 37 children that were imaged bilaterally had bilateral
Comparative Anatomy bifd tendons (13.5%). Out of the total sample of 87 hips, 18
Iliacus has a similar typical presentation in the apes, had two distinct distal iliopsoas tendons (20.7%) (Crompton
extending from the iliac fossa to join with psoas major et al. 2014). Philippon et al. (2014) found that the iliopsoas
to insert onto the lesser trochanter (Champneys 1872; tendon was single-banded in 15 out of 53 cadaver hemipel-
Hepburn 1892; Beddard 1893; Sigmon 1974; Gibbs 1999; ves (28.3%), double-banded in 34 hemipelves (64.2%), and
Diogo et al. 2010, 2012, 2013a,b, 2017). Fibers may extend triple-banded in four hemipelves (7.5%).
distally onto the femoral shaft in gibbons, orangutans, and Based on dissections of 68 cadavers, Spratt et al. (1996)
bonobos (Hepburn 1892; Beddard 1893; Boyer 1935; Miller found four variant slips (5.9%) which included one case of
1952; Sigmon 1974; Diogo et al. 2012, 2013b, 2017). iliacus minimus, another accessory iliacus slip that extended
between the sacrum and the lesser trochanter, and two acces- Comparative Anatomy
sory lateral slips of psoas major. Jelev et al. (2005) found Das and Singh (1950) suggest that iliacus minor cor-
what they define as an accessory iliopsoas in only 1 out of responds to the iliacus externus of lower animals. Citing
108 cadavers over the course of 22 years of dissection (0.9%). three German anatomy texts and Gregory and Camp (1818),
Vázquez et al. (2007) found accessory slips or sheets of Babst et al. (2011) state that iliocapsularis can be found in
iliacus and psoas piercing or covering the femoral nerve primates, rats, reptiles, and birds.
in 19 out of 242 specimens (7.9%) from 121 cadavers. The
femoral nerve was pierced by a muscular slip in 17 speci-
Variations
mens (7%), in a sample composed of 12 iliacus slips, four
psoas slips, and a slip from both in one specimen. The fem- Description
oral nerve was covered by a sheet or slip in two specimens A detached portion or third head of iliacus is referred to as
(0.8%). Astik and Dave (2011) dissected 64 lumbar plexuses iliacus minor (Winslow) (Macalister 1875; Bergman et al.
from 32 cadavers and found accessory slips of iliacus in 1988; Saga and Takahashi 2016). Iliacus minor originates
two plexuses (3.1%). The slips originated from the trans- from the anterior inferior iliac spine, passes deep to ilio-
verse process of the fifth lumbar vertebra and iliolumbar psoas in front of the hip joint capsule, and attaches to the
ligament and split the femoral nerve into two branches. lower portion of the intertrochanteric line on the femur and/
or into the iliofemoral ligament (Macalister 1875; Knott
Anomalies 1883a; Das and Singh 1950; Bergman et al. 1988; Saga and
Description Takahashi 2016). Macalister (1875) notes that the insertion
Iliacus can be very small in cases of congenital absence of is “always above the main iliacus tendon” but Ward et al.
the femur (Manohar 1939). Itoh et al. (1991) describe fetal (2000) and Babst et al. (2011) state that the insertion of ilio-
akinesia/hypokinesia sequence in one male and one female capsularis is distal to the lesser trochanter.
infant, each with a suite of anatomical anomalies. The ilio- Das and Singh (1950) suggest that iliacus minor and ilio-
psoas muscle in the male contained irregular small fibers capsularis are separate muscles, the former inserting com-
while the muscle in the female showed focal atrophy. pletely into the intertrochanteric line and the latter inserting
completely into the iliofemoral ligament. These authors
Prevalence observed a case in which the muscle had a partial attach-
N/A ment into both structures (Das and Singh 1950). Smith-
Petersen (1949) considers iliacus minor to be the acetabular
Clinical Implications origin of iliacus (Ward et al. 2000).
A divided iliopsoas tendon may contribute to snapping at the
Innervation
hip joint (Tatu et al. 2002; Shu and Safran 2011; Philippon et al.
2014; Saga and Takahashi 2016). An accessory iliacus tendon Das and Singh (1950) note that iliacus minor is innervated
can mimic a tear of the iliopsoas tendon on MRI (Nguyen by the nerves to iliacus. Therefore, this muscle is likely
et al. 2013). The presence of accessory muscles or slips, such innervated by branches from the femoral nerve.
as iliacus minimus, may put tension on the femoral nerve that
Prevalence
can result in pain at the hip or knee joints or the L2-L4 derma-
tomes (Spratt et al. 1996). Accessory slips of iliacus can com- Ward et al. (2000) and Babst et al. (2011) suggest that ilio-
press the femoral nerve (Vázquez et al. 2007). Hypertrophy of capsularis is uniformly present in all humans but may be
iliacus and psoas major can distort the shape of the bladder and atrophied in stable hips and hypertrophied in dysplastic hips.
may be mistaken for pelvic lipomatosis (Chang 1978). Ward et al. (2000) found iliocapsularis in all 20 cadaveric
hips they examined, and Babst et al. (2011) found this muscle
in all 85 hips from the 82 patients included in their study.
Iliacus minor (Figure 4.3)
Anomalies
See also: Iliacus
Description
Synonyms Mieden (1982) describes an infant with median cleft lip,
This muscle may also be referred to as iliocapsularis hypotelorism, and alobar holoprosencephaly. Iliacus minor
(Harrison) (Macalister 1875; Bergman et al. 1988; Saga and was present bilaterally, extending between the iliac fossa and
Takahashi 2016), iliotrochantericus (Babst et al. 2011), ilio- the intertrochanteric line. Bersu et al. (1976) describe a male
capsulotrochanteric muscle (Cruveilhier), or iliacus brevis infant with Hanhart syndrome. The femora of this specimen
(Putti) (Ward et al. 2000). were normally developed but distal secondary ossification
centers were absent. The left leg stump had a patella and a
Typical Presentation small rudiment of the proximal tibia but no fibular rudiment.
This muscle may be only present as a variation or anomaly, The right leg stump was less developed and had a patella,
but see prevalence information below, as well as the discus- smaller tibial rudiment, and no fibular rudiment. On both
sions of Ward et al. (2000) and Babst et al. (2011). sides of the body, iliacus minor was present. On the right
Trunk Muscles 223
side, it extended from an origin it shared with rectus femoris major may be completely separate from iliacus (Macalister
to the intertrochanteric line. On the left side, it originated 1875; Bergman et al. 1988) or be completely fused with it
medial and inferior to rectus femoris and inserted onto the (Aleksandrova et al. 2013; Saga and Takahashi 2016). It
lesser trochanter with psoas major. may be divided into longitudinal bundles (Macalister 1875;
Bergman et al. 1988; Jelev et al. 2005; Saga and Takahashi
Prevalence 2016). It may receive a slip from the tendon of psoas minor,
N/A or some of its fbers may be continuous with the feshy
fbers of the diaphragm or the right crus (Macalister 1875;
Clinical Implications Rickenbacher et al. 1985). One or more accessory slips/mus-
In cases of hip dysplasia, the presence of iliocapsularis may cle bundles may be present medial or lateral to this muscle
help to stabilize the femoral head (Ward et al. 2000; Babst (Macalister 1875; Rickenbacher et al. 1985; Bergman et al.
et al. 2011). This muscle serves as an important landmark 1988; Jelev et al. 2005; Saga and Takahashi 2016). The ilio-
in hip surgery (Ward et al. 2000; Babst et al. 2011). Ward psoas tendon may be partially or completely divided into
et al. (2000) note that surgeons may fail to distinguish this one, two, or three tendons (Tatu et al. 2002; Polster et al.
muscle from rectus femoris and iliacus. 2008; Shu and Safran 2011; Crompton et al. 2014; Philippon
et al. 2014; Saga and Takahashi 2016).
Psoas major is associated with several named accessory
psOas MajOr (figure 4.3) slips and muscles. Jelev et al. (2005) describe an accessory
iliopsoas muscle that resulted from the connection between
Synonyms
an accessory iliacus muscle and an accessory psoas major.
This muscle can also be referred to as psoas magnus (Knott Psoas quartus (Figure 4.3) has been observed originating
1883b; Macalister 1875). from the medial aspect of quadratus lumborum and the
Typical Presentation transverse process of the ffth lumbar vertebra on the right
side of one cadaver, and via two slips from the transverse
Description processes of the fourth and ffth lumbar vertebrae on the
Psoas major arises from between the last thoracic and all left side (Clarkson and Rainy 1889). The muscle on both
lumbar vertebrae via fve digitations that attach to the ver- sides fused with the tendons of psoas major and psoas ter-
tebral bodies, transverse processes, and intervertebral discs tius. Tubbs et al. (2006c) note an origin from quadratus
(Standring 2016). Psoas major passes over the pelvic brim lumborum and the transverse process of the third lumbar
and forms the iliopsoas muscle with iliacus as the two insert vertebra. This muscle joins with iliacus and psoas or the
together via a tendon onto the lesser trochanter of the femur tendon of psoas major at the level of the inguinal ligament
(Standring 2016). (Tubbs et al. 2006c; Wong et al. 2019).
Clarkson and Rainy (1889) also describe the presence
Innervation of psoas tertius in the same cadaver, which originated from
Psoas major is primarily innervated by the frst two lumbar the inner aspect of rib twelve and the transverse processes
spinal nerves, with some supply from the third lumbar spi- of the upper four lumbar vertebrae on the right side, passed
nal nerve (Standring 2016). in front of quadratus lumborum and iliacus, and culmi-
nated in tendinous fbers that fused with the psoas major
Comparative Anatomy and psoas quartus tendons. On the left side of this cadaver,
Psoas major has a similar typical presentation in the apes, the muscle originated from the transverse processes of the
extending from the lumbar vertebrae and intervertebral third and fourth lumbar vertebrae and had the same inser-
discs to join with iliacus to insert onto the lesser trochan- tion. Khalid et al. (2017) note a psoas tertius with the same
ter (Champneys 1872; Hepburn 1892; Beddard 1893; Boyer origin as the right side of Clarkson and Rainy’s (1889) spec-
1935; Sigmon 1974; Gibbs 1999; Diogo et al. 2010, 2012, imen that pierced the femoral nerve and joined the tendon
2013a,b, 2017). The origin may extend to the twelfth tho- of iliopsoas.
racic vertebra or frst sacral vertebra in all species (Gibbs
1999; Diogo et al. 2010, 2012, 2013a,b, 2017). Fibers may
extend distally onto the femoral shaft in gibbons, orang- Prevalence
utans, and bonobos (Hepburn 1892; Beddard 1893; Boyer See iliacus entry for prevalence information regarding split-
1935; Miller 1952; Sigmon 1974; Diogo et al. 2012, 2013b, ting of the iliopsoas tendon and the presence of accessory
2017). slips associated with iliacus and psoas major. Astik and
Dave (2011) dissected 64 lumbar plexuses from 32 cadav-
Variations ers and found that psoas major split the femoral nerve into
Description medial and lateral slips in three plexuses (4.7%).
The vertebral attachments of psoas major may vary in num- Knott (1883b) examined psoas major bilaterally in 40
ber (Standring 2016) and the origin from the ffth lumbar subjects and noted that some fbers took origin from the
vertebra may be absent (Rickenbacher et al. 1985). Psoas neck of rib 12 in 3 subjects (7.5%). As this variant occurred
bilaterally in two subjects, it was found in 5 out of 80 sides psOas MinOr (figure 4.3)
(6.25%). In four cases (10%), fbers of origin came from the
right crus of the diaphragm, and in one case (2.5%) from Synonyms
the left crus. The origin from the last lumbar vertebra was This muscle can also be referred to as psoas parvus (Knott
absent in fve subjects (12.5%). As this variant occurred 1883b; Macalister 1875).
bilaterally in three subjects, this absence was noted in 8 out
of 80 sides (10%). Mori (1964) noted that the superiormost Typical Presentation
origin of psoas major was from the frst lumbar vertebra on Description
150 sides (98.7%) and from the second lumbar vertebra on Psoas minor originates from the bodies of the twelfth tho-
two sides (1.3%). This author did not observe an origin from racic and frst lumbar vertebrae, and their shared interverte-
the thoracic vertebrae (Mori 1964). bral disc, courses along the anterior surface of psoas major,
and inserts via a tendon onto the pecten pubis and iliopubic
Anomalies ramus (Standring 2016).
Description
Innervation
On the right side of an anencephalic male fetus, Windle
(1893) observed that psoas major was divided into two por- Psoas minor is innervated by the frst lumbar spinal nerve
tions (one being defcient) that shared an insertion. In a case (Standring 2016).
of congenital absence of the femur, Manohar (1939) reports
that psoas was very small, being well-developed near the Comparative Anatomy
vertebral column and inserting via a tendon onto the middle Psoas minor has a similar typical presentation in the apes,
of the iliac crest. Pirani et al. (1991) describe soft tissue running from the frst lumbar vertebra and often the last
anatomy associated with cases of proximal femoral focal thoracic vertebra to the iliopubic eminence (Champneys
defciency (PFFD). In Aitken type B PFFD, psoas major 1872; Hepburn 1892; Miller 1952; Swindler and Wood 1973;
is smaller than typical. Itoh et al. (1991) describe fetal aki- Sigmon 1974; Gibbs 1999; Diogo et al. 2010, 2012, 2013a,b,
nesia/hypokinesia sequence in one male and one female 2017). An origin from the second lumbar vertebra may be
infant, each with a suite of anatomical anomalies. The ilio- present in gibbons, orangutans, and common chimpanzees
psoas muscle in the male contained irregular small fbers and an origin from the eleventh thoracic vertebra may be
while the muscle in the female showed focal atrophy. present in gorillas (Kohlbrügge 1890–1892; Hepburn 1892;
Bersu et al. (1976) describe a male infant with Hanhart Gibbs 1999; Diogo et al. 2010, 2012, 2013a,b). In gibbons
syndrome. On both sides of this infant, there was an exten- and bonobos, the origin may extend to the third lumbar
sive fusion between the lumbar muscular components of vertebra (Kohlbrügge 1890–1892; Gibbs 1999; Diogo et al.
the diaphragm, psoas major, and quadratus lumborum. The 2012, 2017), and in gibbons alone, this muscle is often fused
femora of this specimen were normally developed but dis- with psoas major (Sigmon 1974; Diogo et al. 2012).
tal secondary ossifcation centers were absent. The left leg
stump had a patella and a small rudiment of the proximal Variations
tibia but no fbular rudiment. The right leg stump was less Description
developed and had a patella, smaller tibial rudiment, and no Psoas minor may be absent (Macalister 1875; Knott 1883b;
fbular rudiment. On both sides of the body, a small muscle Seib 1934; Mori 1964; Sato 1968c; Rickenbacher et al. 1985;
distinct from quadratus lumborum originated from the lat- Bergman et al. 1988; Hanson et al. 1999; Joshi et al. 2010;
eral aspect of psoas major and inserted onto the dorsal third Farias et al. 2012; Guerra et al. 2012; Gandhi et al. 2013;
of the iliac crest. Saga and Takahashi 2016; Standring 2016; Dragieva et al.
2018). It may arise from the frst lumbar vertebra only or
Prevalence the second lumbar vertebra and the disc above it (Bergman
N/A et al. 1988). It can have two heads or be doubled throughout
its length (Macalister 1875; Bergman et al. 1988; Gandhi
Clinical Implications et al. 2013; Saga and Takahashi 2016; Protas et al. 2017).
Hypertrophy of iliacus and psoas major can distort the Protas et al. (2017) describe a two-headed psoas minor
shape of the bladder and may be mistaken for pelvic lipo- with a lateral head that originated from the body of the frst
matosis (Chang 1978). Accessory slips of psoas major lumbar vertebra and a medial head that originated from the
can compress the femoral nerve (Vázquez et al. 2007). A bodies of the fourth and ffth lumbar vertebrae. The two
divided iliopsoas tendon may contribute to snapping at the heads joined at the level of the frst sacral vertebra and
hip joint (Tatu et al. 2002; Shu and Safran 2011; Philippon continued as a single tendon to insert onto the iliopectineal
et al. 2014; Saga and Takahashi 2016). eminence (Protas et al. 2017).
The presence of psoas quartus may split the femo- The psoas minor insertion tendon may also bifurcate
ral nerve and/or contribute to femoral nerve compression (Macalister 1875; Bergman et al. 1988). Its insertion may
(Tubbs et al. 2006c; Wong et al. 2019). vary, ranging from attachments to the inguinal ligament,
Trunk Muscles 225
iliac fascia, femoral neck, lesser trochanter, pectineal line bilaterally in a neonate with trisomy 18 (Aziz 1979). Among
of the femur, pectineal ligament, junction of the third and individuals with trisomy 21, psoas minor was absent on the
fourth lumbar vertebrae, or the junction of the ffth lum- left side of one child, bilaterally in another child, and bilat-
bar vertebra and the sacrum (Knott 1883b; Macalister 1875; erally in a fetus (Bersu 1980).
Bergman et al. 1988; Guerra et al. 2012; Gandhi et al. 2013;
Saga and Takahashi 2016). The ratio of tendinous to muscu- Prevalence
lar fbers may vary, and the entire muscle may be replaced In the literature review conducted by Smith et al. (2015),
by a tendon (Macalister 1875; Guerra et al. 2012; Gandhi the authors state that psoas minor was absent in 9 out of 17
et al. 2013; Saga and Takahashi 2016). Accessory psoas individuals with trisomy 18 (52.9%) and 3 out of 5 individu-
major fbers (“psoas accessorius”) may originate from the als with trisomy 21 (60%).
underside of the psoas minor tendon (Joshi et al. 2010; Saga
and Takahashi 2016). Clinical Implications
Gandhi et al. (2013) note that variations of psoas minor may
Prevalence be mistaken for retroperitoneal lymphadenopathy.
Knott (1883b) found this muscle present in 7 out of 40
subjects (17.5%). As this muscle was present bilaterally in PERINEAL, COCCYGEAL, AND
fve subjects, it was found in 12 out of 80 sides (15%). Seib ANAL MUSCULATURE
(1934) found that on 1000 sides from 500 cadavers, psoas
minor was present in 386 sides (38.6%). Summarizing other cOccygeus (figure 4.4)
research, Seib (1934) also notes that psoas minor is absent
Synonyms
in 57.3% of 5903 sides. Mori (1964) summarized the work
of other Japanese researchers and calculated an absence Coccygeus is sometimes considered to be the ischiococcy-
of psoas minor in 513 out of 958 sides (53.5%). Mori him- geal part of levator ani and is thus sometimes referred to as
self noted the absence of this muscle in 67 out of 100 sides ischiococcygeus (Standring 2016).
(67%). Sato (1968c) reported that psoas minor is absent in
163 out of 424 sides (38.4%) from Kyushu-Japanese males
and absent in 116 out of 256 sides (45.3%) in females. Description
Hanson et al. (1999) found that psoas minor was absent in Coccygeus is a musculotendinous sheet that extends from
3 out of 23 (13%) White male subjects and absent in 19 out the coccyx and the ffth sacral segment to the ischial spine
of 21 (91%) Black male subjects. In a study of 30 Brazilian (Liu and Salem 2016; Standring 2016).
cadaver pelves by Farias et al. (2012), psoas minor was
absent in 22 cases (73.33%). Innervation
In a study of 30 cadavers, Joshi et al. (2010) found that Coccygeus is innervated by branches of the sacral plexus
psoas minor was present in nine subjects (30%). As it was from the third and fourth sacral spinal segments (Standring
present on eight sides bilaterally and one side unilaterally, 2016).
psoas minor was present in 17 out of 60 sides (28.3%). In
4 of the 17 cases (23.5%), “psoas accessorius” was pres- Comparative Anatomy
ent (see description above) (Joshi et al. 2010). In a study of In orangutans, coccygeus is mostly replaced by the sacro-
22 fetuses (11 male, 11 female), Guerra et al. (2012) found spinous ligament (Diogo et al. 2013b). In gibbons, gorillas,
that psoas minor was present in 13 out of 22 fetuses overall and common chimpanzees, coccygeus is well-developed
(59%), being present in eight male fetuses (72.7%) on 15 but primarily tendinous (Elftman 1932; Diogo et al. 2010,
out of 22 sides (68.2%) and present in fve female fetuses 2012, 2013a). In common chimpanzees and bonobos, coc-
(45.5%) on 8 out of 22 sides (36.4%). These authors also cygeus is partially blended with levator ani (Elftman 1932;
noted that the tendon made up 60% of the muscle in females Miller 1952; Diogo et al. 2013a, 2017).
and 54% of the muscle in males (average of 57%) (Guerra
et al. 2012). Dragieva et al. (2018) found that psoas minor Variations
was present in six out of ten cadavers (60%). As the muscle Description
was present bilaterally in three subjects and unilaterally in Coccygeus may be entirely tendinous instead of muscular
three subjects, it was present in 9 out of 20 sides (45%). (Knott 1883b; Bergman et al. 1998; Liu and Salem 2016;
Standring 2016). When tendinous, it is fused with the sacro-
Anomalies spinous ligament (Standring 2016). It may only originate
Description from the sacrum (Macalister 1875). Coccygeus is rarely
absent (Knott 1883b; Bergman et al. 1988; Standring 2016).
In an anencephalic female fetus, Windle (1893) noted the
It may be doubled or tripled (Macalister 1875; Bergman
bilateral presence of an accessory muscle lying in front of
et al. 1998). Coccygeus is associated with two accessory
psoas major that occupied the position of psoas minor and
muscles, curvator coccygeus accessorius or sacrococcygeus
inserted onto the lesser trochanter. Psoas minor was absent
FIGURE 4.4 Pelvic diaphragm muscles in superior view.
anterior (von Luschka 1870; Bergman et al. 1988) (see the Comparative Anatomy
entry for this muscle), and a rectococcygeus muscle of Nair et al. (2011) state that sacrococcygeus anterior repre-
Treitz which originates from the anterior surface of the coc- sents the remnants of tail musculature found in lower ani-
cyx and partially fuses with the muscle coats of the rectum mals. Knott (1883b) notes that Krause suggests that this
(Liu and Salem 2016). muscle is homologous to fexor caudae. Flexor caudae is
present as a vestigial muscle in gorillas and extends from
Prevalence the last sacral vertebra to the middorsal fascia (Raven 1950;
Knott (1883b) found that coccygeus was absent in three Diogo et al. 2010). Remnants of fexor caudae have been
subjects he examined (no information on total sample size). reported in orangutans, but there is no trace of fexor cau-
One individual exhibited bilateral absence of feshy muscle dae in gibbons or chimpanzees (Elftman 1932; Gibbs 1999;
and replacement of this muscle with tendinous fbers. Diogo et al. 2012, 2013a,b, 2017).
N/A Description
Sacrococcygeus anterior is an accessory muscle that origi-
Clinical Implications nates from the anterior surface of the transverse process
As perineal hernias can emerge through defects in the peri- of one or two sacral vertebrae and inserts onto the coccyx
neal muscles, or in gaps between the muscles, congenital (von Luschka 1870; Watson 1880; Knott 1883b; Bergman
variations in muscular anatomy and any abnormal openings et al. 1988; Nair et al. 2011; Liu and Salem 2016). In the
in the muscles or fascia can contribute to the etiology of cases described by Watson (1880), the muscles arose in
perineal hernias (Trackler and Koehler 1968). between the third and fourth sacral foramina. Knott (1883b)
observed an origin from both the fourth and ffth sacral seg-
ments. In the case observed by Nair et al. (2011), the muscle
sacrOcOccygeus anteriOr (figure 4.4)
arose from the anterolateral surface of the sacrum at the
See also: Coccygeus, Sacrococcygeus posterior level of the third sacral vertebra. See the entry for sacrococ-
cygeus posterior for a similar muscle on the posterior aspect
Synonyms of the sacrum.
Sacrococcygeus anterior is also referred to as curvator coc- Niikura et al. (2010) found that this muscle is typi-
cyges (Watson 1880), curvator coccygeus accessorius or cally not present in adults. It appears during fetal week
sacrococcygeus anterior (von Luschka 1870; Bergman et al. 12, and between weeks 18 and 20, it increases in size and
1988), sacrococcygeus anticus (Knott 1883b), or sacrococ- fuses with the levator ani. Nair et al. (2011) conclude that
cygeus ventralis (Niikura et al. 2010). the presence of sacrococcygeus anterior in adult humans
indicates the failure of the fusion of the dorsal part of
Typical Presentation levator ani.
Trunk Muscles 227
Innervation In males, fbers inferolateral to the prostate are referred to

Sacrococcygeus anterior is innervated by a branch from the as puboprostaticus or levator prostate (Standring 2016; Liu
sacral plexus (Nair et al. 2011). and Salem 2016). In females, the fbers attach to the lat-
eral walls of the vagina and are referred to as pubovagi-
Prevalence nalis (Standring 2016; Liu and Salem 2016). Puborectalis
Knott (1883b) found feshy fbers corresponding to “sacro- is sometimes considered part of pubococcygeus (Standring
coccygeus anticus” in 2 out of 16 subjects (12.5%). 2016). It originates from the pubic bones and forms a mus-
cular sling that wraps around the anorectal junction, and
Anomalies some of these fbers may mix with those of the external anal
Description sphincter (Standring 2016; Liu and Salem 2016).
In a child with trisomy 21, Bersu (1980) found sacrococ-
cygeus anterior extending from the lower sacral vertebrae Innervation
to the coccyx. Iliococcygeus, pubococcygeus, and puborectalis are
innervated by branches of the sacral plexus from the
Prevalence third and fourth sacral spinal segments (Standring 2016).
In their literature review, Smith et al. (2015) found that Pubococcygeus may also be supplied by branches of the
sacrococcygeus anterior was present in one out of fve indi- pudendal nerve (Standring 2016).
viduals with trisomy 21 (20%).
Comparative Anatomy
Most portions of levator ani are missing and replaced with
Nair et al. (2011) suggest that fascia can fll the gap between fascia in gibbons, but pubococcygeus has been recorded
sacrococcygeus anterior and the dorsal part of levator ani, in some specimens (Elftman 1932; Diogo et al. 2012). The
creating an area that may be prone to prolapse of the pelvic levator ani of gorillas is largely similar to that in humans
organs. and iliococcygeus sometimes extends to the sacrum (Gibbs
1999; Diogo et al. 2010). In orangutans, most of the levator
levatOr ani (figure 4.4) ani as it appears in humans is present, but iliococcygeus is
usually aponeurotic and puborectalis is homologous with
Synonyms the inferior fbers of pubococcygeus (Elftman 1932; Diogo
Pubococcygeus may also be referred to as pubovisceralis et al. 2013b). The levator ani in common chimpanzees and
(Standring 2016; Liu and Salem 2016). bonobos is similar to that of orangutans (Elftman 1932;
Diogo et al. 2013a), but Diogo et al. (2017) did not record
Typical Presentation information for pubococcygeus in bonobos.
Description
Levator ani forms a large part of the pelvic foor and is com- Variations
prised of small muscles that are not easily differentiated Description
(Standring 2016). The muscles that comprise levator ani A slip named iliosacralis may arise from the posterior part of
vary considerably between individuals, and these muscles iliococcygeus (Standring 2016). Parts of pubococcygeus and
are described differently by different researchers (Bergman puborectalis can be variable in their attachments, while ilio-
et al. 1988). Levator ani is typically described as being coccygeus is typically not (Liu and Salem 2016). Levator ani is
comprised of iliococcygeus, pubococcygeus, and puborec- rarely absent. One absence was reported by Shepherd (1889),
talis, though some include ischiococcygeus (coccygeus, see where levator ani was replaced with pelvic fascia. Substantial
the entry above) as part of the levator ani (Standring 2016), variation in levator ani among women has been noted, and this
or puborectalis as part of pubococcygeus (Standring 2016). variation may be caused by childbirth and aging (Tunn et al.
Iliococcygeus attaches to the inner surface of the ischial 2003; Hoyte et al. 2004; Delancey et al. 2007).
spine and extends to the obturator fascia (Standring 2016).
It also attaches to the sacrum and coccyx and joins with Prevalence
fbers from the opposite side to form a raphe that is con- Tunn et al. (2003) used MRI to study the levator ani in
tinuous with the anococcygeal ligament (Standring 2016). females and found that this muscle did not attach to the
Pubococcygeus is the main part of levator ani (Liu and pubis in 4 out of 20 subjects (20%). Arakawa et al. (2004)
Salem 2016). It attaches to the posterior surface of the found that the connection between puboanalis and its inser-
pubis and its fbers run lateral to the urethra and the ure- tion was via a tight connection with smooth muscle in 37
thral sphincter along the pelvic foor (Standring 2016). out of 46 cadavers (80.4%) and via little or no tissue con-
Fibers that attach to the perineal body are referred to as nection in nine cadavers (19.6%). Tansatit et al. (2013) found
puboperinealis, and some fbers are sometimes referred to that puborectalis attached to the superior pubic ramus in six
as puboanalis as they insert between the external and inter- out of ten hemipelves (60%) and to the obturator internus
nal anal sphincters (Liu and Salem 2016; Standring 2016). fascia in four hemipelves (40%).
Anomalies sphincter ani externus (figure 4.5)

Hoyte et al. (2004) suggest that genetic defects are more N/A
likely to occur in iliococcygeus than other parts of leva-
tor ani. Anorectal malformations are common congenital Typical Presentation
anomalies that are often present with many other anoma-
Description
lies (e.g., cardiovascular, gastrointestinal, musculoskeletal,
spinal cord, and urogenital) and are associated with several The external anal sphincter is a circular muscle that sur-
syndromes (e.g., trisomies and other multisystemic condi- rounds the anus (Liu and Salem 2016; Standring 2016). It
tions) (Alamo et al. 2013). They can prevent the normal extends between the perineal body and the anococcygeal
appearance and function of levator ani and sphincter ani raphe (Liu and Salem 2016; Standring 2016). The deep part
externus until treated with surgery (Alamo et al. 2013). of the external anal sphincter blends with puborectalis (Liu
and Salem 2016; Standring 2016). Fibers from the transverse
Prevalence
perineal muscles and bulbospongiosus pass to the external
Anorectal malformations occur in about 1 out of 5,000 live anal sphincter (Peikert et al. 2015; Standring 2005, 2016).
births and are slightly more common in males (Alamo et al. The anterior portion of the muscle is shorter in females
2013). In 23 patients with anorectal anomalies studied by (Sultan et al. 1994).
Kohda et al. (1985), only ten patients (43.5%) had normal
development of the anorectal sphincter muscles (internal Innervation
anal sphincter, external anal sphincter, and levator ani), Sphincter ani externus is innervated by the inferior rec-
while four had intermediate development of these muscles tal branch of the pudendal nerve, the perineal nerve, or
(17.4%) and the remaining nine had poor development the anococcygeal nerve (Sato 1980; Liu and Salem 2016;
of these muscles (39.1%). Among 20 patients with Down Standring 2016).
syndrome that had anorectal malformations, Torres et al.
(1998) found that only one of them (5%) had poorly devel- Comparative Anatomy
oped perineal muscles (specifc muscles not indicated). Sphincter ani externus is similar in the apes, and the exten-
Clinical Implications sion of some fbers to bulbospongiosus has been observed
Weakness in levator ani and loss of ani volume can contrib- in gibbons, orangutans, and gorillas (Elftman 1932; Diogo
ute to incontinence, constipation, and pelvic organ prolapse et al. 2010, 2012, 2013a,b, 2017).
(Tunn et al. 2003; Hoyte et al. 2004; Delancey et al. 2007; Variations
Alamo et al. 2013). As perineal hernias can emerge through
defects in the perineal muscles, or in gaps between the Description
muscles, congenital variation in muscular anatomy and any Anterior muscle extensions of sphincter ani externus may
abnormal openings in the muscles or fascia can contribute occur to the corpus cavernosum or to the ischial tuberosity
to the etiology of perineal hernias (Trackler and Koehler (Sultan et al. 1994; Liu and Salem 2016; Tubbs and Watanabe
1968). The classifcations of anorectal malformations and 2016). Fibers from the superfcial surface of the sphincter
the surgical approaches to repair them are often depen- ani externus can pass to the medial aspect of ischiocaver-
dent upon their position relative to the levator ani muscle nosus via one or more bundles (Tubbs and Watanabe 2016).
(Nievelstein et al. 1998; Alamo et al. 2013). Slips may also pass from sphincter ani externus to the
FIGURE 4.5 Perineal muscles in inferior view.

Trunk Muscles 229
coccyx (Macalister 1875; Liu and Salem 2016). Retractor Prevalence

scroti refers to a bundle that extends from the superficial Anorectal malformations occur in about 1 out of 5,000 live
portion of sphincter ani externus to bulbospongiosus, or to births and are slightly more common in males (Alamo et al.
the base of the scrotum (Tubbs and Watanabe 2016). 2013). In 23 patients with anorectal anomalies studied by
Kohda et al. (1985), only ten patients (43.5%) had normal
Prevalence
development of the anorectal sphincter muscles (internal
Sultan et al. (1994) used endosonography to study the exter- anal sphincter, external anal sphincter, and levator ani),
nal anal sphincter in 93 nulliparous females and 21 males. while four had intermediate development of these muscles
The authors found that the deep (or proximal) portion of the (17.4%) and the remaining nine had poor development
external sphincter was annular in 72% of females and 76% of these muscles (39.1%). Among 20 patients with Down
of males, the superficial anal sphincter was elliptical in 76% syndrome that had anorectal malformations, Torres et al.
of females and 86% of males, and the subcutaneous portion (1998) found that only one of them (5%) had poorly devel-
was conical in 56% of females and 57% of males (Sultan oped perineal muscles (specific muscles not indicated).
et al. 1994). Anterior muscle extensions from this muscle However, the authors state that the sphincter mechanism in
occurred in 13% of females and 14% of males (Sultan et al. all 20 patients was normal (Torres et al. 1998).
1994).
Peikert et al. (2015) studied the relationship between Clinical Implications
bulbospongiosus and the external anal sphincter through Variations and anomalies in the development and presenta-
MRI of 43 male individuals and the study of six male tion of this muscle can affect continence (Alamo et al. 2013).
cadavers and classified the relationship into five variants.
Variant 1 was present in 2 out of 6 cadavers (~33%) and
14 out of 43 MRI patients (32.6%) and demonstrated a Transversus perinei profundus
bridge-like muscular connection between the two mus- (Deep transverse perinei) (Figure 4.5)
cles with connective tissue separating the muscles crani-
Synonyms
ally. Variant 2 was present in 2 out of 6 cadavers (~33%)
and 6 out of 43 MRI patients (~14%) and demonstrated N/A
direct contact between the two muscles. Variant 3 was
present in none of the cadavers (0%) and in 9 out of 43
MRI patients (~21%) and demonstrated ventral fibers of Description
the external anal sphincter reaching the bulbospongio- The deep transverse perinei comprise a sheet of muscle
sus muscle median raphe via connective tissue without that lies across the urogenital triangle and extends from
forming a muscular continuity. Variant 4 was present in the ischiopubic ramus (Standring 2005). The muscle sheet
1 out of 6 cadavers (~16.7%) and 7 out of 43 MRI patients attaches to the perineal body posteriorly, and its fibers are
(~16.3%) and demonstrated a combination of variants 1 deficient anteriorly (Standring 2005). Its fibers join with
and 2, or 2 and 3. Variant 5 was present in 1 out of 6 the fibers of the corresponding muscle on the opposite side
cadavers (~16.7%) and 7 out of 43 MRI patients (~16.3%) (Standring 2005). Some fibers extend to the deep part of the
and demonstrated no muscular or connective tissue con- external anal sphincter and the sphincter urethrae (Standring
nection between the bulbospongiosus and the external 2005).
anal sphincter (Peikert et al. 2015).
Innervation
Anomalies The deep transverse perinei are innervated by the perineal
Description branch of the pudendal nerve (Standring 2005).
Anorectal malformations are common congenital anoma-
Comparative Anatomy
lies that are often present with many other anomalies (e.g.,
cardiovascular, gastrointestinal, musculoskeletal, spinal There are no descriptions of the deep transverse perinei in
cord, and urogenital) and are associated with several syn- gibbons, orangutans, or bonobos (Diogo et al. 2012, 2013b,
dromes (e.g., trisomies and other multisystemic conditions) 2017). This muscle is present in gorillas and exhibits a pre-
(Alamo et al. 2013). They can prevent the normal appear- sentation similar to that in humans (Gibbs 1999; Diogo
ance and function of levator ani and sphincter ani externus et al. 2010). In common chimpanzees, Elftman (1932) sug-
until treated with surgery (Alamo et al. 2013). gested that some fibers of bulbospongiosus that insert into
In a case of a female infant with trisomy 9 studied by the ischium may be homologous with the deep transverse
Annerén and Sedin (1981), sphincter ani externus was perinei (Diogo et al. 2013a).
missing and the distance between the anus and vulva was
Variations
shortened. Bersu et al. (1976) describe a male infant with
Hanhart syndrome. In this infant, the external anal sphinc- Description
ter was comprised of two flat and parallel muscular bands The deep transverse perineal muscle frequently varies
that extended between the coccyx to the muscles of the among individuals (Bergman et al. 1988). This muscle may
penis with no decussating fibers. be absent, particularly in females (Knott 1883b; Bergman
et al. 1988; Tubbs and Watanabe 2016). An accessory mus- Prevalence

cle named transversus perinei alter may be present in front Knott (1883b) found transversus perinei superfcialis pres-
of this muscle (Tubbs and Watanabe 2016). ent only in 6 out of 30 subjects (20%). Knott (1883b) also
notes that transversus perinei medius—“the muscle more
Prevalence generally known as the t. p. superfcialis”—was absent in 8
Knott (1883b) found transversus perinei profundus absent out of 30 subjects (26.7%).
in 3 out of 30 bodies (10%).
Anomalies
Anomalies N/A
N/A
Clinical Implications As perineal hernias can emerge through defects in the peri-
As perineal hernias can emerge through defects in the perineal muscles, or in gaps between the muscles, congenital
neal muscles, or in gaps between the muscles, congenital variation in muscular anatomy and any abnormal openings
variation in muscular anatomy and any abnormal openings in the muscles or fascia can contribute to the etiology of
in the muscles or fascia can contribute to the etiology of perineal hernias (Trackler and Koehler 1968). Perineal her-
perineal hernias (Trackler and Koehler 1968). nias are defned based on their position (anterior or poste-
rior) relative to the superfcial transverse perineal muscle
transversus perinei superficialis (superficial (Trackler and Koehler 1968), so understanding potential
variations in this muscle is important for classifying these
transverse perinei) (figure 4.5)
hernias.
Synonyms
N/A sphincter urethrae (figure 4.5)
Typical Presentation Synonyms
Description N/A
The superfcial transverse perinei are muscular slips that
course transversely across the superfcial perineal space Typical Presentation
anterior to the anus (Standring 2016). The slips extend Description
between the ischial tuberosity and the perineal body The external urethral sphincter varies in presentation
(Standring 2016). The transverse perineal muscles send between males and females (Jung et al. 2012; Standring
fbers to bulbospongiosus and the external anal sphincter 2016). In males, the fbers inferior to the prostate compose
(Standring 2016). the external sphincter of the membranous urethra (Jung
et al. 2012). The muscle fbers originate from the junction
Innervation of the inferior pubic rami and ischium (Jung et al. 2012).
The superfcial transverse perinei are innervated by a In females, the urethral sphincter is found at the distal end
branch of the pudendal nerve (Standring 2016). of the bladder and is referred to as a urogenital sphincter
that has three components (Jung et al. 2012). The urogenital
Comparative Anatomy sphincter contains (1) the true sphincter that surrounds the
There are no descriptions of transversus perinei superfcia- urethra, (2) a compressor urethral muscle that arises from
lis in gibbons, orangutans, or bonobos (Diogo et al. 2012, the ischiopubic ramus and passes anterior to the urethra
2013b). This muscle is present in some gorillas and exhibits below sphincter urethrae, and (3) the urethrovaginal sphinc-
a presentation similar to that in humans (Gibbs 1999; Diogo ter, a muscle below compressor urethrae that extends from
et al. 2010). Sonntag (1923) states that this muscle is absent the ventral and lateral walls of the urethra to the vagina,
in common chimpanzees. up to the vestibular bulb (Oelrich 1983; Jung et al. 2012;
Standring 2016).
Variations
The superfcial transverse perinei may be reduced or Sphincter urethrae is innervated by a branch of the puden-
absent (Knott 1883b; Bergman et al. 1988; Standring dal nerve (Standring 2016).
2016; Tubbs and Watanabe 2016). It can consist of
many distinct fasciculi (Bergman et al. 1988; Tubbs and Comparative Anatomy
Watanabe 2016). It may connect with puborectalis or There are no descriptions of sphincter urethrae in gibbons,
the ischiopubic rami (Knott 1883b; Tubbs and Watanabe orangutans, or bonobos (Diogo et al. 2012, 2013b, 2017).
2016). It can also be doubled bilaterally (Tubbs and According to Gibbs (1999), this muscle is a true sphincter
Watanabe 2016). in male gorillas with no invasion of the prostate. Roberts
Trunk Muscles 231
and Seibold (1971) state that the fbers of this muscle extend Anomalies
to the vaginal wall in female chimpanzees, as it does in N/A
humans.
Variations N/A
Description
Bergman et al. (1988) state that there is frequent variation ischiOcavernOsus (figure 4.5)
in the muscles that compose the urethral sphincter, but do
not list any specifc variations. Variations of sphincter ure- Synonyms
thrae are not described in Liu and Salem (2016) or Tubbs N/A
and Watanabe (2016). Oelrich (1983) states that in females,
compressor urethrae may sometimes join the urethrovagi- Typical Presentation
nal sphincter near the anterior border of the vagina, instead Description
of joining it ventral to the urethra. In males, ischiocavernosus covers the crus penis and
attaches to the medial surface of the ischial tuberosity
Prevalence and to the ischial ramus (Standring 2016). The muscle is
similar in females and is associated with the crus of the
N/A
clitoris (Standring 2016). Ischiocavernosus is larger in
males (Bergman et al. 1988; Tubbs and Watanabe 2016).
Anomalies
N/A Innervation
Ischiocavernosus is innervated by a branch of the pudendal
Clinical Implications nerve (Standring 2016).
Variation or weakening of the external sphincter ure-
thrae can lead to urinary incontinence (Jung et al. 2012; Comparative Anatomy
Standring 2016). There are no descriptions of ischiocavernosus in gibbons,
orangutans, or bonobos (Diogo et al. 2012, 2013b, 2017).
This muscle is present in gorillas and exhibits a presenta-
ischiOpubicus (nOt illustrateD) tion similar to that in humans (Diogo et al. 2010). Sonntag
(1923) states that the origin of this muscle is from the
Synonyms ascending pubic ramus in common chimpanzees.
N/A
Variations
Description
Ischiocavernosus may receive fbers from sphincter ani
Comparative Anatomy externus (Tubbs and Watanabe 2016). Ischiocavernosus
is associated with two named muscular variations.
Bardeen (1921) states that ischiopubicus is homologous to
Compressor venae dorsalis refers to fasciculi that originate
the compressor of the dorsal vein of the penis or clitoris
from the sheath of the corpus cavernosum urethrae and the
found in lower mammals.
median raphe (Houston 1831; Bergman et al. 1988; Tubbs
Variations and Watanabe 2016). These fasciculi join with their counter-
parts on the other side of the body via a tendon that courses
Description
over the dorsal vein (Houston 1831; Bergman et al. 1988;
Ischiopubicus originates from the sides of the deep peri- Tubbs and Watanabe 2016). Pubocavernosus or levator penis
neal pouch and the inferior rami of the ischium and pubis refers to anterior bundles of ischiocavernosus that attach to
(Bardeen 1921; Tubbs and Watanabe 2016). It inserts the dorsal surface of the penis (Bergman et al. 1988; Tubbs
onto the inferior pubic ligament (arcuate ligament of and Watanabe 2016).
the pubis) (Oelrich 1983; Tubbs and Watanabe 2016).
This muscle is frequently absent in humans (Tubbs and Prevalence
Watanabe 2016). N/A
Innervation Anomalies
N/A N/A
Prevalence
N/A
Clinical Implications MRI of 43 male individuals and the study of six male
N/A cadavers and classifed the relationship into fve variants.
Variant 1 was present in 2 out of 6 cadavers (~33%) and
14 out of 43 MRI patients (32.6%) and demonstrated a
bulbOspOngiOsus (figure 4.5) bridge-like muscular connection between the two mus-
Synonyms cles with connective tissue separating the muscles crani-
ally. Variant 2 was present in 2 out of 6 cadavers (~33%)
This muscle is sometimes referred to as bulbocavernosus
and 6 out of 43 MRI patients (~14%) and demonstrated
(Tubbs and Watanabe 2016).
direct contact between the two muscles. Variant 3 was
Typical Presentation present in none of the cadavers (0%) and in 9 out of 43
MRI patients (~21%) and demonstrated ventral fbers of
Description
the external anal sphincter reaching the bulbospongio-
In males, bulbospongiosus is comprised of two parts joined sus muscle median raphe via connective tissue without
by a median fbrous raphe (Standring 2016). The fbers blend forming a muscular continuity. Variant 4 was present in 1
with the perineal body and are attached to the superfcial out of 6 cadavers (~16.7%) and 7 out of 43 MRI patients
transverse perineal muscle and the external anal sphinc- (~16.3%) and demonstrated a combination of variants 1
ter (Standring 2016). The middle fbers encircle the bulb and 2, or 2 and 3. Variant 5 was present in 1 out of 6
of the penis and corpus spongiosum (Standring 2016). The cadavers (~16.7%) and 7 out of 43 MRI patients (~16.3%)
anterior fbers extend over and insert partly into the sides and demonstrated no muscular or connective tissue con-
of the corpus cavernosum, and also partially insert into a nection between the bulbospongiosus and the external
tendinous expansion over the dorsal vessels of the penis anal sphincter. In this last case, the origin of the bulbo-
(Standring 2016). In females, bulbospongiosus attaches to spongiosus muscle was in the dense connective tissue of
the perineal body but the muscle on each side of the body the body of the perineum (Peikert et al. 2015).
is separate (Standring 2016). It covers the superfcial por-
tions of the vestibular bulbs and greater vestibular glands
(Standring 2016). It attaches onto the corpus cavernosum Anomalies
clitoridis (Standring 2016). N/A
Innervation
Bulbospongisosus is innervated by a branch of the puden-
N/A
dal nerve (Standring 2016).
Comparative Anatomy
Bulbospongiosus has a similar typical presentation in the THORAX, SPINE, AND BACK MUSCLES
apes (Elftman 1932; Diogo et al. 2010, 2012, 2013a,b). DiaphragM (figure 4.6)
There is a connection to the ischium in gibbons, gorillas,
and common chimpanzees (Elftman 1932; Diogo et al. Synonyms
2010, 2012). There is no data on this muscle for bonobos N/A
(Diogo et al. 2017).
Variations
Description
Description
The diaphragm is a continuous musculofbrous sheet that
The posterior part of bulbospongiosus may be separate
has three portions (Standring 2016). The sternal portion
from the anterior part, or is absent altogether, particu-
originates from the xiphoid process (Standring 2016). The
larly in females (Tubbs and Watanabe 2016). Fibers that
costal portion originates from the lower six costal car-
extend from the ischial tuberosity to bulbospongiosus
tilages and their corresponding ribs on each side of the
may be referred to as ischiobulbosi (Tubbs and Watanabe
thorax (Bergman et al. 1988; Standring 2016). The lum-
2016). In males, gluteoperinealis (Krause) refers to a slip
bar portion originates from the medial and lateral arcuate
that originates from the gluteus maximus fascia or upper
ligaments and from the frst three lumbar vertebrae via left
fascia lata and inserts beneath the origin of bulbocaver-
and right crura (Standring 2016). The muscle fbers con-
nosus in the albuginea of the penile urethra (Knott 1883b;
verge into a central tendon near the center of the muscle
Bergman et al. 1988; Nicholson et al. 2016; Tubbs and
(Standring 2016).
Watanabe 2016).
Peikert et al. (2015) studied the relationship between The diaphragm is innervated by the phrenic nerves (Standring
bulbospongiosus and the external anal sphincter through 2016).
Trunk Muscles 233
FIGURE 4.6 Anterior thoracic wall muscles and diaphragm in anterior view.
Comparative Anatomy Bergman et al. 1988). A slip may connect the right crus
The diaphragm has a similar typical presentation in the apes to the duodenum, and a fbrous band can extend from the
(Gibbs 1999; Diogo et al. 2010, 2012, 2013a,b, 2017). The medial borders of the crura to the surface of the esopha-
costal portion originates from the lower seven ribs in goril- gus (Bergman et al. 1988).
las, common chimpanzees, and bonobos (Sonntag 1923; The diaphragm is associated with a few named acces-
Miller 1952; Diogo et al. 2010, 2013a, 2017). Accessory sory muscles. As described by Allen (1950), transpleura-
slips may be present near the lumbar portion in common lis subclavius presents as two bands of muscle that pass
chimpanzees (Sonntag 1923). The sternal portion origi- through the oblique fssure of the right lung. The bands
nates from the xiphoid process in bonobos and from the originate from the sixth rib and attach onto the central ten-
sternum in common chimpanzees (Sonntag 1923; Miller don of the diaphragm, just behind the caval opening (Allen
1952; Diogo et al. 2013a, 2017). 1950; Bergman et al. 1988; Snosek and Loukas 2016).
Diaphragmaticoretromediastinalis (Eppinger) extends
from the superior end of the medial crus to the posterior
Variations mediastinum (Rickenbacher et al. 1985). See the entry for
Description hepatodiaphragmaticus.
The diaphragm is variable in size, position, thickness,
ratio of tendon to muscle, and in its attachments (Knott Prevalence
1883b; Snosek and Loukas 2016; Standring 2016). The In a sample of 50 diaphragms, Botros et al. (1990) found
sternal portion of the muscle varies in attachments and that the right crus divided into a medial, middle, and lat-
may be absent (Bergman et al. 1988; Standring 2016). eral bundle in 98% of cases. They also found that the left
The costal portion can potentially extend to the ffth or crus only had two bundles, a medial and lateral bundle,
sixth rib, and the slip to the twelfth rib may not be pres- in 90% of cases (Botros et al. 1990). Yeh et al. (1990)
ent (Macalister 1875; Bergman et al. 1988). The crura describe variant anatomy of the diaphragm that was
may be divided into separate bundles (Botros et al. 1990; detected with abdominal ultrasonography in 74 patients.
Snosek and Loukas 2016). Fibers from the diaphragm can Among other abnormalities, 34 patients (45.9%) had a
occasionally extend to psoas major or quadratus lumbo- diaphragmatic slip, 10 patients (13.5%) exhibited scal-
rum (Macalister 1875; Bergman et al. 1988). Accessory loping of the diaphragm that was associated with slips,
muscle slips may be found, often near the central tendon and 3 patients (4%) had hypertrophic crura (Yeh et al.
(Macalister 1875; Knott 1883b; Rickenbacher et al. 1985; 1990).

Description This muscle is only present as a variation or anomaly.
The diaphragm may be absent as an anomaly (Macalister
1875), particularly in cases of anencephaly (Rickenbacher et Comparative Anatomy
al. 1985). Congenital or age-related defects of the diaphragm N/A
can lead to thinning or fssure of the musculofbrous sheet
(Macalister 1875; Snosek and Loukas 2016). Accessory dia- Variations
phragms may be present (Hollinshead 1956; Wille et al. 1975; Description
da Silva et al. 2012; Snosek and Loukas 2016). Anomalous Hepatodiaphragmaticus is a muscular band that originates
presentations of the diaphragm can also involve the presence from the central tendon anterior to the esophagus and
of slips of muscle fbers to other structures (Smith et al. 2015). extends to the hilum of the liver (Knott 1883b; Macalister
Bersu et al. (1976) describe a male infant with Hanhart 1875; Von der hellen 1903; Bergman et al. 1988).
syndrome. On both sides of this infant, there was an exten-
sive fusion between the lumbar muscular components of the Innervation
diaphragm, psoas major, and quadratus lumborum. Itoh et N/A
al. (1991) describe fetal akinesia/hypokinesia sequence in
one male and one female infant, each with a suite of ana- Prevalence
tomical anomalies. The diaphragm in both infants showed Knott (1883b) found “hepatico-diaphragmaticus” in 1 out
focal atrophy. of 36 cases (2.8%).
Anomalies
In their literature review, Smith et al. (2015) found several In a neonate with trisomy 13, Pettersen et al. (1979) found
anomalous slips of muscle originating from the diaphragm that “hepaticodiaphragmaticus” extended as a band of stri-
in 1 out of 20 individuals with trisomy 13 (5%). These slips ated muscle from the undersurface of the diaphragm, into the
include a slip from the right crus to psoas major, a slip pres- gastrohepatic ligament and then into the hilum of the liver.
ent on the central tendon, and a slip from the central tendon
to the pericardium just below the left inferior pulmonary Prevalence
vein (Smith et al. 2015). In the study and literature review According to the literature review completed by Smith et al.
completed by Wille et al. (1975), they found that out of 21 (2015), this muscle was present in 1 out of 20 individuals
cases, accessory diaphragms were more frequent on the with trisomy 13 (5%).
right side (20 cases) than the left side (1 case) and also more
common in males (13 cases) than in females (8 cases). Clinical Implications
N/A
Bergman et al. (1988) state that detached or aberrant transversus thOracis (figure 4.6)
muscle slips from the diaphragm can compress the renal
artery. Accessory slips of the diaphragm may be mistaken Synonyms
for intrahepatic masses (Yeh et al. 1990). Congenital or This muscle is also referred to as sternocostalis or trian-
age-related defects of the diaphragm can lead to hernia- gularis sternae (Snosek and Loukas 2016; Standring 2016),
tion of abdominal organs into the thorax (Snosek and infra-costalis anterior, or transversus thoracis anterior
Loukas 2016). Age-related changes of the diaphragm are (Macalister 1875).
strongly associated with emphysema (Caskey et al. 1989).
An accessory diaphragm can lead to pulmonary malde-
velopment and some associated symptoms are respiratory Description
distress, recurrent respiratory infection, and/or chronic Transversus thoracis is located on the inner surface of the
pulmonary infammation (Wille et al. 1975). anterior thoracic wall (Standring 2016). It originates from
the sternum, xiphoid process, and the sternal ends of the
costal cartilages of ribs four through seven (Standring
hepatODiaphragMaticus (nOt illustrateD) 2016). The muscle fbers separate into slips that typically
attach to the costal cartilages of ribs two through six
See also: Diaphragm
(Standring 2016).
Synonyms Innervation
This muscle may also be referred to as hepaticodiaphrag- Transversus thoracis is innervated by adjacent intercostal
maticus (Knox) (Macalister 1875). nerves (Standring 2016).
Trunk Muscles 235
Comparative Anatomy 3 to 6 in 121 cases (39%) and into fve slips from rib 2 to 6
In gorillas, transversus thoracis has been observed to run in 86 cases (27.7%).
between ribs one and eight (Raven 1950; Diogo et al. 2010). Jelev et al. (2011) studied transversus thoracis on 240
In common chimpanzees, it runs between the second and sides from 120 cadavers and found that the most common
sixth ribs (Diogo et al. 2013a). In bonobos, it runs between spread of this muscle was between ribs two and six, and
the second and seventh ribs (Miller 1952; Diogo et al. 2017). that there was asymmetry between the left and right side in
Gibbs (1999) and Diogo et al. (2012, 2013b) have stated 44.2% of the cadavers. The superior most costal cartilage/
that no information is available for gibbons or orangutans. rib of origin was rib 1 in 7.5% of left sides and 0.8% of
However, Satoh (1971) has stated that the superior most cos- right sides, rib 2 in 53.3% of left sides and 37.5% of right
tal attachment is to the second or third rib in the gibbon, sides, rib 3 in 29.2% of left sides and 46.7% of right sides,
and to the third rib in the orangutan. rib 4 in 9.2% of left sides and 14.2% of right sides, and rib
5 in 0.8% of left sides and 0.8% of right sides. The inferior
Variations most costal cartilage/rib of origin was rib 5 in 0.8% of left
Description sides and 1.7% of right sides, rib 6 in 94.2% of left sides and
The muscle attachments can occasionally extend up to the 89.2% of right sides, and rib 7 in 5% of left sides and 9.2%
frst rib or down to the seventh rib (Mori 1964; Satoh 1971; of right sides.
Bergman et al. 1988; Jelev et al. 2011; Snosek and Loukas Anomalies
2016). Attachments can vary between each side of the body
N/A
within an individual (Macalister 1875; Jelev et al. 2011;
Standring 2016). Transversus thoracis may be continuous Clinical Implications
with transversus abdominis (Macalister 1875). Macalister
Jelev et al. (2011) note that understanding the variation in
(1875) notes that this muscle may be absent.
attachments of transversus thoracis is important due to the
Prevalence close anatomical relationship of this muscle to the internal
thoracic artery, which is an important vessel for coronary
Mori (1964) studied the attachments of transversus thoracis
artery bypass grafting (CABG) surgery.
on 310 sides. The superior most costal cartilage/rib of ori-
gin was rib 1 on 7 sides (2.2%), rib 2 on 122 sides (39.3%),
rib 3 on 155 sides (50%), rib 4 on 23 sides (7.2%), and rib intercOstales externi (external
5 on 3 sides (0.9%). The inferior most costal cartilage/rib intercOstals) (figure 4.7)
of origin was rib 5 on 50 sides (16.1%), rib 6 on 237 sides
(76.5%), and rib 7 on 23 sides (7.4%). The most common Synonyms
morphs of this muscle were separation into 4 slips from rib N/A
FIGURE 4.7 Intercostal and subcostal muscles in anterior view.

Typical Presentation Intercostales interni (Internal

Description intercostals) (Figure 4.7)
Each external intercostal muscle extends from the lower Synonyms
margin of one rib to the superior margin of the rib below
N/A
it (Standring 2016). Together, the 11 pairs of external
intercostals pass from the tubercles of the ribs nearly to
the costal cartilages and continue toward the sternum as Typical Presentation
the external intercostal membrane (Standring 2016). Description
Each internal intercostal muscle extends from the inner
Innervation margin of a costal groove and attaches onto the superior
The external intercostal muscles are innervated by the cor- margin of the rib below (Standring 2016). Together, the 11
responding intercostal nerves (Standring 2016). pairs of internal intercostals commence near the sternum
and continue to the posterior costal angles as the internal
intercostal membrane (Standring 2016).
Comparative Anatomy
The external intercostals have a similar typical presenta-
tion in the apes, attaching between the adjacent margins Innervation
of each pair of ribs (Diogo et al. 2010, 2012, 2013a,b, The internal intercostal muscles are innervated by the cor-
2017). Sonntag (1923) states that in common chimpan- responding intercostal nerves (Standring 2016).
zees, the first three and last two external intercostal mus-
cles are fleshy, but the rest are replaced by membranous Comparative Anatomy
tissue. In one of the common chimpanzees examined by
The internal intercostals have a similar typical presentation
Diogo et al. (2013a), the external intercostals had a typi-
in the apes, attaching between the adjacent margins of each
cal presentation.
pair of ribs (Diogo et al. 2010, 2012, 2013a,b, 2017).
One or more external intercostal muscles, particularly the last The internal intercostals may vary in their distance between
muscle, may be replaced by a sheet of connective tissue or their dorsal borders and the costal angle (Rickenbacher
may be completely absent (Macalister 1875; Rickenbacher et al. 1985). These muscles may also show discontinuities as
et al. 1985). Extra muscle bellies may be present when there they thin out on the dorsal part of the thorax (Rickenbacher
are extra ribs (Macalister 1875; Rickenbacher et al. 1985). et al. 1985). Macalister (1875) notes that the internal inter-
Fibers from the external intercostal muscles may blend with costal muscles may join with transversus thoracis (“infra-
the internal intercostal muscles, serratus anterior, and/or the costalis”). Some of the muscles in the eighth through
external oblique (Macalister 1875; Rickenbacher et al. 1985). twelfth intercostal spaces may be continuous with the inter-
nal oblique (Macalister 1875).
N/A N/A
Anomalies Anomalies
Itoh et al. (1991) describe fetal akinesia/hypokinesia Itoh et al. (1991) describe fetal akinesia/hypokinesia
sequence in one male and one female infant, each with a sequence in one male and one female infant, each with a
suite of anatomical anomalies. In the male, the intercostal suite of anatomical anomalies. In the male, the intercostal
muscles were slightly atrophic. Paria et al. (2015) describe a muscles were slightly atrophic. Paria et al. (2015) describe a
case of an infant with congenital absence of the anterior and case of an infant with congenital absence of the anterior and
anterolateral parts of multiple lower ribs and absence of the anterolateral parts of multiple lower ribs and absence of the
adjacent intercostal muscles. adjacent intercostal muscles.
N/A N/A
Clinical Implications Clinical Implications

N/A N/A
Trunk Muscles 237
intercOstales intiMi (innerMOst Typical Presentation

intercOstals) (figure 4.7) Description
Synonyms The subcostal muscles are typically only well-developed
in the lower part of the posterior thorax (Standring 2016).
N/A
Each subcostal muscle originates from the inner surface of
Typical Presentation one rib and crosses one or two intercostal spaces to insert
onto the inner surface of the second or third rib below its
Description rib of origin (Rickenbacher et al. 1985; Standring 2016).
The innermost intercostal muscles are deep to the internal
intercostal muscles and are attached to the internal surfaces Innervation
of two adjacent ribs (Standring 2016). Subcostales muscles are innervated by the corresponding
intercostal nerves (Standring 2016).
Innervation
Comparative Anatomy
The innermost intercostal muscles are innervated by the
corresponding intercostal nerves (Standring 2016). Raven (1950) lists subcostales as absent in the gorilla he
dissected. There is no information on this muscle for the
Comparative Anatomy other apes (Gibbs 1999; Diogo et al. 2010, 2011, 2013a,b,
2017). Satoh (1974) found three bundles of subcostales in 1
There is no information on the innermost intercostals in the
out of 30 macaques (3.3%) and stated that subcostales were
apes (Gibbs 1999), and it is likely that their morphology has
absent in the other specimens. Satoh (1974) concludes that
been characterized along with the internal intercostals due
subcostales are typically absent in non-human primates.
to the close anatomical association between these muscles.
Variations
Variations
Description
Description The subcostales are often variable in their number and extent
The innermost intercostals that occupy the frst three inter- of development (Rickenbacher et al. 1985; Standring 2016).
costal spaces may sometimes extend as far as the vertebral Complete absence of subcostales is rare (Rickenbacher et al.
column (Rickenbacher et al. 1985). Standring (2016) notes 1985). Rickenbacher et al. (1985) state that the bundles of
that the superiormost muscles may be reduced or absent. subcostales that are most constant are those between the sec-
The innermost intercostals may join with the subcostal ond and fourth ribs and those between the ninth and twelfth
muscles posteriorly (Standring 2016). ribs. Satoh (1974) states that the most constant bundles of
subcostales are those between the third and ffth ribs, the
Prevalence ninth and eleventh ribs, and the tenth and twelfth ribs.
N/A The lowest subcostal muscle—which attaches onto the
internal surface and superior border of the twelfth rib—
Anomalies may also have an attachment onto the body of the twelfth
Description thoracic vertebra (Rickenbacher et al. 1985) or the trans-
Itoh et al. (1991) describe fetal akinesia/hypokinesia verse process of the frst lumbar vertebra (Satoh 1974).
sequence in one male and one female infant, each with a Satoh (1974) observed that some subcostal muscles radiated
suite of anatomical anomalies. In the male, the intercostal over the aponeurosis of transversus abdominis or iliopsoas
muscles were slightly atrophic. Paria et al. (2015) describe a instead of inserting onto the rib.
case of an infant with congenital absence of the anterior and Satoh (1974) also categorized bundles of subcostales into
anterolateral parts of multiple lower ribs and absence of the six types. In type A, the bundle of subcostales crossed over
adjacent intercostal muscles. one rib and ended in the next intercostal space without reach-
ing the next rib. In type B, the muscle crossed two intercostal
Prevalence spaces to insert onto the second rib below its rib of origin.
N/A In type C, the muscle spanned 2.5 intercostal spaces. In type
D, the muscle spanned three intercostal spaces and inserted
Clinical Implications onto the third rib below its rib of origin. In type E, the muscle
N/A radiated into the aponeurosis of transversus abdominis or ilio-
psoas. In type F, the muscle inserted onto the transverse pro-
subcOstales (subcOstal Muscles) (figure 4.7) cess of the last thoracic vertebra or the frst lumbar vertebra.
Synonyms Prevalence
This muscle may also be referred to as infracostales, intra- In a sample of 23 Japanese male cadavers, Satoh (1974)
costalis, transversus thoracis posterior, or serratus internus found subcostales present on both sides of the body in
(Macalister 1875). all cases (100%) The number of subcostales muscles per
person ranged from 1 to 14, with an average of fve. Satoh’s muscle was at the frst rib on 105 sides (~4.4%) and at the
(1974) type A was found in 9 out of 276 cases (3.3%), type second rib on 21 sides (~0.9%). This muscle inserted onto
B in 216 cases (78.3%), type C in fve cases (1.8%), type D the third rib on 58 sides (~2.4%) and on the fourth rib on
in 25 cases (9%), type E in ten cases (3.6%), and type F in 68 sides (~2.8%). Tachibana and Miyauchi (1989) report a
11 cases (4%). similar prevalence, noting the presence of this muscle in 14
out of 316 sides (4.4%). Miyauchi (1982b) found supracos-
Anomalies talis anterior in 3 out of 50 sides (6%) from 25 cadavers.
N/A Nelson et al. (1992) found this muscle only once out of 317
cadavers (0.3%).
N/A Anomalies
Description
supracOstalis anteriOr (figure 4.6) In individuals with trisomy 13 and trisomy 21, supracosta-
lis anterior extends from the frst rib to the second, third,
Synonyms or fourth rib (Pettersen 1979; Pettersen et al. 1979; Bersu
This muscle may be referred to as supracostalis anterior 1980; Smith et al. 2015).
anomalous (Bochdalek 1867), supra-costalis superfcialis
(Knott 1883a), or obliques externus thoracis (Keith 1894). Prevalence
Typical Presentation In their literature review, Smith et al. (2015) found that
supracostalis anterior was present in 2 out of 20 individu-
als with trisomy 13 (10%) and 1 out of 5 individuals with
Comparative Anatomy trisomy 21 (20%).
N/A Clinical Implications
N/A
Variations
Description
Supracostalis anterior may be present unilaterally or bilat- supracOstalis pOsteriOr (figure 4.8)
erally (Macalister 1875; Nelson et al. 1992; Bergman et al. Synonyms
1988). When present, supracostalis anterior is situated deep
This muscle is also referred to as supracostalis dorsalis
to the pectoral musculature, typically originating from one
(Mori 1964).
rib and extending across two or more anterior intercostal
spaces to insert onto another rib (Wood 1865; Macalister Typical Presentation
1875; Knott 1883a; Eisler 1912; Nelson et al. 1992; Bakkum
and Miller 2016; Snosek and Loukas 2016). One or more
bundles may be present (Bergman et al. 1988). It may attach Comparative Anatomy
onto ribs one through four, the deep fascia of the neck, or
N/A
the scalene muscles (Macalister 1875; Mori 1964, Bergman
et al. 1988; Nelson et al. 1992; Bakkum and Miller 2016;
Snosek and Loukas 2016). Some argue that it is derived Variations
from remnants of external abdominal oblique (Keith 1894; Description
Bergman et al. 1988; Bakkum and Miller 2016) while Supracostalis posterior may be present unilaterally
others suggest that it arises from the intercostal muscles or bilaterally (Mori 1964). Like its counterpart on the
(Tachibana and Miyauchi 1989; Nelson et al. 1992; Snosek anterior aspect of the thorax, supracostalis posterior
and Loukas 2016). originates from one rib and extends across two or more
intercostal spaces to insert onto another rib (Eisler 1912;
Innervation Mori 1964; Rickenbacher et al. 1985; Bakkum and Miller
This muscle has been observed to be innervated by the 2016). One or more bundles may be present (Mori 1964).
external muscular branch of the frst intercostal nerve Eisler (1912) notes that bundles of supracostalis posterior
(Tachibana and Miyauchi 1989), as well as a collateral are situated medial to the origin of serratus anterior. Mori
branch of the second intercostal nerve (Nelson et al. 1992). (1964) provides a review of the three types of supracosta-
lis posterior that Eisler classifed. Type I originates from
Prevalence one rib and crosses one or two ribs to insert into another
Mori (1964) observed that supracostalis anterior was present rib, and its fbers are oriented in the same direction as
in 126 cases out of 2,400 sides from 1,200 cadavers (~5.3%). the external intercostal muscles. Type II originates from
Mori (1964) also noted that the superior most origin of this a rib via one to three feshy bundles, and its fbers travel
Trunk Muscles 239
FIGURE 4.8 Posterior thoracic wall muscles in posterior view.
superomedially to insert onto another rib. Fibers of type Clinical Implications

II muscles are oriented in the same direction as serra- N/A
tus posterior superior. Type III originate from the eighth,
ninth, and tenth ribs and travel superiorly to insert into
either the fascia of the ribs or into the fascia of the erec- serratus pOsteriOr superiOr (figure 4.8)
tor spinae. Synonyms
Innervation This muscle is also referred to as serratus posticus superior
(Macalister 1875).
Bundles of supracostalis posterior are likely innervated by
the intercostal nerves. Typical Presentation
Mori (1964) observed 65 cases of supracostalis dorsalis and Serratus posterior superior originates from the nuchal liga-
found that type I was present on 19 sides (3.6%), type II ment and the spines of the last cervical and upper two or
was present on 17 sides (3.3%), and type III was present on three thoracic vertebrae (Standring 2016). It inserts via four
29 sides (5.9%). Using these numbers to calculate the total digitations onto the superior margins of the second through
sample size, it is likely that Mori (1964) observed supracos- ffth ribs (Standring 2016).
talis posterior in 65 out of around 520–530 sides (about 12%
Innervation
or 12.5%) (Bakkum and Miller 2016).
Serratus posterior superior is innervated by the second
Anomalies through ffth intercostal nerves (Standring 2016).
Description
Comparative Anatomy
While supracostalis anterior has been observed in individu-
Serratus posterior superior originates from the fourth cervi-
als with trisomy 13 and trisomy 21 (Pettersen 1979; Pettersen
cal vertebra through the frst thoracic vertebra and inserts on
et al. 1979; Bersu 1980; Smith et al. 2015), supracostalis pos-
the second, third, and fourth ribs in gibbons (Diogo et al.
terior has not been observed in individuals with trisomy, birth
2012). This muscle is absent or replaced by tendinous threads
defects, or other congenital disorders, to our knowledge.
in orangutans (Gibbs 1999; Diogo et al. 2013b). In gorillas,
Prevalence the muscle extends from all seven cervical vertebrae to ribs
two through six (Diogo et al. 2010). In common chimpan-
N/A
zees, the muscle extends from the seventh cervical and frst
thoracic vertebrae to ribs two through fve, and occasionally It inserted most frequently onto ribs two through fve
to rib one, which is the closest to the typical human condition on nine sides (45%). Other insertions included the second
(Sonntag 1923; Diogo et al. 2013a). In bonobos, the origin is through fourth ribs on fve sides (25%), the second through
variable and can extend superiorly as far as the third cervical sixth ribs on four sides (20%), and second through seventh
vertebra, and its insertion is onto ribs two through four or two ribs on one side (5%), and the third through sixth ribs on one
through fve (Miller 1952; Diogo et al. 2017). side (5%). Therefore, the upper limit of the insertion was the
second rib on 19 sides (95%) and the third rib on one side
Variations (5%). The lower limit of the insertion was rib fve in nine
Description cases (45%), rib four in fve cases (25%), rib six in fve cases
(25%), and rib seven in one case (5%) (Satoh 1969).
Serratus posterior superior may be absent (Macalister 1875;
Loukas et al. (2008c) studied serratus posterior superior
Standring 2016), and if so, the muscle is replaced by fbrous
on 100 sides of 50 cadavers. They found that the origin of
tissue (Bergman et al. 1988; Bakkum and Miller 2016). The
the muscle was from the seventh cervical through second or
number of digitations may vary, ranging from three to six
third thoracic vertebrae in 65% of specimens, from the sixth
(Macalister 1875; Bergman et al. 1988; Bakkum and Miller
cervical through the second thoracic vertebrae in 30% of
2016; Standring 2016), or even only two digitations (Mori
specimens, and from the ffth cervical through the second
1964). Mori (1964) notes that the origin may extend superi-
or third thoracic vertebrae in 5% of specimens. It inserted
orly up to the fourth, ffth, or sixth cervical vertebra. Satoh
onto the second through ffth ribs in 90% of specimens and
(1969) observed an origin as high as the third cervical ver-
into the second through sixth ribs in 10% of cases.
tebra. Macalister (1875) states that rhomboatloideus may
originate from serratus posterior superior.
Anomalies
Mori (1964) studied the origin and insertion of serratus Serratus posterior superior may be absent as an anomaly
posterior superior in Japanese cadavers. The most com- (Colacino and Pettersen 1978; Smith et al. 2015). In a fetus
mon origin was from between the ffth cervical vertebra with spina bifda with encephaloceles, the serratus posterior
and the frst thoracic vertebra on 41 out of 99 sides (41%). muscles were unidentifable (Wheeler 1918).
Other scopes of origin include from the fourth cervical
Prevalence
vertebra to the frst thoracic vertebra on 33 sides (33%),
the ffth cervical vertebra to the seventh cervical vertebra In their literature review, Smith et al. (2015) found that
on 12 sides (12%), the fourth cervical vertebra to the sev- serratus posterior superior was absent in 1 out of 24 cases
enth cervical vertebra on 4 sides (4%), the ffth cervical of trisomy 13 (4.2%)—a case described by Colacino and
vertebra to the second thoracic vertebra on 4 sides (4%), Pettersen (1978).
the sixth cervical vertebra to the frst thoracic vertebra Clinical Implications
on 3 sides (3%), and the fourth cervical vertebra to the
N/A
second thoracic vertebra on 2 sides (2%). Therefore, the
upper limit of the origin of this muscle is the fourth cervi-
cal vertebra on 42 sides (42%), the ffth cervical vertebra serratus pOsteriOr inferiOr (figure 4.8)
on 57 sides (57%), and the sixth cervical vertebra on 3
sides (3%), while the lower limit of the origin is the sev- Synonyms
enth cervical vertebra on 16 sides (16%), the frst thoracic This muscle is also referred to as serratus posticus inferior
vertebra on 77 sides (77%), and the second thoracic ver- (Macalister 1875).
tebra on 6 sides (6%).
Serratus posterior superior inserted most frequently onto Typical Presentation
ribs 2 through 5 on 148 out of 325 sides (46%). The other Description
insertions included from the second through fourth ribs on Serratus posterior inferior originates from the spines of the
84 sides (26%), the third through ffth ribs on 72 sides (23%), last two thoracic and upper two or three lumbar vertebrae
the second through sixth ribs on 17 sides (5%), and the third (Standring 2016). It inserts via four digitations onto the
and fourth ribs only on four sides (1%) (Mori 1964). inferior margins of ribs 9 through 12 (Standring 2016).
Satoh (1969) described serratus posterior superior on 20
sides of 10 Japanese cadavers. The upper limit of the origin Innervation
of this muscle was the third cervical vertebra on two sides Serratus posterior inferior is supplied by the ninth through
(10%), the fourth cervical vertebra on seven sides (35%), twelfth thoracic spinal nerves (Standring 2016).
the ffth cervical vertebra on nine sides (45%), and the sixth
cervical vertebra on two sides (10%). The lower limit of the Comparative Anatomy
origin was the frst thoracic vertebra on 17 sides (85%) and Serratus posterior inferior originates from the tenth thoracic
the second thoracic vertebra on three sides (15%). vertebra through the second lumbar vertebra and inserts
Trunk Muscles 241
onto ribs 10 through 14 in gibbons (Diogo et al. 2012). It 7 sides (35%), ribs 9 and 10 on 2 sides (10%), ribs 10 and
extends from the thoracolumbar fascia to the last four or 11 on 1 side (5%), and ribs 10 through 12 on 1 side (5%).
fve ribs in orangutans and common chimpanzees (Sonntag Therefore, the upper limit of the insertion was to rib 9 in 18
1923; Diogo et al. 2013ab). In bonobos, it originates from cases (90%) and to rib 10 in 2 cases (10%), while the lower
the thoracolumbar fascia, with vertebral attachments as limit of the insertion was to rib 10 on 2 sides (10%), rib 11
superior as the eighth thoracic vertebra and as inferior as on 8 sides (40%), and rib 12 in 10 cases (50%) (Satoh 1970).
the third lumbar vertebra and attaches to the last four or fve Loukas et al. (2008c) studied serratus posterior inferior
ribs (Miller 1952; Diogo et al. 2017). This muscle is absent on 100 sides of 50 cadavers. They found that the origin of the
in gorillas (Raven 1950; Diogo et al. 2010). muscle was from the eleventh thoracic through second lum-
bar vertebrae in 85% of specimens, from the tenth thoracic
Variations through second lumbar vertebrae in 10% of specimens, and
Description from the tenth thoracic through third lumbar vertebrae in
5% of specimens. The origins were always blended with the
Serratus posterior inferior may be absent (Macalister 1875;
thoracolumbar fascia. It inserted onto the last three ribs in
Rickenbacher et al. 1985; Standring 2016), and if so, the
60% of cases and the last four ribs in 40% of cases.
muscle is replaced by fbrous tissue (Bergman et al. 1988;
Bakkum and Miller 2016). The number of digitations may Anomalies
vary (Macalister 1875; Bergman et al. 1988; Standring
Description
2016), ranging from two to fve digitations (Macalister
1875; Mori 1964; Rickenbacher et al. 1985; Bakkum and Serratus posterior inferior was absent bilaterally in both a
Miller 2016). When it is reduced to only two digitations, neonate with trisomy 18 (Aziz 1979) and a neonate with
often the frst and the fourth digitation are absent (Bergman trisomy 13 (Aziz 1980). In a fetus with spina bifda with
et al. 1988). Mori (1964) notes that the origin may extend encephaloceles, the serratus posterior muscles were uniden-
up to the tenth thoracic vertebra and the insertion may have tifable (Wheeler 1918).
an attachment to the eighth rib. Satoh (1970) notes that the
Prevalence
origin may extend up to the ninth thoracic vertebra.
The insertion of serratus posterior inferior may extend In their literature review, Smith et al. (2015) found that ser-
beneath the external oblique (Rickenbacher et al. 1985). ratus posterior inferior was absent in 4 out of 24 cases of
Serratus posterior inferior may have connections to the exter- trisomy 13 (16.7%) and 1 out of 26 individuals with trisomy
nal intercostal muscles or external oblique (Rickenbacher 18 (3.8%).
et al. 1985) or may be fused with latissimus dorsi (Macalister
1875). Accessory muscle bundles may be present at the infe-
rior border of the digitations, forming arches that extend to N/A
the next lower digitation (Rickenbacher et al. 1985).
levatOres cOstaruM breves anD lOngi (figure 4.8)
Prevalence
Synonyms
Mori (1964) found that the most common upper limit of the
N/A
origin of serratus posterior inferior to be the eleventh thoracic
vertebra on 60 out of 100 sides (60%). The other upper limits Typical Presentation
of the origin included the tenth vertebra on 31 sides (31%), the
Description
twelfth thoracic vertebra on seven sides (7%), the frst lumbar
vertebra on one side (1%), and the second lumbar vertebra on The 12 bundles of levatores costarum originate from the
one side (1%). The most frequent upper limit of the insertion transverse processes of the last cervical vertebra and frst
was to rib 9 on 180 out of 201 sides (90%). The other upper through eleventh thoracic vertebrae (Standring 2016). Each
limits of the insertion included rib 10 on 13 sides (6.5%), rib bundle inserts onto the rib that is immediately below its
8 on 5 sides (2.5%), and rib 11 on 3 sides (1.5%). The lower vertebra of origin (Standring 2016). These bundles insert
limit of the insertion was to rib 12 on 145 out of 206 sides between the tubercles and the costal angles and are referred
(70%) and to rib 11 on 61 sides (30%). Thus, serratus posterior to as levatores costarum breves (Standring 2016). The
inferior inserts most often onto ribs 9 through 12 (Mori 1964). upper eight bundles have only one fascicle. The lowest four
Satoh (1970) described serratus posterior inferior on bundles have two fasciculi, one corresponding to levatores
20 sides of 10 Japanese cadavers. The upper limit of the costarum brevis and the other corresponding to levatores
origin of this muscle was the ninth thoracic vertebra on 2 costarum longus, which inserts onto the second rib below
sides (10%), the tenth thoracic vertebra on 4 sides (20%), its vertebra of origin (Standring 2016).
the eleventh thoracic vertebra on 13 sides (65%), and the
Innervation
twelfth thoracic vertebra on 1 side (5%). It inserted most
frequently onto ribs 9 through 12, which was observed on Levatores costarum are innervated by the thoracic spinal
9 sides (45%). Other insertions include ribs 9 through 11 on nerves (Standring 2016).
Comparative Anatomy tendinous. The most common type of levatores costarum

There is no information for levatores costarum in orang- longi identifed by these authors was type B, found in 51
utans or common chimpanzees (Gibbs 1999; Diogo et al. cases (58.6%).
2013a,b) and little information for the other ape species.
The muscle seems to have a similar typical presentation Anomalies
to humans in gibbons, gorillas, and bonobos, though the N/A
last bundle is attached to the twelfth thoracic vertebra in
gorillas and bonobos (Miller 1952; Gibbs 1999; Diogo et al. Clinical Implications
2010, 2012, 2017). N/A
Variations
interspinales (figure 4.9)
Description
Some bundles of levatores costarum brevis and/or longus Synonyms
may be absent, doubled, or otherwise variable in number N/A
(Satoh and Shu 1968; Rickenbacher et al. 1985; Bakkum
and Miller 2016). Bundles of levatores costarum breves and
longi may unite and present as a sheet (Macalister 1875). Description
Levatores costarum longi may be found in the middle or Interspinales are paired muscular bundles that extend
upper thoracic spine (Satoh and Shu 1968; Rickenbacher between the tips of adjacent spinous processes throughout
et al. 1985). Levatores costarum bundles can send fbers to, the vertebral column (Standring 2016). They are most well-
or fuse with, the external intercostal muscles, scalenus pos- developed in the cervical spine and comprise six pairs of
terior, or iliocostal muscles (Macalister 1875; Rickenbacher muscles (Standring 2016). In the thoracic spine, the muscles
et al. 1985). In some cases, all bundles of levatores costa- are distinct between the frst and second thoracic vertebrae
rum may be absent (Rickenbacher et al. 1985). and between the eleventh and twelfth thoracic vertebrae. In
Satoh and Shu (1968) studied the levatores costarum the lumbar spine, there are four pairs of muscles (Standring
muscles in a Japanese population and note instances where 2016).
a muscle originated via tendinous fbers from the lateral
side of levator costae brevis and passed over the rib above to Innervation
insert into the belly of levator costae brevis in the next inter- The interspinales muscles are innervated by the medial
costal space. These authors suggest that it is an inverted branch of the dorsal ramus of the spinal nerve of the cor-
and incomplete form of levator costae longus. Satoh and responding segment (Rickenbacher et al. 1985).
Shu (1968) also categorized bundles of levatores costarum
longi into four types. They defned type A as an incom- Comparative Anatomy
plete form of the muscle which terminated on the exter- There is no information about interspinales for gibbons,
nal intercostal in the next intercostal space after crossing orangutans, or bonobos (Diogo et al. 2012, 2013b, 2017).
only over one rib. Type B is the typical presentation of this Sonntag (1923) states that interspinales in common chim-
muscle, which crosses over one rib to insert onto the next panzees is similar to those of humans. In gorillas, the
rib and thus spans two intercostal spaces. Type C bundles muscles have a similar typical presentation to humans and
crossed two intercostal spaces and terminated on the exter- the superiormost attachment of interspinales is to the third
nal intercostal muscle in the next intercostal space. Type D cervical vertebra and the inferiormost attachment is to the
bundles extended over three intercostal spaces (Satoh and second thoracic vertebra (Raven 1950; Gibbs 1999; Diogo
Shu 1968). et al. 2010).
Macdonald Brown (1880) describes the bilateral pres-
ence of a muscle termed levator costae primae. Levator Variations
costae primae originates from the superior border of the Description
scapula and partially from the suprascapular ligament The interspinales muscles may be doubled or absent
and inserts onto the frst rib, immediately external to its (Miyauchi 1976; Bakkum and Miller 2016). In the cervical
cartilage (Macdonald Brown 1880; Snosek and Loukas region, the interspinales muscles may extend across more
2016). than two vertebrae (Standring 2016). The muscles in this
part of the spine sometimes blend with neighboring muscles
Prevalence (Rickenbacher et al. 1985). The interspinales in the tho-
The variable muscle described by Satoh and Shu (1968) racic spine vary substantially in number (Miyauchi 1976;
that extended from levator costae brevis to the belly of Rickenbacher et al. 1985). In this region, bundles between
levator costae brevis in the next intercostal space was the second and third thoracic vertebrae are occasionally
found in 2 out of 87 cases (2.3%). In two other cases present (Standring 2016). In the lumbar spine, the interspi-
(2.3%), two muscles classifed as type B were completely nales muscles may be present between the last thoracic and
Trunk Muscles 243
FIGURE 4.9 Interspinal, intertransverse, and caudal spinotransverse muscles in posterior view.
frst lumbar vertebrae (Standring 2016). Interspinales mus- the lower thoracic region, the interspinales were always
cles may be present in the sacral region (Miyauchi 1976; present and the upper limit of these muscles was the tenth
Standring 2016). thoracic vertebra on 22 sides (36.7%), the eleventh thoracic
vertebra on 35 sides (58.3%) and the twelfth thoracic ver-
Prevalence tebra on three sides (5%). The interspinales in the lumbar
Based on a study of 60 sides from 30 cadavers, Miyauchi region were also present in all cases except for the muscle
(1976) found that all interspinales in the cervical region between the fourth and ffth lumbar vertebrae, which was
were present in all cases except for the last (the muscle absent on four sides (6.7%), and the muscle between the
between the seventh cervical and frst thoracic vertebrae), last lumbar and frst sacral vertebrae, which was absent
which was absent on three sides (5%). In the upper tho- on 20 sides (33.3%). A muscle was present between the
racic region, the interspinales between the frst and second frst and second sacral vertebrae on eight sides (13.3%)
thoracic vertebrae were present on only 26 sides (43.3%) (Miyauchi 1976).
and the interspinales between the second and third tho-
racic vertebrae were present on three sides (5%). These Anomalies
muscles were always absent in the mid-thoracic region. In N/A
Clinical Implications Typical Presentation

N/A Description
Intertransversarii connect the transverse processes of adja-
sacrOcOccygeus pOsteriOr (figure 4.9) cent vertebrae (Standring 2016). The intertransversarii
See also: Interspinales; Sacrococcygeus anterior are most distinct in the cervical spine and are composed
of seven pairs of anterior and posterior sets of muscles
Synonyms (Standring 2016). In the thoracic spine, the intertransversa-
This muscle may also be referred to as sacrococcygeus rii are single muscles that connect adjacent transverse pro-
posticus, levator coccygis (Morgagni), extensor coccygis cesses between the last three thoracic vertebrae and the frst
(Macalister 1875; Knott 1883b), or sacrococcygeus dorsalis lumbar vertebra (Standring 2016). In the lumbar spine, the
(Rickenbacher et al. 1985). intertransversarii are composed of medial and lateral sets
of muscles (Standring 2016).
Innervation
This muscle may be only present as a variation, but Macalister
(1875, p. 66) states that it is “a tolerably constant muscle.” The intertransversarii are innervated by either the ventral
or dorsal rami of the spinal nerve of the corresponding seg-
Comparative Anatomy ment (Standring 2016).
This muscle may be homologous with the extensor (leva-
Comparative Anatomy
tor) caudae lateralis of other mammals (Knott 1883b;
Rickenbacher et al. 1985). There is no information on intertransversarii in gibbons
(Gibbs 1999; Diogo et al. 2012), and the intertransversarii
Variations in the other ape species are simply described as being simi-
Description lar to those of humans (Sonntag 1923; Raven 1950; Gibbs
1999; Diogo et al. 2010, 2013ab, 2017).
A variant of interspinales is sacrococcygeus posterior, a
musculotendinous bundle that connects the sacrum to the Variations
coccyx and has an attachment to the dorsal sacrococcygeal
Description
ligaments (Rickenbacher et al. 1985). Sacrococcygeus pos-
terior is covered by strands of the sacrotuberal ligament and Any of the intertransverse muscles may be doubled or
by a portion of the gluteus maximus fascia (Rickenbacher absent (Macalister 1875; Bakkum and Miller 2016). The
et al. 1985). Knott (1883b) observed an origin from the pos- anterior set of intertransversarii muscles between the
terior surface of the third and fourth sacral bones in most frst and second cervical vertebrae are frequently absent
cases and from the posterior inferior iliac spine in one case. (Standring 2016). Accessory intertransverse muscles have
See the entry for sacrococcygeus anterior for a similar mus- been described with attachments from the fourth through
cle on the anterior aspect of the sacrum. sixth to the frst three cervical vertebrae (transversalis cer-
vicis anticus of Retzius) and between the third and ffth cer-
Innervation vical vertebrae (Sandifort’s muscle or musculus singularis)
N/A (Macalister 1875; Knott 1883a; Bakkum and Miller 2016).
The anterior cervical intertransverse muscles may fuse with
Prevalence the scalene muscles (Rickenbacher et al. 1985).
Knott (1883b) found feshy fbers corresponding to “sacro- In rare cases, the intertransverse muscles do not connect
coccygeus posticus” in 4 out of 30 subjects (13.3%). to the transverse processes, and instead fuse with the erec-
tor spinae (Rickenbacher et al. 1985). Adjacent intertrans-
Anomalies verse muscles may partially or completely fuse into strands
While the presence of sacrococcygeus anterior has been that bridge across one or more segments, a variation that
recorded in one individual with trisomy 21 (Bersu 1980), occurs more often in the thoracolumbar region than in the
the presence of sacrococcygeus posterior has not been cervical spine (Rickenbacher et al. 1985). The intertrans-
recorded in individuals with trisomy, birth defects, or other versarii laterales in the lumbar region may fuse with qua-
congenital disorders, to our knowledge. dratus lumborum (Rickenbacher et al. 1985).

N/A N/A
Anomalies
intertransversarii (figure 4.9) N/A
Synonyms
These muscles may also be referred to as intertransversalis
N/A
(Macalister 1875).
Trunk Muscles 245
rOtatOres lOngus anD brevis (figure 4.9) Prevalence

N/A
Synonyms
These muscles may be referred to as rotatores spinae Anomalies
(Macalister 1875). N/A
Typical Presentation Clinical Implications
Description N/A
The rotatores muscles comprise part of the transversospina-
lis muscular group and are most well-developed in the tho-
racic spine (rotatores thoracis) (Standring 2016). Rotatores MultifiDus (figure 4.9)
thoracis is comprised of 11 pairs of muscles, each with a Synonyms
longus and brevis portion (Standring 2016). Rotatores This muscle may also be referred to as multifdus spinae
breves connect the lamina of a vertebra to the transverse (Macalister 1875).
process of the adjacent vertebra below (Standring 2016).
Rotatores longi connect the spinous process of a vertebra to Typical Presentation
the transverse process of the vertebra two segments below Description
(Standring 2016).
Multifdus is one of the muscles that comprises the transver-
Innervation sospinalis muscular group. It is comprised of bundles that
lie in the grooves between the spinous processes and trans-
Rotatores are innervated by medial branches of the dorsal
verse processes along the entire vertebral column (Standring
rami of the spinal nerves of the corresponding segment
2016). Each bundle extends between the lateral aspect of a
(Standring 2016).
vertebral spinous process and the transverse parts of the ver-
Comparative Anatomy tebrae two, four, and fve segments below (Standring 2016).
There is no information on rotatores in gibbons (Gibbs Multifdus thus has attachments to the articular processes of
1999; Diogo et al. 2012). These muscles in the other apes the last four cervical vertebrae, to the transverse processes
have a similar typical presentation to those of humans, but of the thoracic vertebrae, to the mamillary processes of the
it is possible that rotatores longi is present only in gorillas lumbar vertebrae, to the dorsal surface of the sacrum as far
and absent in common chimpanzees, bonobos, and orang- as the fourth sacral foramen, and to the dorsal aspect of the
utans (Sonntag 1923; Raven 1950; Gibbs 1999; Diogo et al. iliac crest (Standring 2016; Rickenbacher et al. 1985).
2010, 2013a,b, 2017). Innervation
Variations Multifdus is innervated by medial branches of the dorsal
rami of the corresponding spinal nerves (Standring 2016).
Description
One or more of the rotatores muscles may be absent from Comparative Anatomy
the upper thoracic vertebrae or the lower thoracic vertebrae There is no descriptive information on multifdus for gib-
(Macalister et al. 1875; Rickenbacher et al. 1985; Standring bons (Gibbs 1999; Diogo et al. 2012) or orangutans, though
2016). The number of rotatores breves in particular may Diogo et al. (2013b) did observe the presence of this muscle
range from 9 to 12 in this region (Rickenbacher et al. 1985). in an orangutan. The muscles in gorillas, common chim-
The rotatores longi may be more poorly developed than the panzees, and bonobos have a similar typical presentation to
rotatores breves (Rickenbacher et al. 1985). The rotatores that in humans, but an attachment may extend to the atlas
may be divided into fasciculi (Macalister 1875). Accessory in bonobos (Raven 1950; Miller 1952; Gibbs 1999; Diogo
bundles or accessory attachments across two adjacent ver- et al. 2010, 2013a, 2017).
tebrae may occur in the thoracic region (Rickenbacher et al.
1985; Cornwall et al. 2011; Bakkum and Miller 2016). Variations
The presence of rotatores muscles in the cervical and
lumbar regions are often variable (Rickenbacher et al. Description
1985), and they may be replaced by deep bundles of mul- Multifdus is often variable in its confguration and the
tifdus (Standring 2016). They may be completely absent muscle may be asymmetrical between the sides of an indi-
(Rickenbacher et al. 1985; Cornwall et al. 2011). Sometimes, vidual (Rickenbacher et al. 1985). One or more digitations
the complete series of rotatores cervicis can be present may be absent (Macalister 1875). The digitation that origi-
(Rickenbacher et al. 1985). When present, the rotatores lum- nates from the seventh cervical vertebra may be absent
borum are represented by strands somewhere between the (Macalister 1875; Rickenbacher et al. 1985). The digitation
last thoracic vertebra and the frst or second sacral vertebrae that extends between the axis and the fourth cervical ver-
and are diffcult to separate from interspinales (Rickenbacher tebra may be attached across the entire breadth of the arch
et al. 1985). of the axis (Macalister 1875; Rickenbacher et al. 1985). In
the cervical region, accessory bundles may be present that onto the transverse processes of the sixth through tenth tho-
attach to the articular and transverse processes and to the racic vertebrae (Standring 2016).
capsules of the vertebral joints (Rickenbacher et al. 1985).
Willard (1997) reports that multifdus in the lumbar region Innervation
attaches to the thoracolumbar fascia via a raphe. Semispinalis capitis is innervated by the greater occipi-
Prevalence tal nerve and the third cervical nerve (Standring 2016).
Semispinalis cervicis and semispinalis thoracis are inner-
N/A
vated by medial branches of the dorsal rami of the corre-
Anomalies sponding spinal nerves (Standring 2016).
Stevenson et al. (2014) describe cadaver with a severe case In common chimpanzees, semispinalis cervicis and thora-
of congenital scoliosis. Multifdus was atrophied on the cis are diffcult to separate and semispinalis capitis is often
left side and could not be identifed above the level of the fused with parts of longissimus and spinalis (Sonntag 1923;
tenth thoracic spinous process. Multifdus on the right side Diogo et al. 2013a). There is no information for semispinalis
had a more typical presentation but many digitations were thoracis in bonobos (Miller 1952; Diogo et al. 2017), but the
replaced by tendinous slips (Stevenson et al. 2014). other portions of semispinalis resemble those of humans.
Sonntag (1924) has noted the presence of the “biventer cer-
Prevalence vicis” bundle in an orangutan.
N/A
Clinical Implications Variations

N/A Description
The slips of semispinalis capitis to the vertebrae may vary
in number (Macalister 1875; Bergman et al. 1988; Bakkum
seMispinalis (figure 4.10) and Miller 2016). A slip may extend from the third cervical
vertebra or from the eighth thoracic vertebra (Rickenbacher
Synonyms
et al. 1985). Semispinalis capitis may be partially divided into
Semispinalis capitis may also be referred to as complexus two bellies with separate tendons of origin that insert together
major (Krause) (Macalister 1875), complexus (Cowper), and onto the occiput (Çelik et al. 1997). Semispinalis capitis may
grand complexus (Cruveilhier) (Bakkum and Miller 2016). receive slips from longissimus or spinalis thoracis (Macalister
Semispinalis cervicis may also be referred to as semispi- 1875; Rickenbacher et al. 1985; Bergman et al. 1988; Bakkum
nalis colli, and semispinalis thoracis as semispinalis dorsi and Miller 2016). Semispinalis capitis is often blended with
(Macalister 1875). The medial portion of semispinalis capi- spinalis capitis (Bakkum and Miller 2016).
tis in the cervical region may be referred to as biventer cer- A second lamella under the medial part of semispinalis
vicis because its superfcial bundles appear distinct from capitis extending from one or more of the thoracic vertebrae
the rest of the muscle and because it has an incomplete ten- to the occipital bone may be present (Bergman et al. 1988;
dinous intersection, giving it the appearance of two distinct Bakkum and Miller 2016). There may be accessory bundles
bellies (Bergman et al. 1988; Standring 2016). that connect to the nuchal ligament or run between semispi-
nalis capitis and rectus capitis posterior major (Macalister
1875; Rickenbacher et al. 1985). An accessory muscle that
Description runs parallel to semispinalis capitis may be present extend-
The semispinalis muscles are part of the transversospi- ing between the transverse process of the axis to the occipi-
nalis muscular group and are divided into three portions tal bone (Macalister 1875; Rickenbacher et al. 1985).
(Standring 2016). Semispinalis capitis originates from the Semispinalis cervicis and semispinalis thoracis can vary
occipital bone between the superior and inferior nuchal in the number of origins and insertions (Macalister 1875;
lines (Standring 2016). It inserts via tendons onto the artic- Rickenbacher et al. 1985). The attachments of semispina-
ular processes of cervical vertebrae four through seven and lis cervicis may include the seventh cervical or the seventh
the transverse processes of the frst six or seven thoracic or eighth thoracic vertebrae (Rickenbacher et al. 1985).
vertebrae (Standring 2016). Semispinalis cervicis is cov- Semispinalis thoracis may also vary in the number of verte-
ered by semispinalis capitis (Standring 2016). It originates brae that its fbers bridge across (Bakkum and Miller 2016).
from the spinous processes of the second to ffth cervi- It may be diffcult to distinguish the bundles of semispina-
cal vertebrae (Standring 2016). Its bundles insert onto the lis cervicis from those of semispinalis thoracis (Macalister
transverse processes of the upper fve or six thoracic verte- 1875; Rickenbacher et al. 1985).
brae (Standring 2016). Semispinalis thoracis originates via Semispinalis lumborum may be present as a variation
tendons from the spinous processes of the last two cervical (Rickenbacher et al. 1985). Typically, remnants of semispinalis
and the frst four thoracic vertebrae and inserts via tendons in the lumbar spine are represented by the mammillo-accessory
Trunk Muscles 247
ligaments (Standring 2016). However, when semispinalis lum- variable vertebral attachments in some species and an
borum is present it is covered by fascia that is attached to the occasional slip to levator scapulae in common chimpanzees
mamillary processes of the last four lumbar vertebrae and the (Sonntag 1923; Diogo et al. 2010, 2012, 2013a,b, 2017).
frst sacral vertebra and is fused to the aponeurosis of longis-
simus (Rickenbacher et al. 1985). This muscle also has tendi- Variations
nous origins from the mamillary processes of the last thoracic
and frst lumbar vertebrae and attachments to the spinous pro- Description
cesses of the last six thoracic vertebrae and the frst lumbar The splenius muscles may be absent (Rickenbacher et al. 1985;
vertebra (Rickenbacher et al. 1985). It may be diffcult to sepa- Bergman et al. 1988). Other variations include proximal dis-
rate from multifdus (Rickenbacher et al. 1985). placement of the origin of the splenius muscles by one or two
vertebrae, or asymmetric origins (Rickenbacher et al. 1985;
Bergman et al. 1988; Bakkum and Miller 2016). Splenius
Prevalence
capitis may be divided into a superior and inferior portion
N/A (Macalister 1875; Bergman et al. 1988; Bakkum and Miller
Anomalies 2016). Splenius capitis may be blended with splenius cervicis
(Macalister 1875; Rickenbacher et al. 1985; Kamibayashi and
Description Richmond 1998; Bakkum and Miller 2016). Slips can con-
On the right side of one male neonate with trisomy 13, nect the splenius muscles to longissimus capitis, iliocostalis,
Pettersen et al. (1979) found that semispinalis cervicis was levator scapulae, serratus anterior, or serratus posterior supe-
connected via an accessory slip to obliquus capitis inferior. rior (Wood 1870; Macalister 1875; Rickenbacher et al. 1985;
Bergman et al. 1988; Bakkum and Miller 2016).
Prevalence
According to the literature review of Smith et al. (2015), Prevalence
the anomalous slip observed by Pettersen et al. (1979) was N/A
present in only 1 out of 20 individuals with trisomy 13 (5%).
Anomalies
Description
N/A
In a trisomy 18 cyclopic fetus, Smith et al. (2015) found that
splenius capitis was closely associated with/fused with tra-
splenius capitis anD splenius cervicis (figure 4.10) pezius at the midline origin on both sides (Smith et al. 2015).
Synonyms Prevalence
Splenius capitis is also referred to as Riemenmuskeln The close connection between splenius capitis and the tra-
(Bakkum and Miller 2016). Splenius cervicis is also pezius observed by Smith et al. (2015) was not present in
referred to as Bauschmuskeln or splenius colli (Macalister any of the trisomy 13, trisomy 18, and trisomy 21 cases
1875; Bakkum and Miller 2016). included in their literature review.
Typical Presentation Clinical Implications
Description N/A
Splenius capitis extends from the mastoid process and the
occipital bone to the spinous processes of the last cervi-
rhOMbOatlOiDeus (figure 4.10)
cal and upper three or four thoracic vertebrae (Standring
2016). Splenius cervicis lies close to the inferior border of See also: Splenius capitis and splenius cervicis
splenius capitis, originating from the transverse processes
of the frst two cervical vertebrae and the posterior tubercle Synonyms
of the third cervical vertebra (Standring 2016). It inserts This muscle is also referred to as rhomboaxoideus (Wood
onto the spinous processes of the third through sixth tho- 1867a), splenius accessorius (Krause), splenius colli acces-
racic vertebrae (Standring 2016). sorius, splenius capitis accessorius, muscularis singularis
splenius accessories (Walther), or adjutor splenii (Macalister
Innervation 1975; Knott 1883a; Bergman et al. 1988; Bakkum and
Splenius capitis is innervated by the second and third cervi- Miller 2016).
cal spinal nerves (Standing 2016). Splenius cervicis is inner-
vated by the lower cervical spinal nerves (Standing 2016). Typical Presentation
Comparative Anatomy
Splenius capitis and splenius cervicis have a similar typical Comparative Anatomy
presentation in the apes, with similar variations including N/A
FIGURE 4.10 Splenius, semispinalis, and nearby variations in posterior view.
Description Macalister (1875) notes that Wood found rhomboatloideus
Rhomboatloideus describes a slip that originates from in 3 out of 36 subjects (8.3%). Knott (1883a) found this mus-
the spines of the lower cervical and upper thoracic ver- cle in 6 out of 75 subjects (8%).
tebrae (anywhere from C6 to T3), close to the attachment
of rhomboid minor, and passes superfcially along the Anomalies
border of splenius to insert on the transverse process of Description
the atlas (Macalister 1866, 1875; Knott 1883a; Bergman Beger et al. (2019) observed rhomboatloideus bilaterally
et al. 1988; Bakkum and Miller 2016). It can also origi- in a 26-week old male fetus with CHARGE syndrome.
nate from the fascia of serratus posterior superior or Rhomboatloideus extended from the transverse process of
rhomboideus minor (Macalister 1975; Bergman et al. the atlas to the spinous processes of the second, third, and
1988, Beger et al. 2018a). It may have attachments to the fourth thoracic vertebrae (Beger et al. 2019).
mastoid process or occipital bone (Bergman et al. 1988).
Rhomboatloideus may have slips that blend with rhom- Prevalence
boideus major and serratus posterior superior (Wood N/A
1867a,b; Bakkum and Miller 2016).
Innervation Rhomboatloideus may be incorrectly diagnosed as a tumor or
N/A can contribute to myofascial trigger points (Beger et al. 2018a).
Trunk Muscles 249
atlantOMastOiDeus (figure 4.10) spinalis dorsi and spinalis cervicis may be referred to as
spinalis colli (Macalister 1875).
Synonyms
This muscle is also referred to as atlanticomastoideus
(Gruber) or rectus lateralis accessorius (Winslow) (Bergman
et al. 1988). Description
Spinalis is partitioned into spinalis thoracis, spinalis cer-
Typical Presentation vicis, and spinalis capitis, which are all continuous with
each other (Standring 2016). Spinalis thoracis is the best-
developed and most constant of the three portions (Standring
Comparative Anatomy 2016). Bundles of spinalis thoracis extend between the spi-
Among the apes, atlantomastoideus has only been reported nous processes of the lower two thoracic and upper two
as a variation in gorillas and common chimpanzees lumbar vertebrae and the spinous processes of the upper
(Gratiolet and Alix 1866; Sommer 1907; Diogo et al. 2010, thoracic vertebrae (Standring 2016).
2012, 2013a,b, 2017). When present, spinalis cervicis extends from the spinous
processes of the second through fourth cervical vertebrae
Variations to the ligamentum nuchae and spine of the last cervical
vertebra (Standring 2016). When present, spinalis capitis is
Description
represented by fbers of semispinalis capitis that attach to
Atlantomastoideus originates from the transverse process the spinous processes of the last cervical and frst thoracic
of the atlas, courses between the rectus capitis lateralis and vertebrae (Standring 2016).
obliquus capitis superior, and attaches to the mastoid pro-
cess (Gruber 1867c; Knott 1883a; Mori 1964; Rickenbacher Innervation
et al. 1985; Bergman et al. 1988; Bakkum and Miller 2016). The erector spinae group is innervated by the lateral
Its insertion may occur at the posterior margin of the mas- branches of the dorsal rami of the corresponding cervical,
toid process or the lateral or medial margin of the mastoid thoracic, and lumbar spinal nerves (Standring 2016).
notch (Rickenbacher et al. 1985). Accessory slips may be
present that connect atlantomastoideus to splenius capitis, Comparative Anatomy
longissimus capitis, or the axis (Mori 1964). Based on their Spinalis has a similar typical presentation in the apes, often
shared innervation, Mori (1964) suggests that atlantomas- with fusion to semispinalis in gorillas (Gibbs 1999; Diogo
toideus derives from the same anlage as longissimus capitis et al. 2010, 2012, 2013a,b, 2017).
and obliquus capitis superior.
Variations
Atlantomastoideus is innervated by the dorsal ramus of the Spinalis is rarely completely absent but is also rarely sym-
frst cervical spinal nerve (Mori 1964). metrical on the left and right sides (Rickenbacher et al.
1985). Spinalis thoracis can vary in its number of digitations
Prevalence and attachments (Macalister 1875; Rickenbacher et al. 1985;
Gruber lists the prevalence of atlantomastoideus as 20%, but Bakkum and Miller 2016; Standring 2016). Spinalis thoracis
Rickenbacher et al. (1985) suggest that this is likely too high. is typically blended with longissimus thoracis and may be
Knott (1883a) found this muscle in 4 out of 33 subjects (12.1%). considered a part of this muscle (Standring 2016). Spinalis
Mori (1964) found atlantomastoideus bilaterally in 5 out of 54 cervicis can send attachments to the spines of the frst two
cadavers (9.3%), on the right side in 5 cadavers (9.3%), and on thoracic vertebrae (Standring 2016) and may be fused with
the left side in 4 cadavers (7.4%). Therefore, out of 108 sides, semispinalis cervicis (Greiner et al. 2004; Bakkum and
atlantomastoideus was present in 19 sides (17.6%). Miller 2016). Spinalis capitis is often fused with semispina-
lis capitis and can also be fused with rectus capitis posterior
Anomalies minor (Rickenbacher et al. 1985; Martin 1994; Greiner et al.
N/A 2004; Bakkum and Miller 2016). Eister (1912) considers the
presence of spinalis capitis to be a variant and not part of the
Clinical Implications typical presentation of spinalis (Rickenbacher et al. 1985).
N/A Spinalis lumborum, which extends between the spinous pro-
cesses of the lower two lumbar vertebrae and some of the
spinalis (figure 4.11) lower thoracic vertebrae, may also be present as a rare varia-
tion (Rickenbacher et al. 1985).
Synonyms Labranche et al. (2017) describe what they believe
Miyauchi (1976) refers to spinalis cervicis as interspinalis to be a distinct and unilateral bundle of spinalis capitis.
cervicalis longus. Spinalis thoracis may be referred to as Acknowledging that this muscle has a presentation that
FIGURE 4.11 Erector spinae muscles in posterior view.
does not match previous descriptions, they state a bundle of vertebra on 5 sides (10.2%). The lower limit of this muscle
spinalis capitis arose deep to semispinalis capitis and was was the fourth cervical vertebra on 2 sides (4.1%), the ffth
on the same fascial plane as spinalis cervicis. The muscle cervical vertebra on 12 sides (24.5%), the sixth cervical ver-
originated from the ligamentum nuchae at the level of the tebra on 18 sides (36.7%), the seventh cervical vertebra on
third and fourth cervical vertebrae and attached to the 11 sides (22.4%), and the frst thoracic vertebra on 6 sides
occiput just superfcial to the rectus capitis posterior minor (12.2%) (Miyauchi 1976).
at the inferior nuchal line. Based on dissections of 142 cadavers (284 sides), Greiner
et al. (2004) found that spinalis cervicis was absent on 102
Prevalence sides (35.9%), blended with semispinalis cervicis on 51 sides
In a study of 60 sides from 30 Japanese cadavers, Miyauchi (18%), and distinct from semispinalis cervicis on 131 sides
(1976) reports that spinalis cervicis was present in 49 sides (46.1%). Based on dissections of the 279 sides that would
(81.7%) and absent on 11 sides (18.3%). Miyauchi (1976) have allowed for the observation of spinalis capitis, Greiner
also found that the upper limit of spinalis cervicis was the et al. (2004) found that spinalis capitis was absent on 196
second cervical vertebra on 36 sides (73.5%), the third cer- sides (70.3%), blended with semispinalis capitis on 80 sides
vical vertebra on 8 sides (16.3%), and the fourth cervical (28.7%), and distinct from semispinalis capitis on 3 sides (1%).
Trunk Muscles 251

Description Longissimus has a similar typical presentation in the apes
Stevenson et al. (2014) describe a cadaver with a severe case (Sonntag 1923; Miller 1952; Gibbs 1999; Diogo et al. 2010,
of congenital scoliosis. The erector spinae muscles were 2012, 2013ab, 2017). It may be fused with iliocostalis and
severely atrophied on the left side and many parts were semispinalis capitis in orangutans (Diogo et al. 2013b).
replaced by tendinous insertions. Erector spinae on the right
side was comprised of typical feshy muscle fbers but the Variations
tendons of iliocostalis inserted in a radial manner due to the Description
abnormal thoracolumbar curvature (Stevenson et al. 2014). Longissimus thoracis can be variable in the number of
its costal attachments and which ribs it is attached to
Prevalence (Macalister 1875; Rickenbacher et al. 1985; Bergman
N/A et al. 1988). Longissimus cervicis can extend to the trans-
Clinical Implications verse processes of the tenth or eleventh thoracic verte-
brae or have a reduced number of bundles (Rickenbacher
Labranche et al. (2017) note that somatic dysfunction of spi-
et al. 1985; Bergman et al. 1988). Longissimus capitis
nalis capitis can contribute to compressive neuropathy of
can be diminished, absent, divided into two parts, or can
the dorsal rami of the cervical spinal nerves. Asymmetry
occasionally be attached to the atlas (Macalister 1875;
of spinalis capitis may unilaterally strain the ligamentum
Rickenbacher et al. 1985; Bergman et al. 1988; Bakkum
nuchae and lead to the development of cervicogenic head-
and Miller 2016). The attachment of longissimus capitis
aches or occipital headaches (Labranche et al. 2017).
may extend to the ffth thoracic vertebra or can be reduced
to just the last two cervical vertebrae (Macalister 1875;
lOngissiMus (figure 4.11) Rickenbacher et al. 1985). Longissimus capitis may also
fuse with the splenius (Macalister 1875; Rickenbacher
Synonyms et al. 1985).
Longissimus thoracis may be referred to as longissimus Accessory slips are occasionally present on the under-
dorsi (Macalister 1875). side of the lumbar portion longissimus (Bergman et al.
1988; Bakkum and Miller 2016). These slips can fuse to
Typical Presentation form a supernumerary muscle termed transversalis dorsi
Description (or transversalis thoracis) that can attach to the frst two
or three thoracic vertebrae (Bergman et al. 1988; Bakkum
Longissimus is partitioned into longissimus thoracis, longis-
and Miller 2016). Accessory slips can also be found near
simus cervicis, and longissimus capitis, which are all contin-
the cervical portion of longissimus and can form a super-
uous with each other (Standring 2016). Longissimus capitis
numerary muscle termed transversalis cervicis posticus
originates from the mastoid process and attaches via tendons
minor (Knott 1883a; Bergman et al. 1988; Bakkum and
to the transverse processes of the frst four thoracic verte-
Miller 2016). Transversalis cervicis posticus minor (tra-
brae and the lower three or four cervical vertebrae (Standring
chelomastoideus minor or trachelomastoideus accessorius)
2016). Longissimus cervicis originates via tendons from the
arises from the transverse processes of the second thoracic
posterior tubercles of cervical vertebrae two through six and
through ffth cervical vertebrae and inserts into the trans-
inserts via tendons onto the transverse processes of the frst
verse process of the frst cervical vertebra and the mastoid
four or fve thoracic vertebrae (Standring 2016).
process (Knott 1883a). Slips from longissimus capitis may
The thoracic portion of longissimus thoracis originates
originate from the transverse process of the frst cervical
from the transverse processes of the frst four thoracic ver-
vertebra, or from the longissimus cervicis tendon near this
tebrae and from the transverse processes and corresponding
vertebra and insert onto the mastoid process (Bergman
ribs in the next eight thoracic segments (Standring 2016).
et al. 1988).
It inserts onto the spinous processes of the lumbar verte-
brae, the sacrum, and the iliac crest (Standring 2016). The Prevalence
lumbar portion of longissimus thoracis originates from the
N/A
accessory process and transverse process of each lumbar
vertebrae, and inserts onto the ilium, either directly or via
Anomalies
a tendon termed the lumbar intermuscular aponeurosis
(Standring 2016). Description
Stevenson et al. (2014) describe a cadaver with a severe case
Innervation of congenital scoliosis. The erector spinae muscles were
The erector spinae group is innervated by the lateral severely atrophied on the left side, and many parts were
branches of the dorsal rami of the corresponding cervical, replaced by tendinous insertions. Erector spinae on the right
thoracic, and lumbar spinal nerves (Standring 2016). In the side was comprised of typical feshy muscle fbers but the
lumbar region, intermediate branches supply longissimus tendons of iliocostalis inserted in a radial manner due to the
(Standring 2016). abnormal thoracolumbar curvature (Stevenson et al. 2014).
Prevalence In the lumbar region, lateral branches supply iliocostalis

Clinical Implications Comparative Anatomy

N/A Iliocostalis has a similar typical presentation in the apes,
with similar variations in costal and vertebral attachments
(Diogo et al. 2010, 2012, 2013a,b, 2017).
iliOcOstalis (figure 4.11)
Synonyms Variations
N/A Description
Iliocostalis can vary in the number of its vertebral and
Typical Presentation costal attachments (Macalister 1875; Rickenbacher et al.
Description 1985; Bakkum and Miller 2016). The digitations to the
Iliocostalis is partitioned into iliocostalis lumborum, frst, eleventh, and twelfth ribs are most commonly absent
iliocostalis thoracis, and iliocostalis cervicis, which are (Macalister 1875). Accessory slips may be attached to the
all continuous with each other (Standring 2016). The middle ribs (Macalister 1875). In the lumbar portion of ilio-
lumbar portion of iliocostalis lumborum originate from costalis, there may be a slip to the lumbocostal ligament
the transverse processes of the frst four lumbar verte- (Rickenbacher et al. 1985).
brae and the thoracolumbar fascia and inserts onto the
Prevalence
iliac crest (Standring 2016). The thoracic portion of
iliocostalis lumborum originates via tendons from the N/A
angles of the lower eight or nine ribs, and its bundles
converge into an aponeurosis that inserts onto the iliac Anomalies
crest (Standring 2016). Iliocostalis thoracis originates Description
from the transverse process of the last cervical vertebra Stevenson et al. (2014) describe a cadaver with a severe case
and from the frst six ribs and inserts onto the lower six of congenital scoliosis. The erector spinae muscles were
ribs (Standring 2016). It also has fascicular attachments severely atrophied on the left side, and many parts were
to the common erector spinae tendon (Gale et al. 2016). replaced by tendinous insertions. Erector spinae on the right
Iliocostalis cervicis originates from the posterior tuber- side was comprised of typical feshy muscle fbers, but the
cles of the transverse processes of cervical vertebrae tendons of iliocostalis inserted in a radial manner due to the
four through six and attaches to the angles of ribs three abnormal thoracolumbar curvature (Stevenson et al. 2014).
through six (Standring 2016).
Prevalence
Innervation N/A
The erector spinae group is innervated by the lateral
branches of the dorsal rami of the corresponding cervi- Clinical Implications
cal, thoracic, and lumbar spinal nerves (Standring 2016). N/A
5 Lower Limb Muscles
Eve K. Boyle
Vondel S. E. Mahon
Rui Diogo
Malynda Williams
CONTENTS
Gluteal and Thigh Muscles��261
Gluteus maximus��261
Synonyms��261
Typical Presentation��261
Comparative Anatomy��262
Variations��262
Anomalies��263
Clinical Implications��263
Gluteus medius��263
Synonyms��263
Variations��263
Anomalies��264
Gluteus minimus (Not Illustrated)�� 264
Synonyms��264
Variations��264
Anomalies��265
Piriformis��265
Synonyms�� 265
Typical Presentation�� 265
Comparative Anatomy�� 265
Variations�� 265
Anomalies�� 267
Clinical Implications�� 267
Scansorius��267
Synonyms�� 267
Typical Presentation�� 267
Comparative Anatomy�� 267
Variations�� 267
Anomalies�� 268
Clinical Implications�� 268
DOI: 10.1201/9781003083535-5 253

Tenuissimus...................................................................................................................................................................268
Synonyms .................................................................................................................................................................268
Variations..................................................................................................................................................................268
Anomalies.................................................................................................................................................................268
Tensor fasciae latae .......................................................................................................................................................269
Synonyms .................................................................................................................................................................269
Variations.................................................................................................................................................................. 270
Anomalies.................................................................................................................................................................270
Sartorius ........................................................................................................................................................................270
Synonyms .................................................................................................................................................................270
Variations..................................................................................................................................................................270
Anomalies.................................................................................................................................................................271
Rectus femoris...............................................................................................................................................................271
Synonyms .................................................................................................................................................................271
Variations..................................................................................................................................................................271
Anomalies.................................................................................................................................................................272
Vastus medialis..............................................................................................................................................................272
Synonyms .................................................................................................................................................................272
Variations..................................................................................................................................................................272
Anomalies.................................................................................................................................................................273
Vastus lateralis...............................................................................................................................................................273
Synonyms .................................................................................................................................................................273
Variations.................................................................................................................................................................. 273
Anomalies.................................................................................................................................................................274
Vastus intermedius ........................................................................................................................................................274
Synonyms ................................................................................................................................................................. 274
Variations..................................................................................................................................................................274
Anomalies.................................................................................................................................................................275
Articularis genus ...........................................................................................................................................................275
Synonyms .................................................................................................................................................................275
Variations..................................................................................................................................................................275
Anomalies.................................................................................................................................................................276
Obturator internus .........................................................................................................................................................276
Lower Limb Muscles 255
Synonyms .................................................................................................................................................................276
Variations..................................................................................................................................................................276
Anomalies.................................................................................................................................................................277
Obturator externus.........................................................................................................................................................277
Synonyms .................................................................................................................................................................277
Variations..................................................................................................................................................................277
Anomalies.................................................................................................................................................................278
Gemellus superior .........................................................................................................................................................278
Synonyms .................................................................................................................................................................278
Variations..................................................................................................................................................................278
Anomalies.................................................................................................................................................................279
Gemellus inferior ..........................................................................................................................................................279
Synonyms .................................................................................................................................................................279
Variations..................................................................................................................................................................279
Anomalies.................................................................................................................................................................280
Quadratus femoris .........................................................................................................................................................280
Synonyms .................................................................................................................................................................280
Variations..................................................................................................................................................................280
Anomalies.................................................................................................................................................................281
Gracilis ..........................................................................................................................................................................281
Synonyms .................................................................................................................................................................281
Variations..................................................................................................................................................................282
Anomalies.................................................................................................................................................................282
Adductor longus ............................................................................................................................................................282
Synonyms .................................................................................................................................................................282
Variations..................................................................................................................................................................282
Anomalies.................................................................................................................................................................283
Adductor brevis .............................................................................................................................................................283
Synonyms .................................................................................................................................................................283
Variations..................................................................................................................................................................283
Anomalies.................................................................................................................................................................283
Adductor magnus ..........................................................................................................................................................284
Synonyms .................................................................................................................................................................284

Variations..................................................................................................................................................................284
Anomalies.................................................................................................................................................................284
Pectineus .......................................................................................................................................................................285
Synonyms .................................................................................................................................................................285
Variations..................................................................................................................................................................285
Anomalies.................................................................................................................................................................285
Semitendinosus .............................................................................................................................................................285
Synonyms .................................................................................................................................................................285
Variations..................................................................................................................................................................287
Anomalies.................................................................................................................................................................287
Semimembranosus ........................................................................................................................................................287
Synonyms .................................................................................................................................................................287
Variations.................................................................................................................................................................. 288
Anomalies.................................................................................................................................................................288
Biceps femoris...............................................................................................................................................................288
Synonyms .................................................................................................................................................................288
Variations..................................................................................................................................................................289
Anomalies.................................................................................................................................................................289
Tensor fasciae suralis ....................................................................................................................................................290
Synonyms .................................................................................................................................................................290
Variations..................................................................................................................................................................290
Anomalies.................................................................................................................................................................291
Posterior Muscles of the Lower Leg ..................................................................................................................................291
Gastrocnemius...............................................................................................................................................................291
Synonyms .................................................................................................................................................................292
Variations..................................................................................................................................................................292
Anomalies.................................................................................................................................................................293
Soleus ............................................................................................................................................................................293
Synonyms .................................................................................................................................................................293
Variations..................................................................................................................................................................294
Anomalies.................................................................................................................................................................294
Peroneotibialis...............................................................................................................................................................295
Synonyms .................................................................................................................................................................295

Variations..................................................................................................................................................................295
Anomalies.................................................................................................................................................................295
Plantaris.........................................................................................................................................................................295
Synonyms .................................................................................................................................................................295
Variations..................................................................................................................................................................295
Anomalies.................................................................................................................................................................296
Popliteus........................................................................................................................................................................297
Synonyms .................................................................................................................................................................297
Variations..................................................................................................................................................................297
Anomalies.................................................................................................................................................................297
Flexor digitorum longus................................................................................................................................................298
Synonyms .................................................................................................................................................................298
Variations..................................................................................................................................................................299
Anomalies.................................................................................................................................................................300
Flexor digitorum accessorius longus.............................................................................................................................300
Synonyms .................................................................................................................................................................300
Variations.................................................................................................................................................................. 300
Anomalies.................................................................................................................................................................301
Fibulocalcaneus internus...............................................................................................................................................302
Synonyms .................................................................................................................................................................302
Variations..................................................................................................................................................................302
Anomalies.................................................................................................................................................................302
Flexor hallucis longus ...................................................................................................................................................303
Synonyms .................................................................................................................................................................303
Variations..................................................................................................................................................................303
Anomalies.................................................................................................................................................................304
Tibialis posterior ...........................................................................................................................................................304
Synonyms .................................................................................................................................................................304
Variations..................................................................................................................................................................305
Anomalies.................................................................................................................................................................305
Anterior and Lateral Muscles of the Lower Leg................................................................................................................305
Tibialis anterior .............................................................................................................................................................305
Synonyms .................................................................................................................................................................305

Variations..................................................................................................................................................................306
Anomalies.................................................................................................................................................................307
Extensor hallucis longus ...............................................................................................................................................308
Synonyms .................................................................................................................................................................308
Variations..................................................................................................................................................................308
Anomalies.................................................................................................................................................................310
Extensor digitorum longus ............................................................................................................................................ 310
Synonyms .................................................................................................................................................................310
Variations..................................................................................................................................................................310
Anomalies.................................................................................................................................................................311
Fibularis tertius..............................................................................................................................................................311
Synonyms .................................................................................................................................................................311
Variations..................................................................................................................................................................312
Anomalies.................................................................................................................................................................313
Anterior fbulocalcaneus ...............................................................................................................................................314
Synonyms .................................................................................................................................................................314
Variations.................................................................................................................................................................. 314
Anomalies.................................................................................................................................................................314
Fibularis longus.............................................................................................................................................................314
Synonyms .................................................................................................................................................................314
Variations..................................................................................................................................................................314
Anomalies.................................................................................................................................................................316
Fibularis brevis..............................................................................................................................................................317
Synonyms .................................................................................................................................................................317
Variations..................................................................................................................................................................317
Anomalies.................................................................................................................................................................317
Fibularis quartus............................................................................................................................................................318
Synonyms .................................................................................................................................................................318
Variations..................................................................................................................................................................318
Anomalies.................................................................................................................................................................319
Fibularis digiti minimi...................................................................................................................................................320
Synonyms .................................................................................................................................................................320
Variations..................................................................................................................................................................320
Anomalies.................................................................................................................................................................321
Muscles of the Foot............................................................................................................................................................321
Abductor hallucis ..........................................................................................................................................................321
Synonyms .................................................................................................................................................................321
Variations..................................................................................................................................................................321
Anomalies.................................................................................................................................................................322
Flexor digitorum brevis.................................................................................................................................................323
Synonyms .................................................................................................................................................................323
Variations..................................................................................................................................................................323
Anomalies.................................................................................................................................................................324
Abductor digiti minimi..................................................................................................................................................324
Synonyms .................................................................................................................................................................324
Variations..................................................................................................................................................................325
Anomalies.................................................................................................................................................................325
Abductor ossis metatarsi digiti quinti ...........................................................................................................................325
Synonyms .................................................................................................................................................................325
Variations.................................................................................................................................................................. 326
Anomalies.................................................................................................................................................................326
Quadratus plantae..........................................................................................................................................................326
Synonyms .................................................................................................................................................................326
Variations.................................................................................................................................................................. 327
Anomalies.................................................................................................................................................................328
Lumbricales (Lumbricals)....................................................................................................................................................328
Synonyms .................................................................................................................................................................328
Variations..................................................................................................................................................................329
Anomalies.................................................................................................................................................................330
Flexor hallucis brevis ....................................................................................................................................................330
Synonyms .................................................................................................................................................................330
Variations..................................................................................................................................................................331
Anomalies.................................................................................................................................................................331
Adductor hallucis ..........................................................................................................................................................331
Synonyms .................................................................................................................................................................331
Variations..................................................................................................................................................................331
Anomalies.................................................................................................................................................................332
Contrahentes pedis ........................................................................................................................................................333
Synonyms .................................................................................................................................................................333
Variations..................................................................................................................................................................333
Anomalies.................................................................................................................................................................333
Opponens hallucis .........................................................................................................................................................333
Synonyms .................................................................................................................................................................333
Variations..................................................................................................................................................................334
Anomalies.................................................................................................................................................................334
Flexor digiti minimi brevis............................................................................................................................................334
Synonyms .................................................................................................................................................................334
Variations..................................................................................................................................................................334
Anomalies.................................................................................................................................................................334
Opponens digiti minimi.................................................................................................................................................335
Synonyms .................................................................................................................................................................335
Variations.................................................................................................................................................................. 335
Anomalies.................................................................................................................................................................335
Interossei plantares (Plantar Interossei) ........................................................................................................................335
Synonyms .................................................................................................................................................................335
Variations..................................................................................................................................................................336
Anomalies.................................................................................................................................................................336
Interossei dorsales (Dorsal Interossei) ..........................................................................................................................336
Synonyms .................................................................................................................................................................336
Variations.................................................................................................................................................................. 337
Anomalies.................................................................................................................................................................338
Extensor digitorum brevis .............................................................................................................................................338
Synonyms .................................................................................................................................................................338
Variations..................................................................................................................................................................338
Anomalies.................................................................................................................................................................339
Extensor hallucis brevis ................................................................................................................................................339
Synonyms .................................................................................................................................................................339
Variations..................................................................................................................................................................340
Anomalies.................................................................................................................................................................340
GLUTEAL AND THIGH MUSCLES Synonyms

N/A
gluteus MaxiMus (figure 5.1)
Entry adapted by permission from Springer Nature Customer Typical Presentation
Service Centre GmbH: Springer Current Molecular Biology Description
Reports, Muscles Lost in Our Adult Primate Ancestors Gluteus maximus originates from the ilium, sacrum, coc-
Still Imprint in US: on Muscle Evolution, Development, cyx, aponeurosis of erector spinae, sacrotuberous ligament,
Variations, and Pathologies. E. Boyle, V. Mahon, R. Diogo, and the gluteal aponeurosis (Standring 2016). It inserts into
2020. the iliotibial tract and onto the gluteal tuberosity of the
See also: Tenuissimus femur (Standring 2016).
FIGURE 5.1 Gluteal muscles in posterior view. The more superfcial muscles of the gluteal region are illustrated on the right side and
the deeper muscles are illustrated on the left side.
Innervation the upper aspect of the greater trochanter or distally down

Gluteus maximus is innervated by the inferior gluteal nerve the femur (Macalister 1875; Stern 1972; Nicholson et al.
(Standring 2016). 2016). Medial fbers of gluteus maximus may have a sepa-
rate insertion onto the lateral margin of the linea aspera
Comparative Anatomy (Bergman et al. 1988). Gluteus maximus may be fused
Gluteus maximus has similar origins and attachments in with tensor fasciae latae (Bergman et al. 1988; du Plessis
the apes, but compared with humans, gluteus maximus in and Loukas 2016).
the apes is reduced in size and thickness and is more devel- Taylor et al. (2015) describe an accessory muscle in a
oped in its distal portions (Hepburn 1892; Sonntag 1924; female cadaver that was surrounded by a fascial sheath. It
Stern 1972; Sigmon 1974; Diogo et al. 2010, 2012, 2013a,b, arose from the deep fbers of the inferior portion of gluteus
2017; Ferrero 2011; Ferrero et al. 2012). maximus and converged into a tendon that had a contribu-
In the apes, it originates from the posterior iliac crest, tion from the iliotibial tract. The tendon inserted onto the
including the posterior superior iliac spine, the thoracolum- proximal aspect of the femur lateral to the intertrochanteric
bar fascia, sacrum, coccyx, sacrotuberal ligament, gluteus crest just superior to the gluteal tuberosity.
medius fascia, and ischial tuberosity, and inserts onto the An additional bundle may be present connecting gluteus
femur in the region of the gluteal tuberosity, into the apo- maximus to the ischial tuberosity (Macalister 1875; Testut
neurosis of vastus lateralis, into the iliotibial tract (when 1884; Bergman et al. 1988; Nicholson et al. 2016; Standring
present), and sometimes onto the hypertrochanteric fossa 2016). When this bundle is an independent muscle, it is
(Champneys 1872; Hepburn 1892; Beddard 1893; Sonntag referred to as ischiofemoralis. This muscle is homologous
1924; Raven 1950; Stern 1972; Sigmon 1974; Brown 1983; to the ischiofemoralis of orangutans (Brown 1983; Ferrero
Gibbs 1999; Diogo et al. 2010, 2012, 2013a,b, 2017; Ferrero 2011, Ferrero et al. 2012; Diogo et al. 2013b) and is likely
2011; Ferrero et al. 2012; Standring 2016). equivalent with, or a variant of, coccygeofemoralis.
The ischial origin of gluteus maximus is recognized as a When present, coccygeofemoralis extends from the
distinct, well-developed muscle in orangutans and is present sacrum and coccyx to the femur just at the lower border of
but frequently blended with gluteus maximus in the other gluteus maximus and may represent a caudal head of this
apes (Ferrero et al. 2012; Diogo et al. 2010, 2012, 2013a,b, muscle (Macalister 1875; Bergman et al. 1988; Nicholson
2017). When present, the ischiofemoralis muscle originates et al. 2016). The adult human coccygeofemoralis (and
from the ischial tuberosity and inserts on the lateral aspect potentially the adult human ischiofemoralis) derives from
of the proximal femoral shaft and the aponeurosis of vastus the embryonic coccygeofemoralis, which originates from
lateralis, with some fbers going to the tensor fasciae latae the coccyx and inserts onto the femur near the distal mar-
(Ferrero et al. 2012; Diogo et al. 2010, 2012, 2013a,b, 2017). gin of gluteus maximus and is innervated by a branch of
The insertion of gluteus maximus inserts more dis- the inferior gluteal nerve. This muscle persists as a dis-
tally onto the femur in the apes than in humans, extend- tinct structure until CR40–45 mm (crown-rump length of
ing two-thirds down the femur in gibbons and orangutans 40–45 mm) and then fuses with gluteus maximus (Tichý
(Stern 1972; Sigmon 1974), four-ffths down the femur in and Grim 1985). As noted by Morimoto (2018), the pres-
siamangs (Hepburn 1892; Sigmon 1974), and almost to the ence of both gluteus maximus and coccygeofemoralis is
distal end of the femur in common chimpanzees and bono- similar to the confguration seen in adult apes, in which
bos (Hepburn 1892; Dwight 1895; MacDowell 1910; Miller gluteus maximus consists of a proximal portion (gluteus
1952; Sigmon 1974; Diogo et al. 2013a, 2017). Gluteus max- maximus proprius) and a distal portion (ischiofemoralis).
imus may also be divided into multiple bellies or bundles in However, Tichý and Grim (1985) suggest that there is
all apes (Hepburn 1892; Sonntag 1924; Gibbs 1999; Diogo little evidence that coccygeofemoralis corresponds to any
et al. 2012, 2013a,b, 2017). muscle in normal adult humans. They note that Testut (1884)
described a coccy-femoralis muscle, and Le Double (1897)
Variations described a femoro-coccygeus muscle. Testut (1884) sug-
Description gests coccy-femoralis is homologous to caudofemoralis in
Gluteus maximus may be split into two laminae (Macalister other mammals, and Tichý and Grim (1985) argue that their
1875; Knott 1883b; Le Double 1897; Standring 2016). It coccygeofemoralis corresponds to this structure in adults
may also be doubled (Macalister 1867b; Nicholson et al. with incomplete fusion of this muscle and gluteus maximus.
2016). The extent of the origin from the sacrum, coccyx, Kirici and Ozan (1999) report a bilateral variation of glu-
or gluteal aponeurosis may vary (Macalister 1875; Le teus maximus in which the muscles had a fbrous sacroiliac
Double 1897; Knott 1883b; Stern 1972; Nicholson et al. part that was separated from a muscular coccygeofemoral
2016). Gluteus maximus may have additional origins from part and was associated with a doubled piriformis muscle
the thoracolumbar fascia, multifdus fascia, sacroiliac lig- on the right side. On both sides of the body, coccygeofemo-
aments, aponeurosis of iliocostalis lumborum, or aponeu- ralis originated from the base of the coccyx. On the right
rosis of latissimus dorsi (Macalister 1875; Knott 1883b; side, tendons arose from this muscle, one inserting into the
Testut 1884; Stern 1972; Nicholson et al. 2016; Standring linea aspera 5 cm below the greater trochanter, and the other
2016). The femoral insertion may extend proximally to inserting onto the origin of the short head of biceps femoris
11 cm below the grater trochanter. On the left side, the muscle inferior gluteal nerve and lead to weakness in gluteus maxi-
inserted onto the gluteal tuberosity 7.5 cm below the greater mus. The accessory muscle described by Taylor et al. (2015)
trochanter. These authors conclude that this morphology (see above) may have caused pain and other symptoms simi-
results from the failure of the fusion of gluteus maximus pro- lar to those of Greater Trochanteric Pain Syndrome (GTPS).
prius (the sacroiliac part) and the coccygeofemoralis muscle
during embryonic development (Kirici and Ozan 1999).
gluteus MeDius (figure 5.1)
Prevalence
Synonyms
Knott (1883b) found that in 5 out of 40 subjects (12.5%), a
thin deep lamina was formed by the gluteus maximus fbers N/A
that originated from the sacrosciatic and posterior sacroiliac
ligaments and was separated by the superfcial portion of the Typical Presentation
muscle by a layer of connective tissue. An accessory slip from Description
the lumbar aponeurosis was also present in fve subjects (total Gluteus medius originates from the ilium between the pos-
sample size not specifed). Taylor et al. (2015) found an acces- terior gluteal line and the anterior gluteal line (Standring
sory muscle extending from the deep inferior fbers of gluteus 2016). It inserts via a tendon onto the lateral aspect of the
maximus to the proximal femur in 1 out of 77 hips (1.3%). greater trochanter (Standring 2016).
Description Gluteus medius is innervated by the superior gluteal nerve
Gluteus maximus may be absent as a rare congenital anom- (Standring 2016).
aly (Tagliapietra et al. 1989). On the right side of a fetus
with craniorachischisis dissected by Alghamdi et al. (2017), Comparative Anatomy
gluteus maximus was diffcult to separate from gluteus Gluteus medius has a similar typical presentation in the
medius. Gluteus maximus may also have extra slips or ten- apes, with similar variations including fusion with nearby
dons (Smith et al. 2015). Pirani et al. (1991) describe soft muscles and tendons (Beddard 1893; Sigmon 1974; Raven
tissue anatomy associated with cases of proximal femoral 1950; Diogo et al. 2010, 2012, 2013a,b, 2017). It is often
focal defciency (PFFD). In Aitken type D PFFD, the glu- fused with piriformis in gibbons and bonobos (Beddard
teal muscles insert onto the proximal femoral ossicle. 1893; Diogo et al. 2017). It may be fused with piriformis,
Bersu et al. (1976) describe a male infant with Hanhart vastus lateralis, or the tensor fasciae latae in orangutans
syndrome. The femora of this infant were normally devel- (Diogo et al. 2013b). Its tendon may be fused with that of
oped, but distal secondary ossifcation centers were absent. piriformis in common chimpanzees (Champneys 1872;
The left leg stump had a patella and a small rudiment of the Hepburn 1892; Dwight 1895).
proximal tibia but no fbular rudiment. The right leg stump
was less developed and had a patella, smaller tibial rudi- Variations
ment, and no fbular rudiment. On the left side, gluteus max- Description
imus had a normal origin but did not insert onto the gluteal Gluteus medius may be divided into two lamellae (Henle
tuberosity. It had a normal attachment to the fascia lata but 1858; Macalister 1875; Bergman et al. 1988) or into mul-
also had a tendinous band that inserted onto the middle third tiple muscle layers (Akita et al. 1993). The extent of the
of the linea aspera. The right gluteus maximus had a normal origin from the ilium and the specifc location of its inser-
origin but inserted via a tendon onto the distal half of the tion onto the greater trochanter may vary (Flack et al. 2012;
linea aspera. A small accessory muscle originated from the Nicholson et al. 2016). Deep fbers from gluteus medius
deep surface of the right gluteus maximus near its origin may have a separate attachment to the upper border of the
from the sacrum and inserted onto the ischial tuberosity. greater trochanter (Bergman et al. 1988; Standring 2016).
Gluteus medius accessorius may originate from the iliac
Prevalence crest and insert onto the tendon of gluteus medius and the
In the literature review conducted by Smith et al. (2015), greater trochanter (Henle 1858; Jazuta 1931; Bergman et al.
they found that extra slips or tendons of gluteus maximus 1988; Nicholson et al. 2016).
were present in only 1 out of 20 individuals with trisomy The posterior border of gluteus medius may blend
13 (5%). with piriformis or send slips to its tendon (Henle 1858;
Macalister 1875; Bergman et al. 1988; Akita et al. 1994;
Clinical Implications Flack et al. 2014; Nicholson et al. 2016; Standring 2016).
Variations in the gluteal muscles including division of the Gluteus medius may also be partially or completely fused
muscle into multiple parts should be acknowledged when with gluteus minimus or its tendon (Macalister 1875; Knott
considering injections into the gluteal musculature (Kirici 1883b; Testut 1884; Akita et al. 1993; Duparc et al. 1997;
and Ozan 1999). Arora et al. (2010) note that accessory slips Flack et al. 2014, Nicholson et al. 2016). The insertion ten-
related to gluteus maximus or piriformis may compress the don of gluteus medius may also join with the tendon of
vastus lateralis (Nazarian et al. 1987; Heimkes et al. 1992; was less developed and had a patella, smaller tibial rudiment,
Nicholson et al. 2016). and no fbular rudiment. On the left side, gluteus medius was
Heimkes et al. (1992) studied the relationships upon inser- normal. On the right side, gluteus medius had a normal ori-
tion between gluteus medius, gluteus minimus, and vastus gin but had an additional insertion onto the gluteal tuberos-
lateralis and classifed these relationships into fve types. In ity. A slip also extended from the posteromedial border of
type I, gluteus medius, gluteus minimus, and vastus lateralis this muscle to the superolateral border of piriformis.
formed an ideal joint tendinous junction. In type II, vastus Pirani et al. (1991) describe soft tissue anatomy associ-
lateralis is connected to gluteus medius and gluteus minimus ated with cases of PFFD. In Aitken type B PFFD, gluteus
while gluteus medius and gluteus minimus were not con- medius and gluteus minimus are shorter than normal and
nected with each other. In type III, vastus lateralis connects extend between the ilium and the proximal femoral metaph-
only to gluteus minimus and gluteus medius has an indepen- ysis. This latter structure indents these muscles from below.
dent insertion onto the greater trochanter. In type IV, vastus In Aitken type D PFFD, the gluteal muscles insert onto the
lateralis connects only to gluteus medius and gluteus mini- proximal femoral ossicle.
mus has an independent insertion onto the greater trochan-
ter. In type V, all three muscles insert independently onto the Prevalence
greater trochanter without connections to each other. N/A

Heimkes et al. (1992) studied the relationships upon inser- Acknowledging the variable relationships between glu-
tion between gluteus medius, gluteus minimus, and vastus teus medius, gluteus minimus, and vastus lateralis and the
lateralis in 52 hip specimens. See above for the description potential independent insertions of these muscles onto the
of the fve types of relationships they identifed. Type I was greater trochanter is important for successful hip opera-
found in 18 cases (34.6%), type II in 13 cases (25%), type tions, particularly when using the transgluteal approach
III in 11 cases (21.1%), type IV in seven cases (13.5%), and (Heimkes et al. 1992).
type V in three cases (5.8%).
Akita et al. (1993) studied gluteus medius in 13 hemipel-
gluteus MiniMus (nOt illustrateD)
ves from 7 cadavers. The anteromedial portion of gluteus
medius was comprised of one muscle layer in six specimens See also: Scansorius
(46%), two independent muscle layers in four specimens
(31%), and three muscle layers in three specimens (23%). In Synonyms
12 specimens (92.3%), the inferior aspect of the most antero- N/A
medial independent muscle layer inserted onto the gluteus
minimus aponeurosis. Flack et al. (2014) studied the mor- Typical Presentation
phology of gluteus medius in 12 lower limbs. In 100% of Description
specimens, the tendons of gluteus medius and gluteus mini- Gluteus minimus originates from the ilium between the
mus fused as they approached the greater trochanter. In four anterior gluteal line and inferior gluteal line and from the
cases (33.3%), gluteus medius fused to piriformis posteriorly. border of the greater sciatic notch (Standring 2016). It inserts
via a tendon onto the greater trochanter (Standring 2016). Its
Anomalies insertion expands to the hip joint capsule (Standring 2016).
Description
Innervation
On the right side of a fetus with craniorachischisis, glu-
teus medius fused with tensor fasciae latae and was dif- Gluteus minimus is innervated by the superior gluteal nerve
fcult to separate from gluteus maximus (Alghamdi et al. (Standring 2016).
2017). On both sides of this specimen, the origin of the
Comparative Anatomy
muscle extended posteriorly so it was wider than normal
(Alghamdi et al. 2017). In a female fetus with trisomy 18, Gluteus minimus has a similar typical presentation in the
gluteus medius fused with tensor fasciae latae bilaterally apes, extending from the ilium to the greater trochanter and
(Alghamdi et al. (2018). In a male neonate with Meckel syn- hip joint capsule (Champneys 1872; Hepburn 1892; Sonntag
drome, gluteus medius originated from the ilium below its 1924; Raven 1950; Miller 1952; Sigmon 1974; Gibbs 1999;
normal origin and did not extend above the upper margin of Diogo et al. 2010, 2012, 2013a,b, 2017).
gluteus maximus (Pettersen 1984).
Bersu et al. (1976) describe a male infant with Hanhart Variations
syndrome. The femora of this infant were normally devel- Description
oped but distal secondary ossifcation centers were absent. Gluteus minimus can be divided into an anterior part and pos-
The left leg stump had a patella and a small rudiment of the terior part (Macalister 1875; Bergman et al. 1988; Standring
proximal tibia but no fbular rudiment. The right leg stump 2016). The specifc location of its insertion onto the greater
trochanter may vary (Flack et al. 2012; Nicholson et al. 2016). Typical Presentation
The insertion onto the hip joint capsule may present as one Description
or more detached muscular bundles or fascicles (Macalister
Piriformis originates from the anterior aspect of the sacrum
1875; Bergman et al. 1988; Flack et al. 2014).
via three digitations, the ilium near the posterior inferior
Gluteus minimus may send slips to gemellus supe-
iliac spine, and from the capsule of the sacroiliac joint
rior, piriformis, tensor fasciae latae, or vastus lateralis
(Standring 2016). It courses through the greater sciatic fora-
(Macalister 1875; Bergman et al. 1988; Nicholson et al.
men to insert onto the greater trochanter (Standring 2016).
2016; Standring 2016). Gluteus medius may be partially
fused, or more rarely completely fused, with the muscle Innervation
belly or tendon of gluteus minimus (Macalister 1875; Knott Piriformis is innervated by the frst two sacral spinal nerves,
1883b; Testut 1884; Akita et al. 1993; Duparc et al. 1997; and sometimes the ffth lumbar spinal nerve (Standring 2016).
Flack et al. 2014, Nicholson et al. 2016). Gluteus minimus
may also be completely or partially fused with piriformis Comparative Anatomy
or its tendon (Knott 1883b; Flack et al. 2014). The insertion Piriformis has a similar typical presentation in the apes,
tendon of gluteus minimus may also join with the proximal extending from the sacrum, and often the ilium, to the
tendon of vastus lateralis (Nazarian et al. 1987; Heimkes greater trochanter (Raven 1950; Miller 1952; Gibbs 1999;
et al. 1992; Flack et al. 2014). Heimkes et al. (1992) studied Diogo et al. 2010, 2012, 2013a,b, 2017). It may be fused
the relationships upon insertion between gluteus medius, with gluteus medius in gibbons, orangutans, and common
gluteus minimus, and vastus lateralis and classifed these chimpanzees (Champneys 1873; Hepburn 1892; Beddard
relationships into fve types. See gluteus medius entry for 1893; Dwight 1895; Sonntag 1924; Sigmon 1974).
more details and prevalence information.
Variations
Prevalence
Description
Flack et al. (2014) studied the morphology of gluteus mini-
mus in 12 lower limbs. In 100% of specimens, the ten- Piriformis may have an origin from the upper part of the
dons of gluteus medius and gluteus minimus fused as they pelvic aspect of the sacrotuberous ligament (Standring
approached the greater trochanter. In four cases (33.3%), the 2016). Its origin from the sacrum may span from one or two
posterior border of gluteus minimus was fused with the ten- segments up to fve segments and may include the coccyx
don of piriformis. Atrophy of gluteus minimus was observed (Macalister 1875; Knott 1883b; Le Double 1897; Bergman
in eight specimens (66.7%), six cases mild and two extensive. et al. 1988; Rickenbacher et al. 1985; Nicholson et al. 2016).
The digitations of piriformis can range from two to four
Anomalies (Bergman et al. 1988). Piriformis may have an insertion onto
Description the hip joint capsule (Macalister 1875; Bergman et al. 1988).
Piriformis may be partially or entirely absent (Macalister
Pirani et al. (1991) describe soft tissue anatomy associ-
1875; Rickenbacher et al. 1985; Bergman et al. 1988; Gibbs
ated with cases of PFFD. In Aitken type B PFFD, gluteus
1999; Ikidag 2019; Caetano and Seeger 2019). Brenner et al.
medius and gluteus minimus are shorter than normal and
(2019) describe the right lower limb of a female cadaver in
extend between the ilium and the proximal femoral metaph-
which piriformis and inferior gemellus were missing.
ysis. This latter structure indents these muscles from below.
Piriformis may be fused with or receive a slip from gluteus
In Aitken type D PFFD, the gluteal muscles insert onto the
medius or its tendon, gluteus minimus or its tendon, supe-
proximal femoral ossicle.
rior gemellus, or obturator internus (Henle 1858; Macalister
Prevalence 1875; Knott 1883b; Le Double 1897; Rickenbacher et al.
N/A 1985; Bergman et al. 1988; Chiba 1992; Akita et al. 1994;
Flack et al. 2014; Nicholson et al. 2016; Standring 2016). It
Clinical Implications may also join with coccygeus (Macalister 1875). The ten-
Acknowledging the variable relationships between glu- don of insertion may be blended to some degree with the
teus medius, gluteus minimus, and vastus lateralis and the common tendon of obturator internus, gemellus superior,
potential independent insertions of these muscles onto the and gemellus inferior (Macalister 1875; Pine et al. 2011;
greater trochanter is important for successful hip opera- Standring 2016) or with the insertion tendon of gluteus
tions, particularly when using the transgluteal approach minimus or gluteus medius (Macalister 1875; Flack et al.
(Heimkes et al. 1992). 2014). The tendon of gemellus superior may insert onto the
piriformis tendon instead of the tendon of obturator inter-
nus (Macalister 1875; Knott 1883b).
pirifOrMis (figures 4.4 anD 5.1) Piriformis may split into two portions, which is often
associated with a high division of the sciatic nerve, often
Synonyms with the peroneal (common fbular) component of the
N/A nerve passing between the two portions and the tibial nerve
passing inferior to the muscle (type B variation accord- piriformis near the upper border of the greater sciatic notch
ing to Beaton and Anson) (Macalister 1875; Rickenbacher was separated by an areolar layer from the sacral part of
et al. 1985; Bergman et al. 1988; Chiba 1992; Natsis et al. piriformis. In three cases (7.5%), piriformis was split by the
2014; Nicholson et al. 2016). Other relationships between greater sciatic nerve. Pine et al. (2011) found that out of 29
piriformis and the sciatic nerve include the normal passage specimens, the tendons of obturator internus and piriformis
of an undivided sciatic nerve below piriformis (type A, were at least partially fused in four specimens (13.8%), with
typical presentation) or above piriformis (type F), the com- three cases (10.3%) demonstrating insertion of the two as
mon fibular nerve passing above piriformis and the tibial a common tendon, while in one case (3.4%) the combined
nerve passing inferior to it (type C), the entire sciatic nerve tendon divided to have separate attachments. Flack et al.
passing through a split piriformis (type D), and the com- (2014) studied the morphology of gluteus medius and glu-
mon fibular nerve passing above a split piriformis and the teus minimus in 12 lower limbs. In four cases each (33.3%),
tibial nerve passing through the muscle (type E) (Beaton gluteus medius fused to piriformis posteriorly and gluteus
and Anson 1937; Nicholson et al. 2016). In a rare case, the minimus was fused with the tendon of piriformis.
sciatic nerve was observed to travel posterior to piriformis Piriformis may be split by the peroneal component
(Sayson et al. 1994). Nicholson et al. (2016) provide a com- of the sciatic nerve in 12.2% of the “normal” population
prehensive review of these relationships and information on (Mortensen and Pettersen 1966; Bersu 1980). Sato (1970)
the relationships between piriformis and other neurovascu- found that piriformis was split into two portions by the
lar structures. Doubling or division of piriformis may also sciatic nerve in 88 out of 418 limbs (21%) from Kyushu
be associated with doubling or division of the gemelli, qua- Japanese males and in 54 out of 268 limbs (20%) from
dratus femoris, or gluteus maximus (Kirici and Ozan 1999; females. Chiba (1992) found that piriformis was split
Nicholson et al. 2016). into three muscle bellies by the branching patterns of the
Kirici et al. (2000) report a double piriformis in left sciatic nerve in 5 out of 514 cases (1%). This author also
lower limbs. In the first case, the common fibular nerve found that when two bundles of piriformis are present, the
passed between the two portions of piriformis. In the sec- tendon of the lower one blends with gemellus superior in
ond case, the common fibular and posterior femoral cutane- five cases (1%).
ous nerves passed between the two portions. This second Natsis et al. (2014) studied the morphology of piri-
case was associated with the passage of the gemelli, obtura- formis on 294 sides from 147 Greek cadavers. Type A
tor internus, pudendal nerve, and vessels behind the mid- anatomy was present on 275 sides (93.6%), and variations
portion of the sacrotuberous ligament. were present in 19 cases (6.4%). Type B was present on 12
An accessory muscle may be present superior to piri- sides (4.1%) and types C, D, and F each on one side (0.3%
formis and may have origins from the sacrotuberous liga- each). These numbers are included in the comprehensive
ment, the greater sciatic foramen, or the posterior end of review by Nicholson et al. (2016, see next paragraph). Four
the ilium, and insertions into the greater trochanter, hip limbs (1.4%) exhibited other variations. In one case (0.3%),
joint capsule, or piriformis tendon (Prasad et al. 2005; piriformis was divided into three muscle bellies, with the
Tubbs and Salter 2006b; Ravindranath et al. 2008; Natsis common fibular nerve passing between the superficial and
et al. 2014; Nicholson et al. 2016). An accessory digas- intermediate muscle bellies and the deep muscle belly pass-
tric muscle may also be present, with a superior belly that ing through a branch of the tibial nerve. In another case
arises from or near piriformis and then continues down (0.3%), the common fibular nerve passed between the
the thigh as a long intermediate tendon, before ending in superficial and deep muscle bellies of a double piriformis
an inferior belly (Moore 1922; Suda and Takahashi 1957; and the tibial nerve passed below piriformis. Lastly, on
Akita et al. 1992; Nicholson et al. 2016). Arora et al. (2010) both sides of one cadaver (0.7%), the sciatic nerve traveled
describe a 5 cm long accessory slip that originated from below piriformis but there was a bilateral supernumerary
the inferomedial portion of gluteus maximus and merged muscle just superior to piriformis, extending from the pos-
with piriformis near the greater trochanter. Although its terior margin of the ilium and inserting onto the greater
fibers were parallel to those of piriformis, thus appearing trochanter. These authors also found that out of the 12 sides
as a second piriformis muscle, the authors suggest that it that exhibited split piriformis muscles, five of them (41.7%)
derived primarily from the gluteus maximus muscle due were positioned as a superficial belly and deep belly and
to its innervation from a branch of the inferior gluteal seven (58.3%) were positioned as a superior belly and infe-
nerve. They term this slip the “gluteopiriformis” muscle rior belly.
(Arora et al. 2010). Nicholson et al. (2016) summarize information about
the relationships between piriformis and the sciatic nerve
Prevalence in 6,466 cases from 43 studies. The normal passage of an
From the examination of 36 bodies, Wood (1868) states that undivided sciatic nerve below piriformis, type A or the
in three males and one female (11%), the tendon of pirifor- typical presentation, was present in 5,392 cases (83.4%).
mis was blended with that of obturator internus and the two Type B variation was present in 827 cases (12.8%), type C
inserted together onto the greater trochanter. Knott (1883b) in 87 cases (1.3%), type D in 34 cases (0.5%), type E in 5
noted that in 2 out of 40 cases (5%), the iliac origin of cases (0.08%), and type F in 7 cases (0.1%). Nicholson et al.
(2016) also review the insertion of piriformis and found that et al. (2019) suggest that since piriformis is an anatomical
when separated into two muscle bellies, each belly has a landmark for ultrasound investigations, hip surgeries, and
tendon that blends with the other to have a common inser- other procedures in the deep gluteal region, its absence can
tion in 41 out of 57 cases (71.9%). infuence orientation in the deep gluteal region and affect
the identifcation of target structures.
Anomalies
Description
scansOrius (figure 5.1)
Anomalous presentations of piriformis include variable
insertions, the presence of accessory slips, complete Entry adapted by permission from Springer Nature Customer
absence, or splitting of the muscle belly by the peroneal Service Centre GmbH: Springer Current Molecular Biology
portion of the sciatic nerve (Bersu et al. 1976; Bersu Reports, Muscles Lost in Our Adult Primate Ancestors
1980; Pirani et al. 1991; Smith et al. 2015). Bersu et al. Still Imprint in US: on Muscle Evolution, Development,
(1976) describe a male infant with Hanhart syndrome. Variations, and Pathologies. E. Boyle, V. Mahon, R. Diogo,
The femora of this infant were normally developed but 2020.
distal secondary ossifcation centers were absent. The left See also: Gluteus minimus
leg stump had a patella and a small rudiment of the proxi-
mal tibia but no fbular rudiment. The right leg stump was Synonyms
less developed and had a patella, smaller tibial rudiment, This muscle is also referred to as gluteus quartus (Haughton),
and no fbular rudiment. On the left side, piriformis had invertor femoris (Owen) (Macalister 1875; Knott 1883a,b;
a normal origin and inserted onto the greater trochanter. Sutton 1888), anterior gluteus minimus (Le Double 1897;
It sent a tendinous slip to the tendon of obturator exter- Testut 1884), the fourth gluteus, marginal gluteus or small
nus. On the right side, piriformis had a normal origin and anterior gluteus (Ochiltree 1912).
inserted onto the posterior surface of the capsule of the
hip joint. It sent a fascial slip to the lateral aspect of the
ischial tuberosity. This muscle is only present as a variation.
Pirani et al. (1991) describe soft tissue anatomy asso-
Comparative Anatomy
ciated with cases of proximal femoral focal defciency
(PFFD). In Aitken type A PFFD, the short external rotators Scansorius is always present in orangutans, frequently
of the hip are larger in cross-sectional diameter than normal present in bonobos, and occasionally present in gorillas,
and insert onto the posteromedial surface of the greater tro- gibbons, and common chimpanzees (Champneys 1872;
chanter. In Aitken type B PFFD, the short external rotators Hepburn 1892; Beddard 1893; Sonntag 1923; Raven 1950;
are smaller than normal. Miller 1952; Sigmon 1974; Gibbs 1999; Ferrero et al.
2012; Diogo et al. 2010, 2012, 2013a,b, 2017). When pres-
Prevalence ent, it extends from the ilium, including the anterior supe-
In their literature review, Smith et al. (2015) found that piri- rior iliac spine, and sometimes from the fascia lata, and
formis was absent in 1 out of 17 individuals with trisomy inserts onto the greater trochanter, occasionally blending
18 (5.9%). In four out of fve individuals with trisomy 21 with gluteus minimus (Champneys 1872; Hepburn 1892;
(80%), piriformis was split by the peroneal portion of the Beddard 1893; Sonntag 1923; Raven 1950; Miller 1952;
sciatic nerve (as described by Bersu 1980). Per limb, this Sigmon 1974; Gibbs 1999; Ferrero et al. 2012; Diogo et al.
morphology was found in fve out of ten limbs (50%) (Bersu 2010, 2012, 2013a,b, 2017).
1980).
Variations
Knott (1883b) noted that in two cases where the sacral Scansorius (Traill) is typically not present in adult humans
digitations of piriformis were reduced to two, these cases (Sutton 1888; Ferrero et al. 2012). When it is present, the
were associated with chronic rheumatoid arthritis on the muscle is represented by a bundle considered to derive
same side. Variation in how the sciatic nerve relates to piri- from the anterior portion of gluteus minimus, or origi-
formis may entrap/compress the sciatic nerve (piriformis nates from between gluteus minimus and gluteus medius
syndrome) (Beaton and Anson 1938; Sayson et al. 1994; (Macalister 1875; Knott 1883b; Sutton 1888; Bergman
Natsis et al. 2014; Nicholson et al. 2016). Accessory slips et al. 1988; Gibbs 1999). It originates from the anterior
or doubling of piriformis can also compress the sciatic, superior iliac spine, iliac crest, and sometimes the ante-
inferior gluteal, or pudendal nerves (Kirici and Ozan 1999; rior margin of gluteus minimus and/or the fascia lata and
Ravindranath et al. 2008; Arora et al. 2010) or contribute to inserts onto the greater trochanter (Macalister 1867b,
superior gluteal nerve entrapment syndrome (Rask 1980). 1875; Knott 1883a,b; Testut 1884; Gruber 1887; Sutton
The absence of piriformis on the right side of a patient 1888; Ochiltree 1912; Bergman et al. 1988; Gibbs 1999, du
examined by Ikidag (2019) allowed for the migration of the Plessis and Loukas 2016; Nicholson et al. 2016; Orthaber
vermiform appendix to the deep gluteal region. Brenner et al. 2020).
It can insert independently onto the greater trochanter or Comparative Anatomy

insert with the tendon of gluteus minimus (Knott 1883b). Among primates, tenuissimus is present in some Central and
Knott (1883b) also notes that scansorius may insert par- South American monkeys (e.g., capuchins) but absent in the
tially onto the side of vastus lateralis. Macalister (1875) lemurs, African and Asian monkeys, and apes (Waterman
notes that when scansorius is present its tendon may either 1929). Green (1931) notes that in most mammals that pos-
be continuous with vastus lateralis or be overlapped by it sess a tenuissimus, the muscle usually originates from the
with the vastus lateralis tendon forming an arch over the coccyx, but the origin in monkeys, as observed by Klaatsch
muscle. Ochiltree (1912) states that the insertion of scanso- (1900), is similar to the origin observed in humans, namely
rius corresponds to the normal insertion of gluteus medius the posterior surface of gluteus maximus and its fascia.
and can displace the insertion of the latter more dorsally When present, the insertion in monkeys is onto the shaft of
and proximally than is typical. Sutton (1888) argues that the the fibula (Green 1931).
outer portion of the ilio-femoral band is the fibrous repre-
sentative of scansorius as (1) the iliofemoral ligament cor- Variations
responds in its attachments to scansorius and (2) monkeys
Description
that have a functional scansorius do not have an iliofemoral
band. Orthaber et al. (2020) describe a right lower limb that There are very few descriptions of muscles that research-
presented with both gluteus medius accessorius and gluteus ers believe correspond to tenuissimus in humans. Green
quartus. Gluteus quartus extended from the anterior por- (1931) describes one case in the right thigh of a cadaver.
tion of gluteus minimus and the fascia lata to the tip of the Tenuissimus presented as a fusiform muscle that arose via
greater trochanter. an expanded aponeurosis and subsequent tendon from the
fascia covering the deep surface of gluteus maximus. The
Innervation muscle belly descended the thigh and inserted onto biceps
Scansorius is innervated by the superior gluteal nerve femoris at the junction of the short head and the long head,
(Sutton 1888; Orthaber et al. 2020). with a slightly more extensive attachment to the long head.
Green (1931) mentions a similar case that was described
Prevalence by Wood (1867b) as a third head of biceps femoris. It arose
Duparc et al. (1997) found a muscle bundle extending via a rounded tendon from the fascia on the deep surface of
from the anterior superior iliac spine to the anterior mar- gluteus maximus and inserted onto the long head of biceps
gin of the greater trochanter in 3 out of 40 cases from a femoris near its junction with the short head.
sample of 24 cadavers (7.5%). Beck et al. (2000) noted Arakawa et al. (2017) describe a case in which they
the presence of scansorius in 2 out of 16 cadaveric hips conclude that tenuissimus, tensor fasciae suralis, and two
(12.5%). Woyski et al. (2012) report a higher prevalence other supernumerary muscles are present on the posterior
for scansorius in humans. Based on examination of 45 right thigh of a cadaver. Tenuissimus passed between the
cadaveric hips (29 female, 16 male), these authors found long and short heads of biceps femoris and fused with the
fibers corresponding to scansorius in 41% of female speci- tensor fasciae suralis, which fused with the posterior sur-
mens and 44% of male specimens. Scansorius was present face of the crural fascia that covered the lateral head of
as a distinct bundle in 50% of males and 31% of females. gastrocnemius. It is possible that the cases described here
The overall prevalence of scansorius was 80% (Woyski may be variants of femorococcygeus or coccygeofemora-
et al. 2012). lis (see section on variations for gluteus maximus) (Green
1931). It is also possible that the accessory digastricus
Anomalies muscle often associated with piriformis may be a variant
N/A of tenuissimus (see section on variations for piriformis)
(Arakawa et al. 2017).
Innervation
Orthaber et al. (2020) suggest that the presence of scanso-
rius may contribute to instability in the hip joint on the con- Tenuissimus is innervated by a branch of the common fibu-
tralateral side. lar nerve (Green 1931; Arakawa et al. 2017).
Prevalence
Tenuissimus (Figure 5.1) N/A
See also: Piriformis, Gluteus maximus, Biceps femoris
Anomalies
This muscle may also be referred to as gluteo-cruralis On the left side of one female neonate with trisomy 18,
(Klaatsch 1900; Waterman 1929). Ramirez-Castro and Bersu (1978) found the tenuissimus
muscle originating from the coccyx and inferior margin of
Typical Presentation gluteus maximus, descending along the posterior aspect of
This muscle is only present as a variation or anomaly. the thigh, and inserting into the edge of the lateral condyle
of the tibia. Aziz (1979) found the muscle on the right side aspera. It inserted onto the tibia just posterior to the inser-
of another female neonate with trisomy 18, which presented tion of the long head of the biceps femoris.
deep to gluteus maximus and originated from the sacrum,
coccyx, and sacrotuberous ligament and descended the leg Prevalence
to join the iliotibial tract. In their literature review, Smith et al. (2015) found that
Bersu et al. (1976) describe a male infant with Hanhart tenuissimus was only present in the two cases of trisomy 18
syndrome. The femora of this infant were normally devel- mentioned above, having a prevalence of 2 out of 17 (11.8%)
oped but distal secondary ossifcation centers were absent. individuals with trisomy 18.
The left leg stump had a patella and a small rudiment of
the proximal tibia but no fbular rudiment. The right leg Clinical Implications
stump was less developed and had a patella, smaller tibial N/A
rudiment, and no fbular rudiment. On the left side, tenui-
ssimus originated from the fascia of the deep surface of tensOr fasciae latae (figure 5.2)
gluteus maximus, continued as a slender muscle belly, and
joined with the long head of the biceps femoris. On the right Synonyms
side, tenuissimus was larger and originated from the fascia This muscle may also be referred to as tensor vaginae femo-
near the insertion tendon of gluteus maximus onto the linea ris (Macalister 1875).
FIGURE 5.2 Anterior thigh muscles in anterior view. The deep anterior thigh muscles are illustrated on the right side and the super-
fcial muscles are illustrated on the left side.
Typical Presentation was less developed and had a patella, smaller tibial rudi-
Description ment, and no fbular rudiment. On both sides of the body,
tensor fasciae latae originated from the fascia at the ante-
Tensor fasciae latae originates from the iliac crest, the ante-
rior border of gluteus medius and had normal insertions.
rior superior iliac spine, and from the fascia lata (Standring
2016). It ends in the upper third of the thigh and inserts via Prevalence
the iliotibial tract onto the tibia (Standring 2016).
N/A
Innervation
Tensor fasciae latae is innervated by the superior gluteal
Unilateral hypertrophy of the tensor fasciae latae can simu-
late a soft tissue tumor (Ilaslan et al. 2003).
Comparative Anatomy
Tensor fasciae latae has a similar typical presentation in the sartOrius (figure 5.2)
apes and has similar variations, including an origin from Synonyms
the gluteal fascia in gibbons and common chimpanzees and
Melling and Zweymüller (1996) refer to a distally divided
occasional fusion with the gluteal muscles in all species
sartorius muscle as sartorius bicaudatus. A proximally
(Hepburn 1892; Raven 1950; Sigmon 1974; Gibbs 1999;
divided sartorius may be referred to as biceps sartorius
Diogo et al. 2010, 2012, 2013a,b, 2017). This muscle may
(Kim and Lee 2019).
be reduced or absent in orangutans and bonobos (Chapman
1880; Sigmon 1974; Diogo et al. 2013b, 2017). Typical Presentation
Description Sartorius originates from the anterior superior iliac spine
(Standring 2016). It inserts onto the medial surface of the
Tensor fasciae latae may be absent (Bergman et al. 1988;
proximal tibia via an aponeurosis (Standring 2016). The
du Plessis and Loukas 2016). The origin of tensor fasciae
aponeurosis forms the pes anserinus with the insertions of
latae may extend to the fascia superfcial to gluteus medius
gracilis and semitendinosus (Standring 2016). It sends a slip
(Standring 2016). Its origin may also be shorter than
to the knee joint capsule and another slip to the deep fascia
is typical and not reach the anterior superior iliac spine
on the medial side of the leg (Standring 2016).
(Macalister 1875; Knott 1883a). The muscular portion of
tensor fasciae latae may extend as far distally as the lateral Innervation
femoral condyle (Standring 2016). Tensor fasciae latae may
Sartorius is innervated by the femoral nerve (Standring
be joined with gluteus maximus (Bergman et al. 1988; du
2016).
Plessis and Loukas 2016). It may be partially or completely
divided into bundles (Macalister 1875; Bergman et al. Comparative Anatomy
1988; du Plessis and Loukas 2016). Accessory slips may be
Sartorius has a similar typical presentation in the apes, orig-
present with attachments to the inguinal ligament, abdomi-
inating from the lateral ilium near the anterior superior iliac
nal aponeurosis, or iliac crest (Macalister 1875; Bergman
spine and inserting onto the medial aspect of the tibial shaft
et al. 1988; du Plessis and Loukas 2016). Macalister (1875)
(Champneys 1872; Hepburn 1892; Beddard 1893; Sonntag
observed a tendinous band extending from the tensor fas-
1924; Raven 1950; Sigmon 1974; Gibbs 1999; Diogo et al.
ciae latae into the base of the greater trochanter.
2010, 2012, 2013a,b, 2017). The insertion may be more dis-
Prevalence tal onto the tibia in gibbons (Diogo et al. 2012).
N/A
Variations
Description Sartorius may be absent (Macalister 1875; Bergman et al.
On the right side of a fetus with craniorachischisis dissected 1988; du Plessis and Loukas 2016; Standring 2016). This mus-
by Alghamdi et al. (2017), tensor fasciae latae was fused cle can vary in its course as it descends in the anterior thigh
with gluteus medius. In the female fetus with trisomy 18 (Macalister 1875). Sartorius may be partially or fully split into
dissected by Alghamdi et al. (2018), gluteus medius was two longitudinal portions, and the additional portion can have
fused with tensor fasciae latae bilaterally. a separate insertion onto the tibia, the femur, the fascia lata,
Bersu et al. (1976) describe a male infant with Hanhart the patellar ligament, the semitendinosus tendon, or the ten-
syndrome. The femora of this infant were normally devel- don of the other sartorius portion (Hallett 1848; Macalister
oped but distal secondary ossifcation centers were absent. 1875; el-Badawi 1987; Bergman et al. 1988; du Plessis and
The left leg stump had a patella and a small rudiment of the Loukas 2016; Coban et al. 2019). This bifurcation may be
proximal tibia but no fbular rudiment. The right leg stump confned to its origin or insertion or comprise over half of the
muscle (Hallett 1848; Macalister 1875; Knott 1883b; Kedzia Prevalence

et al. 2011; du Plessis and Loukas 2016; Coban et al. 2019). N/A
Macalister (1875) notes that Kelch and Kyrtl observed a cen-
tral tendinous intersection in sartorius. An extra head of sar- Clinical Implications
torius may originate from the pubis, pectineal line, femoral Melling and Zweymüller (1996) note that an attachment of
sheath, below the anterior superior iliac spine, or from the sartorius to the medial meniscus stabilizes and protects this
inguinal ligament (Macalister 1875; Bergman et al. 1988; du structure in the knee joint and may prevent herniation or
Plessis and Loukas 2016; Standring 2016). injury during sudden extension of the knee.
The insertion of sartorius near the knee may vary
(Standring 2016), and insertions range from the fascia lata,
fascia of the leg, or knee joint capsule (Macalister 1875; rectus feMOris (figure 5.2)
Knott 1883a,b; Bergman et al. 1988; du Plessis and Loukas Synonyms
2016). Additional insertions may also go to the patella, This muscle may also be referred to as quadriceps extensor
medial femoral condyle, the knee joint capsule, or medial cruris (Knott 1883b), extensor cruris medialis superfcialis,
meniscus (Melling and Zweymüller 1996; Kedzia et al. or rectus anterior.
2011; du Plessis and Loukas 2016).
Mori (1964) categorized the distal portion of sartorius Typical Presentation
into fve types. In type 1, the boundary of the muscular part
Description
and the tendon of insertion is distinct, and the length of the
tendon is moderate. In type 2, the boundary of the muscular Rectus femoris is one component of quadriceps femo-
part and the tendon of insertion is not very clear, but the ris. It has a dual origin, arising from the anterior inferior
length of the tendon is moderate. In type 3, the tendon of iliac spine and from the area just above the acetabulum
insertion is short, and the muscle fbers insert directly onto (Standring 2016). Some fbers originate from the capsule of
the tibia. In type 4, the distal part of the muscle is divided the hip joint (Standring 2016). The muscle ends in a tendon
longitudinally into two portions, and the tendon of the sec- that inserts onto the base of the patella (Standring 2016).
ondary slip is attached to the anterior wall of the adductor
Innervation
canal. In type 5, the muscle fbers typically in the distal
portion of sartorius are replaced by a fbrous band, and this Rectus femoris is innervated by the femoral nerve
band is attached to the vastus medialis fascia. (Standring 2016).
Kim and Lee (2019) describe a case from a left lower Comparative Anatomy
limb in which sartorius split into medial and lateral heads
The typical presentation of rectus femoris in the apes is
and was associated with an accessory sartorius muscle
a muscle with a single head originating from the anterior
that originated from the inguinal ligament. The lateral
superior iliac spine that inserts into the patella (Champneys
head of sartorius inserted onto the medial aspect of the
1872; Hepburn 1892; Beddard 1893; Sonntag 1924; Raven
patella. The medial head merged with the accessory sarto-
1950; Sigmon 1974; Gibbs 1999; Diogo et al. 2010, 2012,
rius and the two inserted together onto the medial surface
2013a,b, 2017). The head from the ilium near the acetab-
of the tibia, forming the pes anserinus. The medial head
ulum is rarely present in orangutans (Gibbs 1999; Diogo
also sent a muscular slip to vastus medialis. Sartorius is
et al. 2013b) occasionally present in common chimpanzees
also associated with the musculus saphenous. It presents
and bonobos (Hepburn 1982; Beddard 1893; Miller 1952;
as a detached slip that attaches to the inguinal ligament
Gibbs 1999; Diogo et al. 2013a, 2017), and frequently pres-
and loops under the saphenofemoral junction, blending
ent in gorillas (Gibbs 1999; Diogo et al. 2010).
with the sartorius laterally and the adductor longus medi-
ally (Tyrie 1894; du Plessis and Loukas 2016; Saga and
Takahashi 2016). Variations
Description
Prevalence The two origins of rectus femoris may be continuous
Mori (1964) classifed the distal portions of sartorius into (Macalister 1875; Knott 1883b; du Plessis and Loukas
fve types (see above for descriptions). Out of 50 thighs, type 2016). A double origin may be present from the ante-
1 was found in 31 cases (62%), type 2 in 13 cases (26%), and rior inferior iliac spine (Macalister 1875; du Plessis and
type 3 in 6 cases (12%). Out of 320 thighs, type 4 was found Loukas 2016). A portion of the insertion tendon may
in 15 cases (4.7%) and type 5 was found in 2 cases (0.6%). split off from the rest and pass in front of the patella
(Macalister 1875). The tendons of the vasti may overlap
Anomalies and form a canal through which the rectus femoris tendon
Description passes (Macalister 1875; Knott 1883b). Vastus lateralis
Pirani et al. (1991) describe soft tissue anatomy associated may insert into the tendon of rectus femoris (Macalister
with cases of PFFD. In Aitken type B PFFD, the cross- 1875). Rectus femoris may be continuous with the vasti
sectional diameter of sartorius is larger than is typical. (Macalister 1875; Bergman et al. 1988). Stringer et al.
(2012) report two cases in which a shortened rectus fem- inferior fbers that extend between the tendon of adductor
oris narrowed into a tendon at the middle of the thigh magnus and the medial aspect of the patella as vastus medi-
before joining the quadriceps tendon. In one of these alis obliquus (Standring 2016).
cases, the tendon also blended with the fascia over vastus
medialis and vastus lateralis. Typical Presentation
The head of rectus femoris that originates from the ace- Description
tabulum/capsule of the hip joint may be absent (Macalister Vastus medialis is one component of quadriceps femoris.
1875; du Plessis and Loukas 2016; Standring 2016). The (Standring 2016). It originates from the femur with attach-
entire rectus femoris muscle or the whole quadriceps femo- ments to the intertrochanteric line, spiral line, linea aspera,
ris mass may be absent (Bergman et al. 1988; Stringer et al. and medial supracondylar line (Standring 2016). It attaches
2012). A third “femoral” head may be present with a deep to the medial aspect of the patella and to the quadriceps
attachment to the iliofemoral ligament, a superfcial attach- femoris tendon (Standring 2016).
ment to the tendon of gluteus minimus, and an insertion into
the greater trochanter (Tubbs et al. 2006d). Rectus accesso- Innervation
rius is an accessory slip that originates from a tendon at the Vastus medialis is innervated by the femoral nerve
margin of the acetabulum and inserts into the ventral border (Standring 2016).
of vastus lateralis (Bergman et al. 1988). An accessory slip
may also be present and extend between the inguinal liga- Comparative Anatomy
ment and vastus medialis (Takeshige et al. 1960). Vastus medialis has a similar typical presentation in the
apes, extending from the posteromedial femoral shaft near
Prevalence the linea aspera and the iliofemoral ligament to the patella,
In a study of 40 specimens, Knott (1883b) reported that with a contribution to the knee joint capsule (Champneys
three of them (7.5%) exhibited direct continuity of the spi- 1872; Hepburn 1892; Beddard 1893; Sonntag 1924; Raven
nous and acetabular origins. In one case (2.5%), a small, 1950; Sigmon 1974; Gibbs 1999; Diogo et al. 2010, 2012,
tendinous accessory slip originated from the anterior 2013a,b, 2017). The origin is more proximal in gibbons than
superior iliac spine. In two cases (5%), the origin from the in the other apes (Sigmon 1974) and can reach the femoral
anterior superior iliac spine was divided into two parts that neck in common chimpanzees (Diogo et al. 2013a).
were separated by a thin layer of connective tissue. In one
of these cases (2.5%), and in three other cases which do not Variations
appear to be included in the sample of 40 specimens, the Description
insertion tendon of rectus femoris passed through a canal The vasti may be bilaminar (Macalister 1875; Knott 1883b;
formed by the overlapping of the tendons of the vasti before Bergman et al. 1988; du Plessis and Loukas 2016). The
inserting onto the patella. In a study of 96 sides from 48 vasti may be continuous with rectus femoris (Mori 1964;
cadavers, Tubbs et al. (2006d) found that 80 sides (83%) had Bergman et al. 1988; du Plessis and Loukas 2016). Vastus
a third “femoral” head of rectus femoris. Two sides (2.1%) medialis is often fused with vastus intermedius (Macalister
had third heads that were bilaminar. 1875; Mori 1964). The vasti may merge into one feshy mus-
cle mass (Bergman et al. 1988). The entire quadriceps fem-
Anomalies
oris muscle may be absent (Bergman et al. 1988). Its lower
Description fbers may have a separate insertion into the medial condyle
In one anencephalic fetus, Windle (1893) found that the straight of the tibia (Macalister 1875). Vastus medialis obliquus (the
(spinous) head of rectus femoris was absent on the left side. inferior portion of the muscle) can vary in its extent of its
Pirani et al. (1991) describe soft tissue anatomy associated with insertion along the medial aspect of the patella (Holt et al.
cases of proximal femoral focal defciency (PFFD). In Aitken 2008).
type B PFFD, rectus femoris is smaller than is typical. Mori (1964) found three types of distal fusion between
vastus medialis and rectus femoris. There was either total
Prevalence fusion of the distal portions of both muscles, fusion between
N/A the distal part of vastus medialis and the tendon of rectus
femoris, or no fusion between the muscles. Mori (1964) also
Clinical Implications found three types of fusion between vastus medialis and
N/A vastus intermedius. There was either total fusion of both
muscles, partial fusion, or no fusion.
vastus MeDialis (figure 5.2)
Prevalence
Synonyms Mori (1964) reports that out of 50 thighs, there was total
This muscle may also be referred to as vastus internus fusion of the distal portions of both vastus medialis and rec-
(Macalister 1875). Some researchers may refer to the most tus femoris in 22% of cases, fusion between the distal part
of vastus medialis and the tendon of rectus femoris in 19%, and inserting into the patella (Hepburn 1892; Beddard
or no fusion between the muscles in 80%. Mori (1964) also 1893; Sonntag 1924; Raven 1950; Sigmon 1974; Gibbs
found that there was total fusion between vastus medialis 1999; Diogo et al. 2010, 2012, 2013a,b, 2017).
and vastus intermedius in 20% of cases, partial fusion in
50%, and no fusion between the muscles in 30%. Variations
Holt et al. (2008) studied the extent of the insertion of Description
vastus medialis obliquus in 65 patients with MRI and in 18
The vasti may be bilaminar, which is an accentuation
cadavers. The mean insertion was 51% of patellar length
of the normal presentation of the fbers in vastus latera-
(ranging from 13% to 95%) for the MRI cohort, and 52% of
lis (Macalister 1875; Knott 1883b; Bergman et al. 1988).
patellar length (ranging from 26% to 81%) in the cadaveric
The entire quadriceps femoris muscle may be absent
study.
(Bergman et al. 1988). The origin of vastus lateralis
Anomalies may be confned to the gluteal tuberosity (du Plessis and
Loukas 2016). Vastus lateralis may also insert into the
Description
tendon of rectus femoris (Macalister 1875; Mori 1964).
In both a male fetus and female fetus with triploidy studied Vastus lateralis may be doubled, presenting as a muscle
by Moen et al. (1984), the vastus heads of quadriceps femo- with two separate heads (Dwight 1887; du Plessis and
ris were fused bilaterally. Pirani et al. (1991) describe soft Loukas 2016).
tissue anatomy associated with cases of proximal femoral The vasti may be continuous with rectus femoris
focal defciency (PFFD). In Aitken type B PFFD, vastus (Macalister 1875; Mori 1964; Bergman et al. 1988). Vastus
medialis is smaller than is typical. lateralis may be connected to the other vasti, particularly
vastus intermedius (Macalister 1875; Mori 1964; Willan
Prevalence
et al. 1990; Becker et al. 2009; du Plessis and Loukas
N/A 2016). Accessory bellies of vastus lateralis may be pres-
ent when this muscle is fused with vastus intermedius (du
Plessis and Loukas 2016). Vastus lateralis may be con-
Variation of the insertion of vastus medialis obliquus into nected with articularis genus when this latter muscle
the patella should be acknowledged when choosing a surgi- consists of two slips (Macalister 1875). The vasti may
cal approach for total knee arthroplasty (Holt et al. 2008). merge into one feshy muscle mass (Bergman et al. 1988).
Heimkes et al. (1992) studied the relationships upon inser-
vastus lateralis (figure 5.2) tion between gluteus medius, gluteus minimus, and vastus
lateralis and classifed these relationships into fve types.
Synonyms See the gluteus medius entry for more details and preva-
This muscle may also be referred to as vastus externus lence information. Rectus accessorius is an accessory slip
(Macalister 1875). Some may refer to the distal portion of that originates from a tendon at the margin of the acetabu-
the muscle that inserts into the patella as vastus lateralis lum and inserts into the ventral border of vastus lateralis
obliquus (du Plessis and Loukas 2016). (Bergman et al. 1988).
Mori (1964) found three types of fusion between vas-
Typical Presentation tus lateralis and vastus intermedius including total fusion,
Description partial fusion, and no fusion. Mori (1964) also categorized
Vastus lateralis is one component of quadriceps femoris four relationships between the insertions of vastus lateralis
(Standring 2016). It originates from the femur with attach- and rectus femoris. These relationships include the distal
ments to the intertrochanteric line, greater trochanter, glu- portion of vastus lateralis situated lateral to the distal por-
teal tuberosity, and linea aspera (Standring 2016). Some tion of rectus femoris, the distal portion of vastus lateralis
fbers originate from the tendon of gluteus maximus and covered by the distal portion of rectus femoris, the distal
the lateral intermuscular septum (Standring 2016). It inserts portion of vastus lateralis fused with the fascia of rectus
onto the lateral aspect of the patella and joins the quadri- femoris, and the distal portion of vastus lateralis fused with
ceps tendon (Standring 2016). the tendon of rectus femoris.
Vastus lateralis is innervated by the femoral nerve Mori (1964) found that in a sample of 50 thighs, vastus late-
(Standring 2016). ralis and vastus intermedius were totally fused in 18% of
cases, partially fused in 55%, and not fused in 26%. Mori
Comparative Anatomy (1964) also found that the distal portion of vastus lateralis
Vastus lateralis has a similar typical presentation in the was situated lateral to the distal portion of rectus femoris
apes, originating from the lateral femur, with attachments in 52% of cases, the distal portion of vastus lateralis was
to the greater trochanter and lateral lip of the linea aspera, covered by the distal portion of rectus femoris in 28%, the
distal portion of vastus lateralis fused with the fascia of rec- surfaces of the proximal femoral shaft (Standring 2016). It
tus femoris in 12%, and the distal portion of vastus lateralis inserts onto the lateral edge of patella and lateral condyle of
fused with the tendon of rectus femoris in 8%. the tibia (Standring 2016).
Willan et al. (1990) examined vastus lateralis and vas-
tus intermedius in 75 lower limbs from 40 cadavers. More Innervation
than three-fourths of the deep surface of vastus lateralis Vastus intermedius is innervated by the femoral nerve
was fused with vastus intermedius in 25 limbs (33.3%), (Standring 2016).
about half of vastus lateralis was fused with vastus inter-
medius in 32 limbs (42.7%), and less than one-half of vas- Comparative Anatomy
tus lateralis was fused with vastus intermedius in 18 limbs Vastus intermedius has a similar typical presentation in
(24%). The anterior edge of vastus lateralis was feshy in the apes, extending from the ventral surface of the femoral
22 limbs (29.3%), while there was an anterior tendinous shaft to the patella (Hepburn 1892; Beddard 1893; Sonntag
lamina wider than 5 mm in 28 limbs (27.3%) and narrower 1924; Raven 1950; Miller 1952; Sigmon 1974; Gibbs 1999;
than 5 mm in 24 limbs (32%). There was a separate deep Diogo et al. 2010, 2012, 2013a,b, 2017). The origin may
tendinous lamina from either vastus lateralis or vastus extend to the femoral neck in gorillas (Raven 1950).
intermedius in 22 limbs (29.3%). There was a separate
deep feshy lamina between vastus lateralis and vastus Variations
intermedius in 27 limbs (36%), with fusion to the tendi- Description
nous lamina in 13 limbs (17.3%), fusion to vastus latera- The vasti may be bilaminar (Macalister 1875; Knott 1883b;
lis in 6 limbs (8%), and fusion to vastus intermedius in 8 Bergman et al. 1988). The entire quadriceps femoris muscle
limbs (10.7%). may be absent (Bergman et al. 1988). The origin of vastus
intermedius may be as high as the intertrochanteric line
Anomalies
(Macalister 1875). The vasti may be continuous with rectus
Description femoris (Bergman et al. 1988). Vastus intermedius may be
Pettersen et al. (1979) found bilateral accessory vastus late- fully or partially fused with vastus lateralis (Macalister 1875;
ralis muscles in a fetus with trisomy 13. On the left side, the Mori 1964; Willan et al. 1990; Becker et al. 2009; du Plessis
muscle originated from the deep surface of the upper part of and Loukas 2016). Vastus intermedius may also be fused
the main vastus lateralis muscle and inserted into the patella proximally with vastus medialis (Macalister 1875; Standring
via feshy fbers deep to the attachment of rectus femoris. 2016; du Plessis and Loukas 2016). Vastus intermedius may
On the right side, the muscle was similar but had originated join with articularis genus, and if the latter muscle is absent,
separately from vastus lateralis. In both a male fetus and the deep fbers of vastus intermedius can attach in rare cases
female fetus with triploidy studied by Moen et al. (1984), the to the synovial membrane of the knee (Macalister 1875; du
vastus heads of quadriceps femoris were fused bilaterally. Plessis and Loukas 2016). The vasti may merge into one
feshy muscle mass (Bergman et al. 1988).
Prevalence
In their literature review, Smith et al. (2015) found that Prevalence
accessory vastus lateralis muscles have only been recorded Mori (1964) found that in a sample of 50 thighs, vastus
in 1 out of 20 individuals with trisomy 13 (5%). lateralis and vastus intermedius were totally fused in 18%
of cases, partially fused in 55%, and not fused in 26%.
Mori (1964) also found that there was total fusion between
Acknowledging the variable relationships between glu- vastus medialis and vastus intermedius in 20% of cases,
teus medius, gluteus minimus, and vastus lateralis and the partial fusion in 50%, and no fusion between the muscles
potential independent insertions of these muscles onto the in 30%.
greater trochanter is important for successful hip opera- Willan et al. (1990) examined vastus lateralis and vastus
tions, particularly when using the transgluteal approach intermedius in 75 lower limbs from 40 cadavers. More than
(Heimkes et al. 1992). three-fourths of the deep surface of vastus lateralis was
fused with vastus intermedius in 25 limbs (33.3%), about
vastus interMeDius (figure 5.2) half of vastus lateralis was fused with vastus intermedius in
32 limbs (42.7%), and less than one-half of vastus lateralis
Synonyms was fused with vastus intermedius in 18 limbs (24%). There
This muscle may also be referred to as cruraeus (Macalister was a separate deep tendinous lamina from either vastus
1875). lateralis or vastus intermedius in 22 limbs (29.3%). There
was a separate deep feshy lamina between vastus lateralis
Typical Presentation and vastus intermedius in 27 limbs (36%), with fusion to
Description the tendinous lamina in 13 limbs (17.3%), fusion to vastus
Vastus intermedius is one component of quadriceps femoris lateralis in six limbs (8%), and fusion to vastus intermedius
(Standring 2016). It originates from the anterior and lateral in eight limbs (10.7%).
Anomalies There is variation in the number of bundles that comprise

Description articularis genus, with some reports finding only one bundle
and others finding more than ten bundles (DiDio et al. 1967,
In both a male fetus and female fetus with triploidy studied
1969; Ahmad 1975; Kimura and Takahashi 1987; Puig et al.
by Moen et al. (1984), the vastus heads of quadriceps femo-
1996; Toscano et al. 2004; Woodley 2016; Woodley et al.
ris were fused bilaterally.
2012; Sakuma et al. 2014; Grob et al. 2017). The bundles
Prevalence may be organized into multiple layers, often recognized as
N/A split into superficial and deep layers, and sometimes with a
recognized intermediate layer (Kimura and Takahashi 1987;
Clinical Implications Woodley et al. 2012; Sakuma et al. 2014; Grob et al. 2017).
The complex and variable anatomical relationships between Varying definitions of which fibers constitute articularis
articularis genus, vastus intermedius, and subsequently vas- genus (Grob et al. 2017) or underestimation of bundles due
tus medialis, are important to acknowledge when planning to using MRI (Puig et al. 1996; Woodley 2016; Woodley
knee surgery (Grob et al. 2017). et al. 2012) may explain why the number of bundles reported
in the literature varies greatly among studies.
Articularis genus (Figure 5.2) Prevalence

Knott (1883b) found that articularis genus was absent in 3
Synonyms
out of 40 limbs (7.5%). DiDio et al. (1967) studied articu-
This muscle may also be referred to as sub-cruraeus laris genus in 156 limbs from 78 cadavers. One bundle of
(Macalister 1875), subcrurales, tenseur de la synoviale, articularis genus was found in 70 cases (44.9%), two bun-
supragenualis, or suprageniculares (DiDio et al. 1997). dles in 52 cases (33.3%), three bundles in 23 cases (14.7%),
four bundles in 9 cases (5.8%), and six bundles in 2 cases
(1.3%). The articularis genus muscles were rectangular in
Description 87 cases (55.8%), trapezoidal in 42 cases (26.9%), and had
Articularis genus is comprised of several bundles that arise an inverted trapezoidal outline in 27 cases (17.3%). The ori-
deep to vastus intermedius (Standring 2016). The bundles gin of articularis genus was the lateral and medial aspects
originate from the distal femoral shaft and insert onto the of the distal femur in 83 cases (53.5%), the anterior aspect
proximal reflection of the synovial membrane (suprapatel- in 26 cases (16.7%), anterior and lateral aspects in six cases
lar bursa) of the knee joint (Standring 2016). (3.8%), medial aspect in four cases (2.6%), anterior and
medial aspects in four cases (2.6%), and lateral aspect in
Innervation two cases (1.3%). Its insertion of its bundles into the supra-
Articularis genus is innervated by the femoral nerve patellar bursa varied from the center of its proximal bor-
(Standring 2016). der in 126 out of 401 times (31.4%), medial aspect of its
proximal border 117 times (29.2%), lateral aspect 110 times
Comparative Anatomy (27.4%), posterior aspect 25 times (6.2%), and the anterior
Hepburn (1892) claims that articularis genus is present in aspect 23 times (5.7%).
all apes, but this muscle has not been reported or has been DiDio et al. (1969) studied articularis genus in 66 indi-
found absent in all apes except gorillas (Sonntag 1923; viduals, ranging from fetuses to adults. The shape of the
Diogo et al. 2012, 2013a,b, 2017). In gorillas, it extends from muscle was triangular in 44% of cases, had a lambdoidal
the distal part of the femur to the proximolateral aspect of outline in 27%, was rectangular in 24%, and had an unclas-
the articular capsule of the knee joint (Raven 1950; Gibbs sified shape in 5%. One bundle was found in 66.7% of
1999; Diogo et al. 2010). cases, two partially divided bundles in 27.3% of cases, two
bundles in 1.5% of cases, and in 4.5% of cases the num-
Variations ber of bundles was not determined. Kimura and Takahashi
Description (1987) studied the number of bundles of articularis genus in
Articularis genus may blend with vastus intermedius 40 thighs from 44 cadavers. One bundle was found in two
(Macalister 1875; du Plessis and Loukas 2016; Standring cases (5%), two bundles in 11 cases (27.5%), three bundles
2016; Grob et al. 2017). Articularis genus may be reduced, in eight cases (20%), four bundles in seven cases (17.5%),
absent, or divided (Macalister 1875; du Plessis and Loukas five bundles in five cases (12.5%), six bundles in three cases
2016). It is also variable in shape and size (e.g., DiDio (7.5%), seven bundles in one case (2.5%), and in three cases
et al. 1967, 1969; Toscano et al. 2004). Its insertion onto the (7.5%) the bundles were unable to be properly counted.
suprapatellar bursa and the knee joint capsule can vary in Toscano et al. (2004) studied articularis genus in 15 lower
its specific location (DiDio et al. 1967; Grob et al. 2017). In limbs. The muscles were trapezoidal in shape in 40% of cases,
addition to its typical insertion into the upper portion of the rectangular in 33%, and in the remaining cases, the muscles
synovial membrane, it can insert onto either side of the knee were either triangular or irregular. In 57.1% of cases, articu-
(Macalister 1875; du Plessis and Loukas 2016). laris genus inserted onto the anterior aspect of the proximal
edge of the suprapatellar bursa. The most common number ramus, the ischial ramus, the pelvic surface of the ilium,
of bundles was four bundles (33.3% of cases). Woodley et al. and the greater sciatic foramen (Standring 2016). It also
(2012) used MRI and dissection to study articularis genus originates from the obturator membrane and the obturator
in 18 lower limbs. Using MRI, these authors found that two fascia (Standring 2016). Its fibers converge into four or five
bundles were present in one case (5.6%), three bundles in tendinous bands that pass through the lesser sciatic fora-
five cases (27.8%), four bundles in nine cases (50%), and five men to insert as a single tendon onto the greater trochanter
bundles in three cases (16.7%). Using dissection on these (Standring 2016). The superior gemellus and inferior gemel-
same specimens, they found that there were four bundles in lus fuse with this tendon prior to insertion (Standring 2016).
one case (5.6%), five bundles in four cases (22.2%), six bun-
dles in one case (5.6%), seven bundles in six cases (33.3%), Innervation
eight bundles in two cases (11.1%), nine bundles in two cases Obturator internus is innervated by the nerve to obturator
(11.1%), and ten bundles in two cases (11.1%). internus (Standring 2016).
Grob et al. (2017) studied articularis genus in 18 lower
limbs from 12 cadavers. They found that bundles of the Comparative Anatomy
superficial layer, and in 11 specimens (61.1%) the bundles of Obturator internus has a similar typical presentation in the
the intermediate layer, arose from vastus intermedius and apes, arising from the bones around the internal aspect of
the anterior and anterolateral surfaces of the femurs. The the obturator foramen and from the obturator membrane,
bundles of the deep layer, and in the remaining seven speci- passing through the lesser sciatic foramen, and inserting
mens (38.9%), the bundles of the intermediate layer arose with the gemelli onto the trochanteric fossa of the femur
from the anterior surface of the femur only. Out of the 18 (Champneys 1872; Hepburn 1892; Beddard 1893; Raven
lower limbs, the articularis genus muscle from one limb had 1950; Sigmon 1974; Diogo et al. 2010, 2012, 2013a,b, 2017).
three bundles (5.6%), four limbs had four bundles (22.2%),
six limbs had five bundles (33.3%), and seven limbs had Variations
articularis genus muscles with six bundles (38.9%). Description
Obturator internus may be divided into two parts (Bergman
Anomalies
et al. 1988). Accessory slips may be present (Macalister
N/A 1875; Bergman et al. 1988; Nicholson et al. 2016). These slips
may originate from the lower margin of the ilio-pectineal
line, from the tendon of psoas minor, the ridge above the
The complex and variable anatomical relationships between inner margin of the ischial tuberosity, the inner aspect of
articularis genus, vastus intermedius, and subsequently vas- the ischium, the third sacral vertebra, the pelvic fascia, or
tus medialis, are important to acknowledge when planning the sacrotuberous ligament (Macalister 1875; Knott 1883b;
knee surgery (Grob et al. 2017). Nicholson et al. 2016). Some fibers from obturator internus
may be continuous with those of gemellus superior or gemel-
Obturator internus (Figures 4.4 a nd 5.3) lus inferior (Aung et al. 2001; Bergman et al. 1988; Nicholson
et al. 2016). The tendon of obturator internus may blend
Synonyms with the tendon of piriformis (Wood 1868; Pine et al. 2011).
N/A Obturator internus may be absent (Sato 1970).
Description From the examination of 36 bodies, Wood (1868) states that
Obturator internus originates from bony surfaces that sur- in three males and one female (11%), the tendon of pirifor-
round the obturator foramen including the inferior pubic mis was blended with that of obturator internus—the two
FIGURE 5.3 Lateral rotators of the hip in posterior view.

inserted together onto the greater trochanter. Though total Innervation

sample size is unclear, Knott (1883b) noted an origin from Obturator externus is innervated by the posterior branch of
the sacrotuberous ligament in three cases and a slip from the obturator nerve (Standring 2016).
the inner surface of the ischium above the spine in two sub-
jects. Sato (1970) found that obturator internus was absent Comparative Anatomy
in 30 out of 410 limbs (7.3%) from Kyushu Japanese males Obturator externus has a similar typical presentation in the
and in 29 out of 250 limbs (11.6%) from females. Aung et al. apes, arising from the bones around the external aspect of
(2001) found that out of 101 hemipelves from 60 cadavers, the obturator foramen and from the obturator membrane
two specimens (2%) exhibited a distal extension of obtura- and inserting onto the trochanteric fossa of the femur
tor internus muscle fibers that continued to join the fibers (Champneys 1872; Hepburn 1892; Beddard 1893; Raven
of inferior gemellus. Pine et al. (2011) found that out of 29 1950; Sigmon 1974; Diogo et al. 2010, 2012, 2013a,b, 2017).
specimens, the tendons of obturator internus and piriformis It may fuse with obturator internus in gibbons, orangutans,
were at least partially fused in four specimens (13.8%), with and common chimpanzees (Hepburn 1892; Beddard 1893)
three inserting as a common tendon and one dividing to and both gemelli in orangutans (Diogo et al. 2013b). It can
have separate attachments. have an attachment onto the femoral neck in gorillas and
common chimpanzees (Hepburn 1892; Gibbs 1999) and
Anomalies from the hip joint capsule in bonobos (Diogo et al. 2017).
Description
Bersu et al. (1976) describe a male infant with Hanhart syn- Variations
drome. The femora of this infant were normally developed Description
but distal secondary ossification centers were absent. The left Obturator externus may have a connection with adductor
leg stump had a patella and a small rudiment of the proxi- brevis (Bergman et al. 1988; du Plessis and Loukas 2016).
mal tibia but no fibular rudiment. The right leg stump was Macalister (1875) notes more specifically that adductor
less developed and had a patella, smaller tibial rudiment, and brevis may send a slip to the tendon of obturator externus.
no fibular rudiment. On both sides, obturator internus had Macalister (1875) and Knott (1883b) note that the obturator
normal origins. On the left side, it inserted onto the inferior externus tendon may have an attachment or some degree of
aspect of the sacrotuberous ligament. On the right side, it connection onto the capsule of the hip joint. The muscle may
inserted into the periosteum of the lesser sciatic notch. be split into superior and inferior portions by the obturator
Pirani et al. (1991) describe soft tissue anatomy associated nerve and vessels (Macalister 1875; Knott 1883b; Bergman
with cases of proximal femoral focal deficiency (PFFD). In et al. 1988; du Plessis and Loukas 2016). The superior por-
Aitken type A PFFD, the short external rotators of the hip tion may arise from the horizontal ramus of the pubis below
are larger in cross-sectional diameter than normal and insert pectineus and is separated by the rest of the muscle by the
onto the posteromedial surface of the greater trochanter. In obturator nerve and vessels (Macalister 1875; Knott 1883b).
Aitken type B PFFD, the short external rotators are smaller Obturator externus may be split into three parts (Bergman
than normal. In a female fetus with trisomy 18, obturator et al. 1988; du Plessis and Loukas 2016).
internus, superior gemellus, and inferior gemellus were fused Some authors report accessory muscle slips and/or fas-
laterally on both sides of the body (Alghamdi et al. 2018). ciculi closely associated with obturator externus. Nakamura
et al. (1992) and Yatsunami et al. (2004) report accessory
Prevalence
muscles occurring between adductor brevis and adductor
N/A minimus, originating from the upper part of the inferior
pubic ramus and running closely with obturator externus
before inserting onto the pectineal line, the lesser trochan-
N/A ter, or the anterior aspect of the aponeurotic insertion of
adductor minimus. Nakamura et al. (1992) suggest that
Obturator externus (Figure 5.3) these muscles are formed when the superficial fascicle of
obturator externus separates from the rest of the muscle
Synonyms during development and forms the supernumerary muscle
N/A when it has an insertion other than the trochanteric fossa.
In addition to similar accessory muscles, Yatsunami et al.
Typical Presentation (2004) report accessory fasciculi fused to the posterior sur-
Description face of adductor minimus, which they suggest are likely
Obturator externus originates from the structures that sur- homologous to the superficial layer of the main belly of
round the external/anterior surface of the obturator foramen obturator externus.
including the pubic ramus, the ischial ramus, and the obtu- Upasna et al. (2013) report an accessory muscle in the
rator membrane (Standring 2016). It inserts via a tendon left leg of a cadaver that also exhibited an adductor lon-
onto the trochanteric fossa of the femur (Standring 2016). gus and pectineus with a combined origin from the pecten
pubis, and an adductor brevis that was divided in two and dissociated femoral head and proximal femoral metaphy-
had distal fibers that merged with those of adductor longus. sis before inserting onto the posteromedial surface of the
The accessory muscle was located under adductor brevis proximal end of the shaft of the femur. It is muscular up to
and lied on top of the obturator externus and the posterior its insertion. In Aitken type D PFFD, obturator externus
division of the obturator nerve. Its fibers extended between inserts onto the proximal femoral ossicle.
the inferior pubic ramus and merged with the aponeurosis
of adductor magnus. A similar muscular slip was also pres- Prevalence
ent on the right side but was not described in detail. N/A
Knott (1883b) observed that obturator externus was split N/A
into two portions by the obturator nerve and vessels in 4 out
of 20 cases (20%). Nakamura et al. (1992) found a super-
numerary muscle in 33 out of 100 thighs from 50 cadavers Gemellus superior (Figure 5.3)
(33%). The muscles had an insertion onto the anterior sur- Synonyms
face of the insertion aponeurosis of adductor minimus in 17 N/A
out of 33 thighs (51.5%), the upper portion of the pectineal
line in nine thighs (27.3%), and the posterior aspect of the Typical Presentation
base of the lesser trochanter in seven thighs (21.2%).
Description
Yatsunami et al. (2004) found that out of 73 thighs from
45 cadavers, the obturator nerve pierced obturator externus Gemellus superior originates from the ischial spine, blends
in 50 cases (68.5%) and ran over the muscle in 23 cases with the obturator internus tendon, and inserts onto the
(31.5%). These authors found accessory muscles occurring greater trochanter (Standring 2016).
between adductor brevis and adductor minimus in 37 thighs
Innervation
(50.7%) and accessory fasciculi fused to the posterior sur-
face of adductor minimus in 18 thighs (24.7%). They suggest Gemellus superior is innervated by the nerve to obturator
that during ontogeny, the superior fasciculus of obtura- internus (Standring 2016).
tor externus retained its normal morphology in 23 thighs
Comparative Anatomy
(31.5%), a portion of the superior fasciculus converted into
the accessory muscle in 27 thighs (37%), the entire superior Gemellus superior has a similar typical presentation in
fasciculus converted into the accessory muscle in 10 thighs most apes, arising from the ischium near the ischial spine
(13.7%), and the entire superior fasciculus disappeared in and having a common insertion onto the trochanteric fossa
13 thighs (17.8%). with the tendon of obturator internus (Diogo et al. 2012,
2013a,b, 2017). Gemellus superior is absent in most gorillas
Anomalies (Gibbs 1999; Diogo et al. 2010).
Description
Bersu et al. (1976) describe a male infant with Hanhart Variations
syndrome. The femora of this infant were normally devel- Description
oped but distal secondary ossification centers were absent. Gemellus superior may be reduced or absent (Hallett 1848;
The left leg stump had a patella and a small rudiment of Wood 1868; Macalister 1875; Knott 1883a,b; Bergman
the proximal tibia but no fibular rudiment. The right leg et al. 1988; Duda et al. 1996; Nicholson et al. 2016; Standring
stump was less developed and had a patella, smaller tibial 2016). Absence of both gemelli is rare (Bergman et al. 1988).
rudiment, and no fibular rudiment. On both sides, obturator Gemellus superior can be separated into two parts (Knott
externus had normal origins. On the left side, it inserted 1883b). It may be fused with piriformis or gluteus mini-
into a tendon it shared with a slip from piriformis. On the mus (Macalister 1875; Bergman et al. 1988; Chiba 1992;
right side, it had a broad aponeurotic insertion onto the Nicholson et al. 2016). Some fibers from obturator internus
lesser trochanter. may be continuous with those of gemellus superior (Aung
Pirani et al. (1991) describe soft tissue anatomy associ- et al. 2001; Bergman et al. 1988; Nicholson et al. 2016).
ated with cases of proximal femoral focal deficiency (PFFD). Macalister (1875) observed it as sometimes inseparable
In Aitken type A PFFD, obturator externus is straight. It from obturator internus and reports it joining with the lower
courses through the concavity of the subtrochanteric varus border of gluteus minimus. Gemellus superior may have an
deformity of the proximal femoral metaphysis and inserts origin from the greater sciatic notch (Aasar 1947; Nicholson
onto the posteromedial surface of the greater trochanter and et al. 2016). This muscle may have an insertion onto the hip
is muscular nearly up to its insertion. In types B, C, and D, joint capsule (Macalister 1875; Bergman et al. 1988).
obturator externus is L-shaped. For example, in Aitken type Instead of forming a common tendon with obturator
B PFFD, the distal part of obturator externus is oriented internus and inferior gemellus, the tendon of gemellus supe-
90° cranially and 45° posteriorly and passes between the rior may insert onto the piriformis tendon (Macalister 1875;
Knott 1883b). Furthermore, the gemelli may instead cover surface of the greater trochanter. In Aitken type B PFFD, the
the superficial surface of the tendon of obturator internus or short external rotators are smaller than normal.
extend over this tendon as an aponeurosis (Shinohara 1995;
Nicholson et al. 2016). Babinski et al. (2003) describe a Prevalence
case in which a high division of the sciatic nerve caused the In their literature review, Smith et al. (2015) found that
common fibular nerve to pass superior to superior gemellus superior gemellus was absent in 1 out of 24 individuals
and the tibial nerve to pass inferior to superior gemellus. (4.2%) with trisomy 13 and 3 out of 26 (11.5%) individuals
Gemellus superior may also be doubled (Macalister with trisomy 18.
1875; Bergman et al. 1988; Arifoglu et al. 1997; Tanyeli
et al. 2006; Nicholson et al. 2016). Ortega et al. (2013) Clinical Implications
describe a case in which gemellus superior was bilaterally If components of the sciatic nerve surround superior gemel-
duplicated. The main belly of gemellus superior arose from lus, this morphology could implicate this muscle in pirifor-
the ischial spine and inserted onto the greater trochan- mis syndrome, coccygodynia, or muscle atrophy (Babinski
ter with piriformis via a common tendon. The duplicated et al. 2003). Doubling of superior gemellus may compress the
belly of gemellus superior—along with obturator internus sciatic nerve and contribute to piriformis syndrome (Arifoglu
and inferior gemellus—extended from the ischial tuberos- et al. 1997). Ortega et al. (2013) note that displacement of a
ity to the quadrate tubercle but was displaced and situated duplicated gemellus superior and nearby muscles can lead to
directly on the superior surface of quadratus femoris. weakness of lateral hip rotation and abduction of the femur.
Prevalence
Terry (1942) combines the observations from Wood (1867b Gemellus inferior (Figure 5.3)
and 1868) and finds that out of 70 total subjects, superior Synonyms
gemellus was absent in four bodies (5.7%). It was absent N/A
bilaterally in two males and one female, and absent on the
right side in one other male and was thus absent in 7 out of Typical Presentation
140 sides (5%). Terry (1942) found that superior gemellus Description
was absent in 31 out of 254 total individuals (12.2%) and
Gemellus inferior originates from the ischial tuberosity,
was absent more often in African Americans (22 out of 150
blends with the obturator internus tendon, and inserts onto
cases, or 14.7%) than in Whites (9 out of 104, or 8.7%). Sato
the greater trochanter (Standring 2016).
(1970) found that gemellus superior was absent in 21 out
of 390 limbs (5.4%) from Kyushu Japanese males and in Innervation
19 out of 260 limbs (7.3%) from females. Shinohara (1995)
Gemellus inferior is innervated by the nerve to quadratus
studied obturator internus and the gemelli on 14 sides from
femoris (Standring 2016).
12 cadavers. This author found that on five sides (36%) the
muscle fibers of the superior and inferior gemelli termi-
Comparative Anatomy
nated on the obturator internus tendon (the typical presen-
tation). On the nine other sides (64%), the gemelli covered Gemellus inferior has a similar typical presentation in
the superficial surface of the obturator internus tendon. the apes, arising from the ischial tuberosity and inserting
with obturator internus into the trochanteric fossa, with
Anomalies occasional fusion with nearby muscles (Champneys 1872;
Description Hepburn 1892; Beddard 1893; Raven 1950; Sigmon 1974;
Diogo et al. 2010, 2012, 2013a,b, 2017).
Gemellus superior may be absent in individuals with trisomy
13 or trisomy 18 (Smith et al. 2015). In a female fetus with
Variations
trisomy 18, obturator internus, gemellus superior, and gemel-
lus inferior were fused laterally on both sides of the body Description
(Alghamdi et al. 2018). Bersu et al. (1976) describe a male Gemellus inferior may be divided into two or three fasciculi
infant with Hanhart syndrome. The femora of this infant (Knott 1883b). It may also be doubled, and in rare cases, it
were normally developed, but distal secondary ossification has been reported absent (Hallett 1848; Knott 1883b; Sato
centers were absent. The left leg stump had a patella and a 1970; Bergman et al. 1988; Tanyeli et al. 2006; Nicholson
small rudiment of the proximal tibia but no fibular rudiment. et al. 2016). It is absent less often than is the superior
The right leg stump was less developed and had a patella, gemellus (Macalister 1875; Bergman et al. 1988; Duda
smaller tibial rudiment, and no fibular rudiment. Gemellus et al. 1996). Absence of both gemelli is rare (Bergman et al.
superior was absent bilaterally. Pirani et al. (1991) describe 1988). Wood (1868) found inferior gemellus absent on the
soft tissue anatomy associated with cases of proximal fem- right side and superior gemellus absent bilaterally in one
oral focal deficiency (PFFD). In Aitken type A PFFD, the female examined by him. Brenner et al. (2019) describe the
short external rotators of the hip are larger in cross-sectional right lower limb of a female cadaver in which piriformis
diameter than normal and insert onto the posteromedial and inferior gemellus were missing.
Gemellus inferior may be fused with quadratus femo- Quadratus femoris (Figure 5.3)
ris (Macalister 1875; Knott 1883b; Bergman et al. 1988;
Nicholson et al. 2016). Some fibers from obturator internus Synonyms
may be continuous with those of gemellus inferior (Aung N/A
et al. 2001; Bergman et al. 1988; Nicholson et al. 2016).
Macalister (1875) also observed it as sometimes insepa- Typical Presentation
rable from obturator internus. Macalister (1875) reports a Description
case described by Meckel, in which quadratus femoris was Quadratus femoris originates from the ischial tuberosity
absent and gemellus inferior was enlarged. Instead of form- and inserts onto the quadrate tubercle and onto the femur
ing a common tendon with obturator internus, the gemelli just below this tubercle (Standring 2016).
may instead cover the surface of the tendon of obtura-
tor internus or extend over this tendon as an aponeurosis Innervation
(Shinohara 1995; Nicholson et al. 2016). Quadratus femoris is innervated by the nerve to quadratus
femoris (Standring 2016).
Prevalence
Wood (1868) found inferior gemellus absent in 1 out of Comparative Anatomy
36 individuals (2.8%). Sato (1970) found that gemellus Quadratus femoris has a similar typical presentation in the
inferior was absent in 14 out of 390 limbs (3.6%) from apes, extending from the ischial tuberosity to the intertrochan-
Kyushu Japanese males and in 12 out of 264 limbs (4.5%) teric crest, with occasional extension onto the lesser or greater
from females. Shinohara (1995) studied obturator inter- trochanters (Champneys 1872; Hepburn 1892; Beddard 1893;
nus and the gemelli on 14 sides from 12 cadavers. This Sonntag 1924; Raven 1950; Miller 1952; Sigmon 1974; Diogo
author found that on five sides (36%) the muscle fibers of et al. 2010, 2012, 2013a,b, 2017). There is an attachment to the
the superior and inferior gemelli terminated on the obtura- hip joint capsule in bonobos (Diogo et al. 2017).
tor internus tendon (the typical presentation). On the nine
other sides (64%), the gemelli covered the superficial sur- Variations
face of the obturator internus tendon. Aung et al. (2001) Description
found that out of 101 hemipelves from 60 cadavers, two Quadratus femoris may be absent (Macalister 1875; Knott
specimens (2%) exhibited a distal extension of obturator 1880, 1883a,b; Sato 1970; Bergman et al. 1988; Nicholson
internus muscle fibers that continued to join the fibers of et al. 2016; Standring 2016). Tubbs and Salter (2006b)
inferior gemellus. describe a case in which an accessory piriformis muscle
was present on the same side of a limb where quadratus
Anomalies femoris was absent. Liu et al. (2011) found that in one case,
Description bilateral absence of quadratus femoris was also associated
Bersu et al. (1976) describe a male infant with Hanhart with bilateral absence of semimembranosus. Macalister
syndrome. The femora of this infant were normally devel- (1875) reports a case described by Meckel, in which quadra-
oped, but distal secondary ossification centers were absent. tus femoris was absent and gemellus inferior was enlarged.
The left leg stump had a patella and a small rudiment of Bergman et al. (1988) note that it can also be replaced by a
the proximal tibia but no fibular rudiment. The right leg large obturator internus.
stump was less developed and had a patella, smaller tibial The extent of its insertion onto the femur can vary (Duda
rudiment, and no fibular rudiment. Gemellus inferior was et al. 1996; Nicholson et al. 2016). It may also be fused with
absent bilaterally. Pirani et al. (1991) describe soft tissue inferior gemellus or adductor magnus (Macalister 1875;
anatomy associated with cases of proximal femoral focal Knott 1883b; Bergman et al. 1988; Nicholson et al. 2016).
deficiency (PFFD. In Aitken type A PFFD, the short exter- Quadratus femoris can send a slip to the origin of semi-
nal rotators of the hip are larger in cross-sectional diameter membranosus or have an attachment to the joint capsule of
than normal and insert onto the posteromedial surface of the hip (Macalister 1875; Knott 1883b). Quadratus femo-
the greater trochanter. In Aitken type B PFFD, the short ris may be split at its insertion, having a posterior part that
external rotators are smaller than normal. In a female fetus inserts normally and an anterior part that inserts onto the
with trisomy 18, obturator internus, gemellus superior, and intertrochanteric crest (Bergman et al. 1988). Quadratus
gemellus inferior were fused laterally on both sides of the femoris may be doubled (Tanyeli et al. 2006). Macalister
body (Alghamdi et al. 2018). (1875) reports a case observed by Jancke in which quadra-
tus femoris was split into 30 fasciculi.
Prevalence
N/A Prevalence
Sato (1970) found that quadratus femoris was absent in 4
Clinical Implications out of 418 limbs (1%) from Kyushu Japanese males and in
N/A 13 out of 262 limbs (5%) from females.
Anomalies Clinical Implications

Description Liu et al. (2011) suggest that absence of quadratus femo-
Bersu et al. (1976) describe a male infant with Hanhart ris may present as muscle weakness of hip lateral rotation.
syndrome. The femora of this infant were normally devel- Girolami et al. (2019) describe a case of isolated sciatica in
oped, but distal secondary ossification centers were absent. a left leg that was associated with a quadratus femoris mus-
The left leg stump had a patella and a small rudiment of cle with an enlarged belly and narrow origin and insertion.
the proximal tibia but no fibular rudiment. The right leg
stump was less developed and had a patella, smaller tibial Gracilis (Figure 5.4)
rudiment, and no fibular rudiment. Quadratus femoris was
absent bilaterally. Pirani et al. (1991) describe soft tissue Synonyms
anatomy associated with cases of proximal femoral focal This muscle may also be referred to as adductor gracilis or
deficiency (PFFD). In Aitken type A PFFD, the short rectus femoris internus (Knott 1883b).
external rotators of the hip are larger in cross-sectional
diameter than normal and insert onto the posteromedial Typical Presentation
surface of the greater trochanter. In Aitken type B PFFD, Description
the short external rotators are smaller than normal. In a Gracilis originates from the body of the pubis, the inferior
fetus with craniorachischisis, quadratus femoris on the pubic ramus, and the ischial ramus (Standring 2016). It
right lower limb was smaller than normal and less differ- inserts via a tendon onto the medial surface of the proxi-
entiated (Alghamdi et al. 2017). mal tibia just below the medial condyle (Standring 2016).
The insertion tendon contributes to the pes anserinus, and
Prevalence some fibers blend with the deep fascia of the lower leg
FIGURE 5.4 Medial thigh muscles in anterior view.

Innervation Innervation
Gracilis is innervated by the obturator nerve (Standring Adductor longus is innervated by the anterior division of
2016). the obturator nerve (Standring 2016).
Comparative Anatomy Comparative Anatomy

Gracilis has a similar typical presentation in the apes, Adductor longus has a similar typical presentation in the
extending from the pubis, pubic ramus, and adjoining apes, extending from the pubis to the middle of the medial
ischial ramus to the medial aspect of the proximal tibia lip of the linea aspera or posteromedial femoral shaft
(Champneys 1872; Hepburn 1892; Beddard 1893; Sonntag (Champneys 1872; Hepburn 1892; Beddard 1893; Sonntag
1924; Raven 1950; Miller 1952; Sigmon 1974; Gibbs 1999; 1924; Raven 1950; Miller 1952; Sigmon 1974; Gibbs 1999;
Diogo et al. 2010, 2012, 2013a,b, 2017). In bonobos, gracilis Diogo et al. 2010, 2012, 2013a,b, 2017). Adductor longus
has been observed to fuse distally with sartorius, semiten- inserts more proximally in gibbons than in the other apes
dinosus, or adductor brevis (Miller 1952; Diogo et al. 2017). (Appleton and Ghey 1929). In orangutans, gibbons, com-
mon chimpanzees, and bonobos the muscles may be com-
Variations pletely or partially fused with the other adductors, vastus
Description medialis, or pectineus (Champneys 1872; Beddard 1893;
Gracilis may be absent (Sainsbury and Wagget 1984). Gracilis Sonntag 1924; Miller 1952; Gibbs 1999; Diogo et al. 2017).
may send fibers to the fascia lata or fascia of the leg (Macalister
Variations
1875; Knott 1883b; Bergman et al. 1988; du Plessis and Loukas
2016). The extent of its origin may vary (Macalister 1875). It Description
may have a common origin with adductor longus (Tuite et al. Adductor longus may be doubled (Standring 2016) or bilam-
1998). The origin of gracilis may be divided (biceps gracilis), inar (Macalister 1875; Knott 1883b). It may be fused with
with an upper head that originates from the pubis and a lower adductor magnus, adductor brevis, or both muscles (Mori
head that originates from the ischial ramus (Bergman et al. 1964; Bergman et al. 1988; du Plessis and Loukas 2016).
1988; du Plessis and Loukas 2016). It may also be split at its Adductor longus may insert more distally, as far as the knee
insertion (Macalister 1875). Gracilis is frequently associated (Macalister 1875; Bergman et al. 1988; du Plessis and Loukas
with a slip that merges with the tendon of the medial head of 2016). It may receive a slip from pectineus (Macalister 1875).
gastrocnemius (Standring 2016). Adductor longus may also send a fasciculus to the inner side
of vastus medialis (Macalister 1875; Knott 1883b).
Prevalence Thomson (1885) reports a case in which adductor lon-
Tuite et al. (1998) studied the origin of adductor longus in gus had a wide origin that extended onto the ilio-pectin-
37 cadavers. In one case (2.7%), adductor longus had a com- eal line for 1 inch and blended with the superficial fibers
mon origin with gracilis. of pectineus. Tuite et al. (1998) found that the origin of
adductor longus may have two heads, may be fused with
Anomalies the origin of gracilis, or may be partially muscular. Upasna
N/A et al. (2013) describe bilateral variations in the lower limbs
of a female cadaver. Adductor longus and pectineus had a
Clinical Implications combined origin from the pecten pubis, and an adductor
If the slip of gracilis that blends with the tendon of the brevis was divided in two and had distal fibers that merged
medial head of gastrocnemius is undivided from the main with those of adductor longus. These variations were asso-
portion of this muscle, there may be complications during ciated with bilateral accessory slips located under adduc-
surgical harvest of the gracilis tendon (Standring 2016). tor brevis.
Prevalence
Adductor longus (Figure 5.4) Knott (1883b) found that adductor longus was bilaminar
in 1 out of 40 cases (2.5%). Mori (1964) categorized the
Synonyms fusion of the adductors into eight types. All three adduc-
N/A tors fused near their origins in 4% of cases, only adductor
longus and adductor brevis fused near their origins in 4% of
cases, all three adductors fused near their insertions in 8%
Description of cases, only adductor longus and adductor magnus fused
Adductor longus originates via a tendon from the front of near their insertions in 38% of cases, only adductor brevis
the pubis in between the crest and the symphysis (Standring and adductor magnus fused near their insertions in 6% of
2016). It has an aponeurotic insertion into the linea aspera cases, adductor brevis was completely fused with adductor
on the middle third of the femur and typically blends with magnus in 8% of cases, adductor magnus fused completely
vastus medialis, adductor brevis, and adductor magnus with adductor minimus in 8% of cases, and there was no
(Standring 2016). fusion between any of the adductors in 14% of cases.
Tuite et al. (1998) studied the origin of adductor longus Sigmon 1974; Gibbs 1999; Diogo et al. 2010, 2012, 2013a,b,
in 37 cadavers. In nine cadavers (24.3%), muscular fibers 2017). Adductor brevis may be fused with adductor magnus
comprised the lateral origin of adductor longus, ranging in in all apes (Hepburn 1892; Sigmon 1974; Gibbs 1999; Diogo
width from 5 to 11 mm. In one case (2.7%), adductor longus et al. 2017). In orangutans, it may be divided, with one part
had two heads. In one other case (2.7%), adductor longus arising from the superior pubic ramus and the other from
had a common origin with gracilis. the inferior pubic ramus (Hepburn 1892; Sonntag 1924;
Sigmon 1974; Ferrero 2011; Ferrero et al. 2012; Diogo et al.
Anomalies 2013b). It may also be divided into two parts in gorillas
Description (Gibbs 1999; Diogo et al. 2010) and common chimpanzees
Aziz (1980) notes that on both sides of one neonate with tri- and bonobos (Champneys 1872; Hepburn 1892; Beddard
somy 13, a muscular slip arose superficial to the main belly 1893; Diogo et al. 2017).
of adductor longus and inserted on the medial surface of
the proximal tibia close to the insertion of sartorius. On the Variations
right lower limb of a fetus with craniorachischisis dissected Description
by Alghamdi et al. (2017), some fibers of adductor magnus Adductor brevis may be partially or completely divided into
fused with adductor longus. Adductor longus was doubled two or three parts (Macalister 1875; Knott 1883b; Ochiltree
on the right side in a male fetus with triploidy dissected by 1912; Bergman et al. 1988; du Plessis and Loukas 2016;
Moen et al. (1984). Pirani et al. (1991) describe soft tissue Standring 2016). This muscle may be fused with or incor-
anatomy associated with cases of proximal femoral focal porated into adductor magnus (Macalister 1875; Mori 1964;
deficiency (PFFD). In Aitken type B PFFD, adductor lon- Bergman et al. 1988; du Plessis and Loukas 2016; Standring
gus is smaller than normal and oriented more perpendicular 2016). It may also be fused with both adductor longus and
to the femur than is typical. In Aitken type D PFFD, adduc- adductor magnus or only adductor longus (Mori 1964).
tor longus inserts onto the distal femoral remnant. Adductor brevis may also have a connection with obturator
externus (Bergman et al. 1988; du Plessis and Loukas 2016).
Prevalence Macalister (1875) notes more specifically that adductor bre-
An accessory slip associated with adductor longus (Aziz 1980) vis may send a slip to the tendon of obturator externus.
was only observed in 1 out of 20 individuals with trisomy 13 Upasna et al. (2013) found bilateral variations in the
(5%) included in the literature review of Smith et al. (2015). lower limbs of a female cadaver. Adductor longus and pec-
tineus had a combined origin from the pecten pubis, and
Clinical Implications an adductor brevis was divided in two and had distal fibers
N/A that merged with those of adductor longus. These variations
were associated with bilateral accessory slips. The authors
Adductor brevis (Figure 5.4) describe the accessory muscle on the left side in detail, not-
ing it was located under adductor brevis and overlied obtu-
Synonyms rator externus and the posterior division of the obturator
N/A nerve. Its fibers extended between the inferior pubic ramus
and merged with the aponeurosis of adductor magnus.
Adductor brevis originates from the body and inferior Mori (1964) categorized the fusion of the adductors into
ramus of the pubis (Standring 2016). It has an aponeurotic eight types. All three adductors fused near their origins
insertion into the femur from the lesser trochanter to the in 4% of cases, only adductor longus and adductor brevis
linea aspera, and onto the upper aspect of the linea aspera fused near their origins in 4% of cases, all three adductors
behind pectineus (Standring 2016). fused near their insertions in 8% of cases, only adductor
longus and adductor magnus fused near their insertions in
Innervation 38% of cases, only adductor brevis and adductor magnus
Adductor brevis is innervated by the obturator nerve fused near their insertions in 6% of cases, adductor brevis
(Standring 2016). was completely fused with adductor magnus in 8% of cases,
adductor magnus fused completely with adductor minimus
Comparative Anatomy in 8% of cases, and there was no fusion between any of the
Adductor brevis has a similar typical presentation in the adductors in 14% of cases.
apes, originating from the pubis and inferior pubic ramus,
and sometimes the superior pubic ramus, with an insertion Anomalies
onto the femur below the lesser trochanter and onto the Description
upper third of the medial lip of the linea aspera on the mid- Bersu et al. (1976) describe a male infant with Hanhart syn-
dorsal femoral surface (Champneys 1872; Hepburn 1892; drome. The femora of this infant were normally developed,
Beddard 1893; Sonntag 1924; Raven 1950; Miller 1952; but distal secondary ossification centers were absent. The
left leg stump had a patella and a small rudiment of the and in orangutans, it may be divided proximally (Ferrero
proximal tibia but no fibular rudiment. The right leg stump 2011; Ferrero et al. 2012; Diogo et al. 2013b). Adductor
was less developed and had a patella, smaller tibial rudi- minimus is only present as a distinct muscle in about two-
ment, and no fibular rudiment. On the left side, adductor thirds of great apes (Hepburn 1892; Sigmon 1974; Gibbs
brevis inserted via a tendon it shared with the superior and 1999; Diogo et al. 2013, 2013b, 2017). In gibbons, adduc-
middle bundles of adductor magnus into the middle third of tor minimus is usually absent or not distinct from the other
the linea aspera. On the right side, the origin of this muscle adductors (Hepburn 1892; Sigmon 1974; Diogo et al. 2012).
was displaced inferiorly.
Pirani et al. (1991) describe soft tissue anatomy asso- Variations
ciated with cases of proximal femoral focal deficiency Description
(PFFD). In Aitken type B PFFD, adductor brevis is smaller The portion of the muscle referred to as adductor mini-
than normal and oriented more perpendicular to the femur mus can be variably separated from the rest of the muscle
than is typical. In Aitken type D PFFD, adductor brevis (Macalister 1875; Mori 1964; Tubbs et al. 2011). The por-
inserts onto the distal femoral remnant. tion of adductor magnus that extends between the ischial
tuberosity and the medial condyle of the femur is also vari-
Prevalence ably separated from the rest of the muscle (Macalister 1975;
N/A Knott 1883b; du Plessis and Loukas 2016; Standring 2016).
Adductor magnus may fuse with quadratus femoris, adduc-
tor brevis, and/or adductor longus (Macalister 1875; Mori
N/A 1964; Bergman et al. 1988; du Plessis and Loukas 2016).
There may be accessory slips of adductor magnus that
Adductor magnus (Figure 5.4) originate from the linea aspera and insert into the adductor
magnus tendon and the adductor tubercle (Macalister 1875;
Synonyms Hildebrand 1978; Bergman et al. 1988; Tubbs and Zehren
The portion of adductor magnus that extends between the 2006; du Plessis and Loukas 2016).
pubic ramus and the gluteal tuberosity may be referred to
as adductor minimus, adductor quartus (Macalister 1875; Prevalence
Knott 1883a; Standring 2016; du Plessis and Loukas 2016), Knott (1883b) found that the portion of adductor magnus
pars lateralis, caput superius externum, or premier faisceau that inserts onto the medial condyle of the femur was com-
du grand adducteur (Tubbs et al. 2011). pletely separate from the rest of the muscle in 1 out of 40
cases (2.5%).
Typical Presentation Mori (1964) categorized the fusion of the adductors into
Description eight types. All three adductors fused near their origins
Adductor magnus originates from the inferior pubic ramus, in 4% of cases, only adductor longus and adductor brevis
ischial ramus, and the ischial tuberosity (Standring 2016). fused near their origins in 4% of cases, all three adductors
The fibers that originate from the pubic ramus insert onto fused near their insertions in 8% of cases, only adductor
the gluteal tuberosity of the femur (Standring 2016). The longus and adductor magnus fused near their insertions in
fibers that originate from the ischial ramus have an aponeu- 38% of cases, only adductor brevis and adductor magnus
rotic insertion onto the linea aspera and medial supracondy- fused near their insertions in 6% of cases, adductor brevis
lar line (Standring 2016). The fibers that originate from the was completely fused with adductor magnus in 8% of cases,
ischial tuberosity insert via a tendon onto the adductor tuber- adductor magnus fused completely with adductor minimus
cle on the medial condyle of the femur (Standring 2016). in 8% of cases, and there was no fusion between any of the
adductors in 14% of cases. Adductor magnus was divided
Innervation into three portions in 70% of cases and divided into two
Adductor magnus is innervated by both the obturator nerve portions in 20% of cases.
and the tibial division of the sciatic nerve (Standring 2016). Tubbs et al. (2011) studied adductor minimus on 40 sides
from 20 adult cadavers and in five fetuses. Adductor mini-
Comparative Anatomy mus was distinct from adductor magnus on 21 sides (52.5%)
Adductor magnus has a similar typical presentation in the in the adults. Five out of these 21 sides (24%) had partial
apes, originating from the inferior pubic ramus, ischial fusion to adductor magnus. Adductor minimus was distinct
ramus, and ischial tuberosity and inserting onto the medial from adductor magnus on all sides of the fetuses (100%).
lip of the linea aspera and the adductor tubercle (Champneys
1872; Hepburn 1892; Beddard 1893; Sonntag 1924; Raven Anomalies
1950; Miller 1952; Sigmon 1974; Gibbs 1999; Diogo et al. Description
2010, 2012, 2013a,b, 2017). In gibbons, it is often divided On the right lower limb of a fetus with craniorachischi-
into two parts distally (Sigmon 1974; Diogo et al. 2012), sis dissected by Alghamdi et al. (2017), some fibers of
adductor magnus fused with adductor longus. Bersu et al. in orangutans (Sigmon 1974) or bonobos (Miller 1952). It
(1976) describe a male infant with Hanhart syndrome. is divided into two layers/parts in common chimpanzees
The femora of this infant were normally developed but (Macalister 1871; Gibbs 1999), and also in some gorillas,
distal secondary ossification centers were absent. The left according to Ochiltree (1912).
leg stump had a patella and a small rudiment of the proxi-
mal tibia but no fibular rudiment. The right leg stump was Variations
less developed and had a patella, smaller tibial rudiment, Description
and no fibular rudiment. On the left side, the inferior bun- Pectineus may be divided into two layers or parts (Macalister
dle of adductor magnus originated from the ischial tuber- 1875; Knott 1883b; Ochiltree 1912; Bergman et al. 1988; du
osity, along with semitendinosus and biceps femoris, and Plessis and Loukas 2016; Standring 2016). It may receive
inserted onto the junction of the middle and lower thirds fibers from adductor longus (Macalister 1875; Bergman
of the linea aspera just above the adductor tubercle. Pirani et al. 1988). Its origin may be connected with the origin of
et al. (1991) describe soft tissue anatomy associated with adductor brevis (Ochiltree 1912). Accessory bundles may
cases of proximal femoral focal deficiency (PFFD). In connect pectineus with obturator externus, iliacus, the hip
Aitken type B PFFD, adductor magnus is oriented more joint capsule, or the lesser trochanter (Macalister 1875;
perpendicular to the femur than is typical. In Aitken type Knott 1883b; Bergman et al. 1988; du Plessis and Loukas
D PFFD, adductor magnus inserts onto the distal femoral 2016; Standring 2016). Upasna et al. (2013) describe bilat-
remnant. eral variations in the lower limbs of a female cadaver.
Adductor longus and pectineus had a combined origin from
Prevalence the pecten pubis, and an adductor brevis was divided in two
N/A and had distal fibers that merged with those of adductor lon-
gus. These variations were associated with bilateral acces-
Clinical Implications sory slips located under adductor brevis.
Accessory slips of adductor magnus may compress the
proximal popliteal vein and cause an aneurysm (Tubbs and Prevalence
Zehren 2006; du Plessis and Loukas 2016). Knott (1883b) found that pectineus was divided into two
parts in 1 out of 40 specimens (2.5%).
Pectineus (Figure 5.4) Anomalies
N/A Bersu et al. (1976) describe a male infant with Hanhart
syndrome. The femora of this infant were normally devel-
Typical Presentation oped but distal secondary ossification centers were absent.
Description The left leg stump had a patella and a small rudiment of
Pectineus originates from the pecten pubis (Standring the proximal tibia but no fibular rudiment. The right leg
2016). It inserts onto the femur between the lesser trochan- stump was less developed and had a patella, smaller tibial
ter and the linea aspera (Standring 2016). rudiment, and no fibular rudiment. On the right side, the
origin of pectineus was shifted medially, and it inserted
Innervation via a tendon onto the middle third of the linea aspera in
Pectineus is typically innervated by the femoral nerve close association with adductor brevis and longus. Pirani
(Standring 2016). It may also be innervated by the acces- et al. (1991) describe soft tissue anatomy associated with
sory obturator nerve, when present, and sometimes from a cases of proximal femoral focal deficiency (PFFD). In
branch of the obturator nerve (Standring 2016). If divided Aitken type B PFFD, pectineus is smaller than is typical.
into two parts or layers, the deep layer (also referred to as In Aitken type D PFFD, pectineus inserts onto the proxi-
dorsal or medial) is innervated by the obturator nerve, and mal femoral ossicle.
the superficial layer (also referred to as ventral or lateral) is
innervated by the femoral nerve (Ochiltree 1912; Bergman Prevalence
et al. 1988; Standring 2016; du Plessis and Loukas 2016). N/A

Pectineus has a similar typical presentation in the apes, N/A
extending from the pubis and/or superior pubic ramus to the
femur just below the lesser trochanter (Champneys 1872; Semitendinosus (Figure 5.5)
Hepburn 1892; Beddard 1893; Sonntag 1924; Raven 1950;
Miller 1952; Sigmon 1974; Gibbs 1999; Diogo et al. 2010, Synonyms
2012, 2013a,b, 2017). It may be fused with adductor longus N/A
FIGURE 5.5 Posterior thigh muscles in posterior view.
Typical Presentation Comparative Anatomy

Description Semitendinosus has a similar typical presentation in the
Semitendinosus is one of the three hamstring muscles apes, originating from the ischial tuberosity with biceps fem-
(Standring 2016). It originates via a tendon from the ischial oris and inserting via a narrow tendon onto the medial tibia
tuberosity (Standring 2016). Its tendon inserts onto the (Champneys 1872; Hepburn 1892; Beddard 1893; Sonntag
medial aspect of the proximal tibia (Standring 2016). It 1924; Raven 1950; Miller 1952; Sigmon 1974; Gibbs 1999;
joins with the tendon of gracilis upon insertion and has an Diogo et al. 2010, 2012, 2013a,b, 2017). A tendinous inter-
expansion to the deep fascia of the leg and the medial head section in the muscle is sometimes present in all apes except
of gastrocnemius (Standring 2016). gorillas (Macalister 1871; Hepburn 1892; Gibbs 1999). The
common origin may include semimembranosus in common
Innervation chimpanzees, bonobos, and orangutans (Champneys 1872;
Semitendinosus is innervated by the tibial division of the Hepburn 1892; Beddard 1893; Sonntag 1924; Sigmon 1974;
sciatic nerve (Standring 2016). Gibbs 1999; Diogo et al. 2017). The insertion is more distal
onto the tibia in common chimpanzees, bonobos, orang- normal. In infants with trisomy 18, Ramirez-Castro and
utans, and gibbons than in gorillas and humans (Champneys Bersu (1978) found two types of accessory bellies associ-
1872; Hepburn 1892; Gibbs 1999). ated with semitendinosus. One type originated between
semitendinosus and the long head of biceps femoris and
Variations inserted onto the semitendinosus tendon. The second type
Description originated from the sacrotuberous ligament and inserted
A tendinous inscription is often present near the midpoint onto the belly of semitendinosus near its origin (Ramirez-
of the muscle (du Plessis and Loukas 2016; Standring Castro and Bersu 1978; Smith et al. 2015).
2016), and it may be reduced or doubled (Macalister 1875;
Knott 1883b). Semitendinosus may be completely sepa- Prevalence
rated from biceps femoris (Macalister 1875; Bergman et In their literature review, Smith et al. (2015) found that the
al. 1988; du Plessis and Loukas 2016). Semitendinosus accessory belly between semitendinosus and the long head
may receive a slip from the long head of biceps femo- of biceps femoris was present in 3 out of 17 (17.6%) indi-
ris (Bergman et al. 1988; Chakravarthi 2013; du Plessis viduals with trisomy 18. An accessory belly between semi-
and Loukas 2016; Standring 2016; Hoban et al. 2019). It tendinosus and the sacrotuberous ligament was present in 2
may be partially fused with semimembranosus (Bergman out of 17 (11.8%) individuals with trisomy 18.
et al. 1988; du Plessis and Loukas 2016). It may have an
origin from the sacrotuberous ligament (Bergman et al. Clinical Implications
1988; du Plessis and Loukas 2016). Accessory slips of Attachments of semitendinosus to biceps femoris and gas-
semitendinosus may also originate from the sacrotuber- trocnemius may cause complications when surgically har-
ous ligament, coccyx, or ischial tuberosity (Bergman et al. vesting the semitendinosus tendon (Standring 2016). A
1988; Chakravarthi 2013; Fraser et al. 2013; du Plessis and supernumerary semitendinosus muscle may compress the
Loukas 2016). sciatic nerve or popliteal artery (Paraskevas et al. 2010).
Gray (1945) reports two cases of semitendinosus varia- Variation in the origins of the hamstring muscles may
tions. In one case, the three hamstring muscles originated predispose individuals to strains and posterior thigh pain
from a common tendon that had a superior attachment to (Fraser et al. 2013). An origin of semitendinosus from the
the sacrotuberous ligament and fascial sheath of pirifor- ischial tuberosity may predispose individuals to posterior
mis. The tendon received slips from quadratus femoris, the thigh or pelvic floor pain (Fraser et al. 2013). A slip con-
ischium, and the femur and gave rise to the muscle bellies necting biceps femoris and semitendinosus could compress
of the hamstrings at mid-thigh. In the second case, bilat- the sciatic nerve, causing pain, or interfere with popliteal
eral supernumerary muscles arose from the linea aspera sciatic nerve block (Hoban et al. 2019). Accessory slips or
between the short head of biceps femoris and adductor muscles associated with semitendinosus may simulate soft-
magnus and inserted onto the dorsal aspect of the capsule of tissue tumors (Chakravarthi 2013).
the knee joint and into the fascia covering the medial head
of gastrocnemius (Gray 1945). Gray (1945) suggests that
these bilateral supernumerary muscles should be regarded
Semimembranosus (Figure 5.5)
as femoral heads of semitendinosus. Synonyms
Paraskevas et al. (2010) report a supernumerary semi- N/A
tendinosus muscle that originated from the lateral lip of the
linea aspera between adductor magnus and the short head Typical Presentation
of biceps femoris and inserted onto the medial tibial con- Description
dyle. The distal portion of this muscle was covered by the
Semimembranosus is one of the three hamstring muscles
long head of biceps femoris. These authors suggest that an
(Standring 2016). It originates via a tendon from the ischial
anomaly during the 90° medial rotation of the lower limb
tuberosity (Standring 2016). Some fibers blend with biceps
bud between weeks 6 and 8 of gestation may have created
femoris and semitendinosus (Standring 2016). Its insertion
the supernumerary muscle (Paraskevas et al. 2010).
tendon is divided into five parts (Standring 2016). The main
Prevalence part inserts onto a tubercle on the medial condyle of the tibia
(Standring 2016). The other parts of the insertion go to the
N/A
medial border of the tibia, the posterior side of the medial
tibial condyle, the fascia over popliteus, the femoral inter-
Anomalies condylar line, and lateral femoral condyle (Standring 2016).
Description
Pirani et al. (1991) describe soft tissue anatomy associated Innervation
with cases of proximal femoral focal deficiency (PFFD). Semimembranosus is innervated by the tibial division of
In Aitken type B PFFD, semitendinosus is smaller than the sciatic nerve (Standring 2016).
Comparative Anatomy Anomalies

Semimembranosus has a similar typical presentation in the Description
apes, extending from the ischial tuberosity to the posterior sur- Bersu et al. (1976) describe a male infant with Hanhart
face of the medial tibial condyle, with some attachments to the syndrome. The femora of this infant were normally devel-
fascia and ligaments of the knee (Champneys 1872; Hepburn oped but distal secondary ossification centers were absent.
1892; Beddard 1893; Sonntag 1924; Raven 1950; Sigmon The left leg stump had a patella and a small rudiment of
1974; Gibbs 1999; Diogo et al. 2010, 2012, 2013a,b, 2017). the proximal tibia but no fibular rudiment. The right leg
stump was less developed and had a patella, smaller tibial
Variations rudiment, and no fibular rudiment. Semimembranosus was
Description absent bilaterally. Pettersen et al. (1979) found extra slips of
Semimembranosus is variable in size and may be reduced semimembranosus in two neonates with trisomy 13. In one
or absent (Macalister 1875; Bergman et al. 1988; Moncayo case, the slip was on the right side and originated from the
et al. 2010; du Plessis and Loukas 2016; Standring 2016). ischial tuberosity anterior to the origin of the normal semi-
When absent, it may be replaced by a tendon with a small membranosus, and its fibers fused with the main muscle at
central muscle belly (Macalister 1875). Liu et al. (2011) found midthigh. On the left side of the second case, the accessory
that in one case, bilateral absence of semimembranosus was slip ran along the length of the normal semimembranosus
associated with bilateral absence of quadratus femoris. The and the two fused upon insertion. On the right side of the
absence of semimembranosus may also be associated with second case, the accessory slip was fused anterior to the
variations in the length and shape of the other hamstring main muscle. Pirani et al. (1991) describe soft tissue anat-
muscles (Moncayo et al. 2010; Sussmann 2019). omy associated with cases of proximal femoral focal defi-
Its proximal attachment may extend to the coccyx ciency (PFFD). In Aitken type B PFFD, semimembranosus
(Bergman et al. 1988). Its origin may be primarily from the is smaller than normal.
sacrotuberous ligament in some cases (Macalister 1875).
Prevalence
Slips from its insertion may be continuous with the arcu-
ate ligament or end in the popliteal fat (Macalister 1875; In their literature review, Smith et al. (2015) found that
Knott 1883b). The distal tendon may course along the tibial extra slips of semimembranosus were present in 2 out of 20
collateral ligament and insert more distally than normal individuals (10%) with trisomy 13. An extra slip between
(Moncayo et al. 2010). There may be a sixth insertion/ten- the hamstrings was found in 1 out of 17 individuals with
dinous expansion of semimembranosus onto the posterior trisomy 18 (5.9%).
horn of the lateral meniscus (Kim et al. 1997).
Semimembranosus may send a slip to adductor magnus
or the femur (Standring 2016). It may have a slip continuous Liu et al. (2011) suggest that the absence of semimembra-
with quadratus femoris (Macalister 1875). It may be partially nosus may present as muscle weakness of knee flexion or
fused with semitendinosus (Bergman et al. 1988; du Plessis hypermobility of the posteromedial knee joint. Variation
and Loukas 2016). Gray (1945) reports a case in which the in the origins of the hamstring muscles may predispose
three hamstring muscles originated by a common tendon that individuals to strains and posterior thigh pain (Fraser et al.
had a superior attachment to the sacrotuberous ligament and 2013). Accessory semimembranosus bellies may become
fascial sheath of piriformis. The tendon received slips from entrapped by the semitendinosus tendon and cause pain
quadratus femoris, the ischium, and the femur. The tendon (Zeren et al. 2009). A tendon attachment to the lateral
gave rise to the muscle bellies of the hamstrings at mid-thigh. meniscus may be misdiagnosed as a lateral meniscus tear
Semimembranosus may also be doubled or divided longi- (Kim et al. 1997).
tudinally (Bergman et al. 1988; du Plessis and Loukas 2016;
Standring 2016). The division may be confined to the tendon
of origin in some cases (Knott 1883b). When doubled, the
Biceps femoris (Figure 5.5)
accessory belly may present as a fleshy slip with proximal See also: Tenuissimus, Tensor fasciae suralis
and distal tendons or may arise from the sacrotuberous liga-
ment (Bergman et al. 1988; du Plessis and Loukas 2016). Synonyms
The distal tendon of the accessory belly can have a variable This muscle may also be referred to as biceps flexor cruris
attachment (Bergman et al. 1988; du Plessis and Loukas (Macalister 1875; Knott 1883a,b).
2016). Semimembranosus can be comprised of up to four
bellies (Bergman et al. 1988; du Plessis and Loukas 2016). Typical Presentation
Description
Prevalence Biceps femoris is one of the three hamstring muscles
Kim et al. (1997) found a tendinous insertion of semimem- (Standring 2016). The long head originates by a tendon
branosus onto the posterior horn of the lateral meniscus in from the ischial tuberosity and from the sacrotuberous
16 out of 37 knees (43.2%). ligament (Standring 2016). The short head originates from
the linea aspera (Standring 2016). Biceps femoris inserts Loukas 2016). See the entry for tensor fasciae suralis for
mainly onto the head of the fibula (Standring 2016). There more details. An accessory digastric muscle can also pres-
are additional insertions onto the fibular collateral ligament ent with a superior belly arising from or near piriformis,
and lateral tibial condyle (Standring 2016). continuing down the thigh as a long intermediate tendon,
and connecting to an inferior belly that is closely associ-
Innervation ated with biceps femoris (Moore 1922; Suda and Takahashi
The long head of biceps femoris is innervated by the tibial divi- 1957; Akita et al. 1992; Nicholson et al. 2016). Tenuissimus,
sion of the sciatic nerve, and the short head is innerved by the another supernumerary muscle, originates from the fascia
common fibular division of the sciatic nerve (Standring 2016). on the deep surface of gluteus maximus and often inserts
at the junction of the short and long head of biceps femoris
Comparative Anatomy (see the entry for this muscle) (Green 1931).
Biceps femoris has a similar typical presentation in the
apes. The long head originates with semitendinosus from Prevalence
the ischial tuberosity and the short head from the lateral lip Mori (1964) found that the muscle of the long head of biceps
of the linea aspera and the intermuscular septum. The long femoris was completely isolated from the muscle of the short
head inserts into the tibial head, lateral tibial condyle, fibular head in 30% of cases. The insertion tendon of the long head
head, iliotibial tract, knee joint capsule, and fascia of the leg fused with the muscle of the short head in 70% of cases.
while the short head inserts into the fibular head and fascia
of the leg (Champneys 1872; Hepburn 1892; Beddard 1893;
Anomalies
Sonntag 1924; Raven 1950; Miller 1952; Sigmon 1974; Gibbs
1999; Diogo et al. 2010, 2012, 2013a,b, 2017). The short and Description
long heads are fused in most cases (Hepburn 1892; Sigmon Bersu et al. (1976) describe a male infant with Hanhart syn-
1974; Gibbs 1999). The origin of the short head extends more drome. The femora of this infant were normally developed,
distally in great apes than in gibbons or humans (Sigmon but distal secondary ossification centers were absent. The
1974). Accessory bellies have been noted in common chim- left leg stump had a patella and a small rudiment of the prox-
panzees and bonobos (Diogo et al. 2013a, 2017). imal tibia but no fibular rudiment. The right leg stump was
less developed and had a patella, smaller tibial rudiment,
Variations and no fibular rudiment. On both sides, the long head of
Description biceps femoris inserted onto the lateral surface of the tibia.
The short head is sometimes absent (Macalister 1875; Knott The short head of the biceps was absent on the right side. On
1883a,b; Bergman et al. 1988; du Plessis and Loukas 2016; the left side, it inserted into the posterior surface of the knee
Standring 2016). The short head may also split into mul- capsule and the lateral surface of the tibia, independently of
tiple fasciculi (Macalister 1875). The long head and short the long head. On the right side, an accessory slip originated
head can be partially or completely separated (Mori 1964; from the medial portion of the belly of the long head and
Bergman et al. 1988; du Plessis and Loukas 2016) and may inserted posterior to the insertion of the main tendon.
only unite at their insertion (Macalister 1875; Chakravarthi In infants with trisomy 18, Ramirez-Castro and Bersu
2013). The two heads may insert separately (Macalister (1978) found two types of accessory bellies associated with
1875). Biceps femoris may have a more prominent attach- semitendinosus. One type originated between semitendino-
ment onto the proximal tibia (Macalister 1875; Knott 1883b; sus and the long head of biceps femoris and inserted onto
Date et al. 2012; du Plessis and Loukas 2016). It may also the semitendinosus tendon (Ramirez-Castro and Bersu 1978;
not have a fibular insertion and attach entirely onto the prox- Smith et al. 2015). Pettersen et al. (1979) found an extra belly
imal tibia (Kristensen et al. 1989; Hernandez et al. 1996). of the long head (third head) of biceps femoris running lateral
The long head of biceps femoris may send a slip to to the normal long head in one neonate with trisomy 13. This
semitendinosus (Bergman et al. 1988; Chakravarthi 2013; accessory belly inserted via a tendon onto the short head.
du Plessis and Loukas 2016; Standring 2016; Hoban et al. Pirani et al. (1991) describe soft tissue anatomy associated
2019). Accessory slips or a third head of biceps femoris with cases of proximal femoral focal deficiency (PFFD). In
originate most commonly from the ischial tuberosity, but Aitken type B PFFD, biceps femoris is smaller than normal.
may also arise from the coccyx, sacrum, sacrotuberous
ligament, gluteus maximus, linea aspera, gluteal tuberos- Prevalence
ity, fascia lata, adductor magnus, vastus lateralis, gastroc- In their literature review, Smith et al. (2015) found that an
nemius, medial supracondylar line, or the lateral condyle extra belly of biceps femoris was present in 1 out of 20 indi-
of the femur (Macalister 1875; Knott 1883b; Bergman et al. viduals (5%) with trisomy 13 (case described by Pettersen
1988; du Plessis and Loukas 2016; Standring 2016). et al. 1979). An extra slip between the hamstrings was found
There may be a crural extension of biceps femoris, acces- in 1 out of 17 individuals with trisomy 18 (5.9%). Smith
sory muscle tensor fasciae suralis, that presents as a slip to et al. (2015) also found that the accessory belly between
the fascia of the leg, the Achilles tendon, and/or gastroc- semitendinosus and the long head of biceps femoris was
nemius (Knott 1883b; Bergman et al. 1988; du Plessis and present in 3 out of 17 (17.6%) individuals with trisomy 18.
Clinical Implications rise to a third head of gastrocnemius that inserted into the
Variation in the origins of the hamstring muscles may junction of the two heads of gastrocnemius.
predispose individuals to strains and posterior thigh pain Tensor fasciae suralis ends in a tendon that joins the cru-
(Fraser et al. 2013). An anomalous insertion of the biceps ral fascia of the lower leg over the gastrocnemius muscle
femoris tendon may cause pain and snapping at the knee (Knott 1883b; Schaeffer 1913; Barry and Bothroyd 1924;
(Kristensen et al. 1989; Hernandez et al. 1996; Date et al. Miyauchi et al. 1985; Bergman et al. 1988; Tubbs et al.
2012). A slip connecting biceps femoris and semitendino- 2006e; du Plessis and Loukas 2016; Arakawa et al. 2017;
sus could compress the sciatic nerve, causing pain, or inter- Bale and Herrin 2017; George et al. 2019). Its insertion
fere with popliteal sciatic nerve block (Hoban et al. 2019). may also pass distally between the two heads of gastroc-
Accessory bellies of biceps femoris may compress the nemius and join with the Achilles tendon (Kelch 1813;
common fibular nerve and cause foot drop (Kaplan et al. Gruber 1873c, 1879; Halliburton 1881; Turner 1884–1885;
2008). Accessory slips or muscles associated with biceps Somayaji et al. 1998).
femoris may simulate soft-tissue tumors (Chakravarthi Tensor fasciae suralis may be digastric (Halliburton 1881;
2013). Gruber 1879; Arakawa et al. 2017). Gruber (1879) describes a
bilateral case. On the right side, the superior belly originated
from the long head of the biceps and ended in a tendon in the
Tensor fasciae suralis (Figure 5.5)
popliteal fossa. Across from the intercondylar space, the ten-
See also: Semitendinosus, Biceps femoris, Gastrocnemius don gave rise to the inferior belly which coursed through the
groove between the heads of gastrocnemius and inserted into
Synonyms the Achilles tendon. On the left side, the muscle was similar,
This muscle may also be referred to as ischioaponeuroticus but the superior belly received a tendinous bundle from the
(Bergman et al. 1988). George et al. (2019) propose fem- tendinous inscription of semitendinosus.
eroaponeuroticus for the variant of tensor fasciae suralis Halliburton (1881) describes a similar case in which
described in their study. the superior belly of a supernumerary muscle arose from
the long head of biceps femoris. At the top of the popli-
Typical Presentation teal space, it gave rise to a tendon that flattened into an
This muscle is only present as a variation. aponeurosis and attached to the fibular head. The inferior
belly originated from the tendon where it expanded into the
Comparative Anatomy aponeurotic slip and was situated in the groove between
Martinoli et al. (2010) suggest that tensor fasciae suralis the two heads of gastrocnemius. The inferior belly ended
may represent a vestigial extension of the hamstrings, as in a tendon that inserted into gastrocnemius just below the
hamstring insertion in some mammals extends more dis- union of the two heads. Its tendinous fibers ultimately con-
tally to facilitate permanent flexion of the knee. tinued into the Achilles tendon.
Arakawa et al. (2017) describe a case in which they con-
Variations clude that tenuissimus, tensor fasciae suralis, and two other
Description supernumerary muscles are present on the posterior right thigh
When present, tensor fasciae suralis originates from the long of a cadaver. Tenuissimus passed between the long and short
head of biceps femoris, occasionally originating from semi- heads of biceps femoris and fused with the tensor fasciae sura-
tendinosus (Kelch 1813; Gruber 1873c, 1879; Knott 1883b; lis. Tensor fasciae suralis was digastric with a superior belly
Turner 1884–1885; Barry and Bothroyd 1924; Bergman that arose from the medial surface of the long head of biceps
et al. 1988; Tubbs et al. 2006e; du Plessis and Loukas 2016; femoris in the distal third of the muscle. The superior head
Bale and Herrin 2017). It may also originate from a tendon ended in an aponeurosis, which rotated about 90° and gave rise
that is continuous with the biceps femoris tendon (Schaeffer to the inferior belly that fused with the posterior surface of the
1913). George et al. (2019) report an origin from the lower- crural fascia that covered the lateral head of gastrocnemius.
most portion of the linea aspera, arising with the fibers of There may be accessory muscles in the popliteal fossa
the short head of biceps femoris. that could be variants of tensor fasciae suralis or deriva-
It can also arise via two tendons. Miyauchi et al. (1985) tives of nearby muscles (Okamoto et al. 2004; Kim et al.
and Somayaji et al. (1998) describe cases in which one 2009b; Kim et al. 2014). Okamoto et al. (2004) describe
tendon arose from semitendinosus and the other from the a case in which a flat, rectangular muscle extended trans-
long head of biceps femoris (Miyauchi et al. 1985). Barry versely between the lateral border of the medial head of
and Bothroyd (1924) also report an origin via two fascial gastrocnemius and the tendon of biceps femoris, covering
slips, one from the fascia lata at the point of divergence of the structures in the popliteal fossa. As it was innervated by
the biceps femoris and semitendinosus, and the other from the common fibular nerve, Okamoto et al. (2004) suggest
the fascia covering the medial and deep surfaces of biceps that it may be derived from the short head of biceps femoris.
femoris. Gupta and Bhagwat (2006) report a case in which These authors also note a similar case described by Parsons
tensor fasciae suralis originated via two slips, one from (1919–1920), who suggested the muscle was the conversion
semitendinosus and the other from biceps femoris, and gave of the popliteal fascia to fleshy fibers.
Kim et al. (2009b) describe a remarkably similar muscle Anomalies

with innervation from the lateral sural cutaneous nerve. N/A
Kim et al. (2014) describe another similar case in which the
flat transverse muscle was innervated by the tibial nerve.
The authors regard it as a third head of gastrocnemius (gas-
trocnemius tertius) (Kim et al. 2014). Tensor fasciae suralis may appear as a mass or swelling
in the popliteal fossa (Montet et al. 2002). As this muscle
Innervation crosses the popliteal fossa, it may compress the popliteal
Tensor fasciae suralis is typically innervated by a branch vein (Somayaji et al. 1998) and/or other neurovascular
from the tibial nerve (Schaeffer 1913; Barry and Bothroyd structures in the fossa. George et al. (2019) describe a
1924; Somayaji et al. 1998; Tubbs et al. 2006e; Arakawa laterally displaced variant of tensor fasciae suralis and
et al. 2017). George et al. (2019) found innervation from the suggest that its placement may cause compression neu-
common fibular nerve. ropathy to the common fibular nerve and lateral sural
cutaneous nerve.
Prevalence
Somayaji et al. (1998) found only one case of tensor fas-
ciae suralis out of 300 lower limbs (0.3%) from 150 cadav- POSTERIOR MUSCLES OF THE LOWER LEG
ers compiled from 15 years of dissection. In a study of 236
Gastrocnemius (Figure 5.6)
lower limbs, Bale and Herrin (2019) found three cases of
tensor fasciae suralis (1.3%). See also: Tensor fasciae suralis
FIGURE 5.6 Superficial muscles of the posterior lower leg in posterior view. The muscles that are typically present are illustrated on
the left side. Popliteus and two variations are illustrated on the right side.
Synonyms intermedio-medial and intermedio-lateral heads. Rodrigues

N/A et al. (2016) report a case in which the medial and lateral
heads each arose via three bundles from their normal points
Typical Presentation of origin, giving the appearance of a six-headed muscle.
Description The bundles remained separate until they merged to form
The medial head of gastrocnemius originates from the the calcaneal tendon.
posterior aspect of the medial condyle of the femur and A third head of gastrocnemius—gastrocnemius
from the popliteal surface (Standring 2016). The lat- tertius—may originate from the popliteal surface of the
eral head originates from the lateral surface of the lateral femur (Macalister 1875; Knott 1883a,b; Iwai et al. 1987;
femoral condyle (Standring 2016). Fibers from both heads Bergman et al. 1988, 1995; Koplas et al. 2009; Dave et al.
also arise from the knee joint capsule (Standring 2016). 2012; Lambert 2016; Standring 2016). The third head may
Gastrocnemius joins with the tendon of the soleus to form also originate from the long head or tendon of biceps femo-
the calcaneal (Achilles) tendon, which inserts onto the cal- ris, the capsule of the knee joint, the crural fascia, the fib-
caneus (Standring 2016). ula, linea aspera, either supracondylar line, or the lateral
epicondyle of the femur (Macalister 1875; Knott 1883b; Le
Innervation Double 1897; Frey 1919; Mori 1964; Bergman et al. 1988,
Gastrocnemius is innervated by the tibial nerve (Standring 1995; Lambert 2016). It may insert into one or both heads of
2016). gastrocnemius, or at the junction of these heads (Frey 1919;
Mori 1964; Iwai et al. 1987; Bergman et al. 1995; Koplas
et al. 2009; Yildirim et al. 2011; Dave et al. 2012; Lambert
Comparative Anatomy
2016). Plantaris can present as a third head of gastrocne-
Gastrocnemius in the apes is not as enlarged as it is in mius and may blend with the two heads at their junction,
humans, but it similarly originates via two heads from the merge with only one head, or join the fascia on the deep
medial and lateral femoral condyles and the knee joint cap- surface of gastrocnemius (Bergman et al. 1988).
sule and inserts with soleus into the calcaneus (Hepburn Yildirim et al. (2011) describe a cadaver with bilateral
1892; Beddard 1893; Frey 1913; Sonntag 1924; Raven 1950; gastrocnemius tertius and a unilateral accessory soleus
Miller 1952; Gibbs 1999; Diogo et al. 2010, 2012, 2013a,b, muscle. On the right side, gastrocnemius tertius originated
2017). Sesamoid bones are present in both heads in gibbons from the lateral condyle of the femur and inserted onto the
(Hepburn 1892; Frey 1913; Gibbs 1999). The calcaneal ten- medial head of gastrocnemius. On the left, gastrocnemius
don is more defined in gibbons and orangutans than in the tertius had two bellies. The deep belly originated from the
other apes (Diogo et al. 2012, 2013a,b, 2017). tendon of plantaris and the superficial belly from just above
this tendon. The deep belly inserted onto the inner surface
Variations of the medial head of gastrocnemius, and the superficial
Description belly inserted onto the external surface of the lateral head.
The lateral head or the entire muscle may be absent Gastrocnemius is associated with accessory muscle ten-
(Bergman et al. 1988; Lambert 2016; Standring 2016). The sor fasciae suralis (see the entry for this muscle), which typi-
lateral head may also be reduced and replaced by a fibrous cally extends between the hamstrings and gastrocnemius, the
cord (Macalister 1875; Bergman et al. 1988; Lambert 2016). fascia over gastrocnemius, or the Achilles tendon. Given the
The lateral head may have origins from the lateral collat- anatomical similarities between tensor fasciae suralis and
eral ligament or the posterior oblique ligament of the knee gastrocnemius tertius, some authors may designate muscles
(Macalister 1875; Knott 1883b). The medial head may described as tensor fasciae suralis as the third head of gas-
originate higher than usual, from the medial supracondy- trocnemius (Lambert 2016). Gupta and Bhagwat (2006)
lar line (Stuart 1879). Small accessory slips may be present report a case in which tensor fasciae suralis originated via
near the origin of either head (Takase et al. 1997; Liu et al. two slips, one from semitendinosus and the other from biceps
2005; Rochier and Sumpio 2009). The length of the gas- femoris, and gave rise to a third head of gastrocnemius that
trocnemius aponeurosis and the extent of its attachments inserted into the junction of the two heads of gastrocnemius.
to soleus vary (Blitz and Eliot 2007). The two heads may Okamoto et al. (2004) describe a case in which a flat,
remain separate until their insertion onto the calcaneus or rectangular muscle extended transversely between the lat-
remain separate from soleus and its tendon (Macalister eral border of the medial head of gastrocnemius and the
1875; Le Double 1897; Bergman et al. 1988; Lambert 2016). tendon of biceps femoris, covering the structures in the
A sesamoid bone may be found in either head (Macalister popliteal fossa. As it was innervated by the common fibular
1875; Mori 1964). The whole muscle may be bilaminar nerve, Okamoto et al. (2004) suggest that it may be derived
(Macalister 1875; Le Double 1897). from the short head of biceps femoris. These authors also
One or both heads may be doubled (Bergman et al. note a similar case described by Parsons (1919–1920), who
1988; Ashaolu et al. 2014). Ashaolu et al. (2014) found suggested the muscle was the conversion of the popliteal
numerous cases of a “four-headed” gastrocnemius that fascia to fleshy fibers. Kim et al. (2009b) describe a remark-
included the normal medial and lateral heads and smaller ably similar muscle with innervation from the lateral sural
cutaneous nerve. Kim et al. (2014) also describe another Prevalence

similar case in which the flat transverse muscle was inner- In their literature review, Smith et al. (2015) found that an
vated by the tibial nerve. The authors regard it as a third extra slip of gastrocnemius was present in 1 out of 20 neo-
head of gastrocnemius (Kim et al. 2014). nates with trisomy 13 (5%).
In a sample of 50 legs, Mori (1964) found a sesamoid bone A high origin of the medial head of gastrocnemius was asso-
in the lateral head of gastrocnemius in 14 cases (28%). Mori ciated with an abnormal course and aneurysm of the popli-
(1964) also found a third head of gastrocnemius, extend- teal artery in a case described by Stuart (1879). Accessory
ing between the capsule of the knee joint and the medial or slips of the medial or lateral head of gastrocnemius may
lateral head in 2.8% of cases. There was muscular fusion lead to popliteal vascular entrapment syndrome (Takase
between gastrocnemius and soleus in 22 cases (44%) and et al. 1997; Liu et al. 2005; Rochier and Sumpio 2009;
tendinous fusion between the two muscles in 28 cases Sirasanagandla et al. 2013a). If the third head of gastrocne-
(56%). Soleus received a small bundle from the lateral head mius extends across the popliteal fossa, it may compress or
of gastrocnemius in 2.5% of cases (Mori 1964). entrap the neurovascular structures in this area and impact
Blitz and Eliot (2007) examined the gastrocnemius apo- nerve function and/or cause intermittent claudication,
neurosis in 53 cadaveric specimens. A long aponeurosis venous stasis, arterial stasis, aneurysms, or thromboembo-
was found in 28 specimens (53%), a short aponeurosis was lism (Hamming 1959; Iwai et al. 1987; Bergman et al. 1995;
found in five specimens (9%), and direct attachments were Koplas et al. 2009; Yildirim et al. 2011; Sirasanagandla
found between the deep surface of one of the gastrocnemius et al. 2013a; Lambert 2016). Extra heads of gastrocnemius
heads and soleus in 20 specimens (38%). In an MRI study may also cause sural nerve entrapment (Rodrigues et al.
of 1,039 knees, Koplas et al. (2009) observed a third head of 2016). Variation in the length of the gastrocnemius aponeu-
the gastrocnemius arising from the midline of the popliteal rosis and its attachments to soleus must be considered when
surface of the femur in 21 knees (2%). Of these, 20 cases planning gastrocnemius recession (Blitz and Eliot 2007).
(1.9%) joined the medial aspect of the lateral head and one
(0.1%) joined the medial head. Two accessory heads of gas-
trocnemius were present in one case (0.1%). Ashaolu et al. Soleus (Figure 5.6)
(2014) studied gastrocnemius in 60 legs from 30 Nigerian See also: Peroneotibialis
cadavers. Typical two-headed gastrocnemius muscles were
found in 21 legs (35%). Three-headed gastrocnemius mus- Synonyms
cles were found in eight legs (13.3%). Four-headed gastroc- N/A
nemius muscles were found in 31 legs (51.7%).
Description Soleus originates from the head and the proximal shaft of
Bersu et al. (1976) describe a male infant with Hanhart the fibula (fibular head) and from the soleal line and the
syndrome. The femora of this infant were normally devel- medial margin of the tibia (tibial head) (Standring 2016).
oped, but distal secondary ossification centers were absent. It also takes origin from a fibrous band connecting the two
The left leg stump had a patella and a small rudiment of bones, known as the tendinous arch of the soleus (Standring
the proximal tibia but no fibular rudiment. The right leg 2016). Soleus joins with the tendon of gastrocnemius to
stump was less developed and had a patella, smaller tibial form the calcaneal (Achilles) tendon, which inserts onto the
rudiment, and no fibular rudiment. There were no muscle calcaneus (Standring 2016).
masses suggestive of gastrocnemius at the knee.
In one neonate with trisomy 13, Pettersen et al. (1979) Innervation
found extra slips of gastrocnemius bilaterally. On the right Soleus is innervated by the tibial nerve (Standring 2016).
side, accessory slips originated from the medial and lateral
epicondyles of the femur and attached to the normal lateral Comparative Anatomy
head of gastrocnemius. On the left side, the extra slip was Soleus inserts into the calcaneus with gastrocnemius in all
associated with the lateral head. Mieden (1982) describes apes, but the tibial origin of this muscle is typically absent
two male fetuses with cyclopia and alobar holoprosenceph- in gibbons, orangutans, and gorillas (Chapman 1878;
aly. On the left side of one specimen, the medial head of Hepburn 1892; Beddard 1893; Raven 1950; Lewis 1962;
gastrocnemius was split into two heads. In one fetus with Gibbs 1999; Ferrero 2011; Ferrero et al. 2012; Diogo et al.
trisomy 18 and cyclopia, Smith et al. (2015) found that gas- 2010, 2012, 2013b). The tibial head is sometimes present
trocnemius and soleus on the right side of the body were in common chimpanzees and bonobos (Champneys 1872;
fused toward their insertion into the Achilles tendon, while Hepburn 1892; Frey 1913; Miller 1952; Lewis 1962; Gibbs
the two muscles on the left side were fused along the major- 1999; Diogo et al. 2013a, 2017). Accessory soleus muscles
ity of their length. have been found in orangutans (Diogo et al. 2013b).
Variations of the accessory soleus muscle was 14 out of 689 cases, or

Description around 2%. The prevalence was 2.4% in males and 2.1% in
females. One hundred and fifteen cases of accessory soleus
The tibial head of soleus may be absent (Mori 1964;
muscles were identified in the clinical literature, 64 of them
Bergman et al. 1988). The fibular head may be bilaminar
were male patients (55.6%), 28 were female patients (24%),
(Macalister 1875; Knott 1883b). The entire soleus or one of
and 23 were unidentified (20.4%). These authors also found
the heads may be absent or doubled (Macalister 1875; Le
that the most common insertion was onto the medial aspect
Doublet 1897; Bergman et al. 1988; Lambert 2016). Soleus
of the calcaneus via a separate tendon (26.1% of cases). A
may remain separate from gastrocnemius and its tendon up
tendinous insertion into the calcaneal tendon was found in
to its insertion (Macalister 1875; Knott 1883b; Bergman
3.5% of cases, and a fleshy attachment to the medial surface
et al. 1988; Lambert 2016). Soleus is also associated with
of the calcaneus was found in 4.3% of case. The remaining
several named accessory muscles. See the entry for pero-
66.1% of cases had unidentified attachment types. In their
neotibialis. Tensor fasciae plantaris or tibiotarsalis is a slip
own dissections of 100 legs from 50 cadavers, accessory
that originates from the soleal line below the attachment
soleus was found in three legs (3%). In their own imaging
of soleus and inserts onto the plantar fascia (Wood 1864;
study of 100 clinical patients, accessory soleus was found in
Macalister 1875; Anderson 1880; Knott 1883b; Bergman
three cases (3%) (Hatzantonis et al. 2011).
et al. 1988; Lambert 2016).
Soleus accessorius or accessory soleus is typically found
medially and distally in the ankle, often situated between Anomalies
the calcaneal tendon and the flexor tendons or distal tibia Description
(Percy and Telep 1984; Bergman et al. 1988; Sekiya et al. Accessory soleus muscles are sometimes found in individu-
1994; Brodie et al. 1997; Standring 2016). It may originate als with congenital clubfoot (talipes equinovarus) and may
from the soleal line, the proximal fibula, or from the deep cause resistance to correction of this deformity (Sodre et al.
surface of soleus (Gordon and Matheson 1973; Lorentzon 1994; Chotigavanichaya et al. 2000). In a male neonate with
and Wirrell 1987; Bergman et al. 1988; Sekiya et al. 1994; Meckel syndrome, Pettersen (1984) observed that soleus
Yu and Resnick 1994; Yildirim et al. 2011; Lambert 2016). was fleshy distal to the usual site of tendon formation at
It may insert into the calcaneal tendon, upper or medial sur- the ankle. In a fetus with trisomy 18 and cyclopia, Smith
faces of the calcaneus, or tibial collateral ligament of the et al. (2015) found that gastrocnemius and soleus on the
ankle (Gordon and Matheson 1973; Lorentzon and Wirrell right side of the body were fused toward their insertion into
1987; Bergman et al. 1988; Sekiya et al. 1994; Yu and the Achilles tendon, while the two muscles on the left side
Resnick 1994; Brodie et al. 1997; Hatzantonis et al. 2011; were fused along the majority of their length.
Yildirim et al. 2011; Lambert 2016; Standring 2016).
Tibiocalcaneus internus (Figure 5.6) originates from Prevalence
medial crest of the tibia and from the soleal line and inserts Sodre et al. (1994) noted that accessory soleus muscles
onto the medial aspect of the calcaneus (Testut 1892; Hecker were found in 6 out of 72 patients with congenital club foot
1924; Sammarco and Conti 1994; Sookur et al. 2008; (8.3%). Porter (1996) reports that an “anomalous calf flexor
Lambert 2016). It also has an origin from the deep fascia muscle” (with a description corresponding to that of soleus
of the leg (Sammarco and Conti 1994). While the accessory accessorius) was present in 7 out of 125 feet of children
soleus muscle is superficial to the flexor retinaculum, tibio- with congenital clubfoot (5.6%).
calcaneus internus passes deep to the flexor retinaculum and
superficial to neurovascular structures in the tarsal tunnel Clinical Implications
(Hecker 1924; Sookur et al. 2008; Lambert 2016).
Soleus accessorius becomes hypertrophic with physical
activity and therefore often presents in athletes or active indi-
Prevalence viduals as a soft mass in the medial ankle (Percy and Telep
In a study of 50 legs, Mori (1964) found that the tibial head 1984; Nelimarkka et al. 1988; Yu and Resnick 1994; Brodie
of soleus was absent in seven cases (14%). The muscle fibers et al. 1997; Lambert 2016). The presence of this muscle may
of soleus extended distal to the talocrural joint in two cases cause pain or swelling during exercise, potentially due to
(4%). There was muscular fusion between gastrocnemius restricted blood supply to the muscle (Gordon and Matheson
and soleus in 22 cases (44%) and tendinous fusion between 1973; Percy and Telep 1984; Nelimarkka et al. 1988; Brodie
the two muscles in 28 cases (56%). Soleus received a small et al. 1997; Lambert 2016). Symptomatic accessory soleus
bundle from the lateral head of gastrocnemius in 2.5% of muscles may simulate retrocalcanear bursitis or peritendi-
cases (Mori 1964). nitis of the Achilles tendon (Nelimarkka et al. 1988). Luck
Luck et al. (2008) found 35 cases of accessory soleus et al. (2008) report a high association between the presence
muscles from 4,771 MRI examinations of the ankle (0.73%) of accessory soleus muscles and Achilles tendinopathy.
over a 10-year duration from a sample of symptomatic Buschmann et al. (1991) found an association between acces-
adults. Hatzantonis et al. (2011) conducted a literature sory soleus muscles and pain and tenderness of the tibialis
review and determined that the overall cadaveric prevalence posterior tendon. Accessory soleus may cause compression
neuropathy of the posterior tibial nerve and tarsal tunnel syn- Typical Presentation
drome (DosRemedios and Jolly 2000; Kinoshita et al. 2003). Description
Plantaris originates from the lateral supracondylar line
Peroneotibialis (Figure 5.6) of the femur and from the oblique popliteal ligament
See also: Popliteus, Soleus (Standring 2016). Its tendon courses between gastrocne-
mius and soleus and inserts onto the calcaneus medial to
Synonyms the insertion of the Achilles tendon (Standring 2016).
N/A
Innervation
Typical Presentation Plantaris is innervated by the tibial nerve (Standring 2016).
Comparative Anatomy
Comparative Anatomy When present in the apes, plantaris originates runs from the
Peroneotibialis is present in some monkeys (e.g., howler lateral supracondylar line of the femur, lateral epicondyle,
monkeys, macaques). It lies deep to popliteus, originating lateral tibial condyle, or lateral head of gastrocnemius to
from the head or neck of the fibula and inserting onto the the calcaneus (Miller 1952; Vereecke et al. 2005; Ferrero
posterior aspect of the proximal tibial shaft (Howell and 2011; Ferrero et al. 2012; Diogo et al. 2012, 2013a,b, 2017).
Straus 1933; Grand 1968). Grand (1968) describes the mus- Plantaris is typically absent in gibbons, orangutans, and
cle as a vestigial rotator of the fibula on the tibia. gorillas (Gibbs 1999). It is often present in common chim-
panzees and bonobos (Gibbs 1999; Diogo et al. 2013a, 2017).
Variations
Peroneotibialis originates from the medial aspect of the Description
proximal fibula and inserts into the tendinous arch of the Plantaris may be absent (Knott 1883b; Bergman et al. 1988;
soleus or into the superior end of the soleal line of the tibia Daseler and Anson 1943; Mori 1964; Sato 1970; Moss 1988;
(Gruber 1878; Knott 1883a; Bergman et al. 1988; Lambert Saxena and Bareither 2000; Ashaolu et al. 2014; Lambert
2016; Standring 2016). Gruber (1878) refers to this muscle 2016; Standring 2016; Olewnik et al. 2018b). When absent,
as a tensor of the tendinous arch of the soleus (Bergman it may be replaced by a fibrous band (Bergman et al. 1988).
et al. 1988; Lambert 2016). Bergman et al. (1988) and Only the tendon may be absent, the muscle ending in an
Standring (2016) note that it runs deep to popliteus. In the aponeurosis or fascia between gastrocnemius or soleus
case photographed by Lambert (2016), the muscle has an (Macalister 1875; Sugavasi et al. 2013). Plantaris may
origin via a tendinous slip from popliteus. blend with the lateral head of gastrocnemius (Macalister
1875; Freeman et al. 2008; Lambert 2016). It may also
Innervation send a fibrous expansion to the patella (Freeman et al.
N/A 2008; Lambert 2016). The origin may be shifted more dis-
tally (Macalister 1875). Plantaris may have an origin from
Prevalence
the lateral retinaculum of the knee (Koplas et al. 2009).
Gruber (1878) found peroneotibialis in 128 out of 860 legs Plantaris may present as a third head of gastrocnemius and
(14.9%), and Knott (1883a) found the muscle in 4 out of 49 may blend with the two heads at their junction, merge with
subjects (8.2%). only one head, or join the fascia on the deep surface of gas-
trocnemius (Wood 1868; Bergman et al. 1988).
Anomalies
The tendon of plantaris may fuse with the calcaneal
N/A tendon upon insertion (Bergman et al. 1988; Nayak et al.
2010; Standring 2016; Olewnik et al. 2018b). It may also
join with the flexor retinaculum, tibial collateral ligament
Lambert (2016) notes that since peroneotibialis courses of the ankle, or the fascia of the leg (Bergman et al. 1988;
over the neurovascular bundle in the posterior lower leg, it Nayak et al. 2010; Standring 2016; Olewnik et al. 2018b).
may possibly lead to entrapment of the tibial nerve and the The tendon may have an attachment onto, or send a slip to,
posterior tibial artery. the plantar fascia (Macalister 1875; Bergman et al. 1988;
Lambert 2016).
Plantaris (Figure 5.6) An accessory head may be present, making plantaris
bicipital (Wood 1868; Macalister 1875; Bergman et al. 1988;
Synonyms Lambert 2016). Plantaris may be doubled, i.e., the accessory
Accessory plantaris may be referred to as plantaris minor head may be entirely separate from the main muscle (Knott
(Krause), popliteus superior s. minor (Calori) (Knott 1883b), 1883b; Standring 2016). Potential origins of the accessory
or bicipital plantaris (Hall) (Lambert 2016). head, or origins of a variable plantaris muscle, include the
normal origins of plantaris, the fascia of popliteus, the pop- of the knee joint and lateral head of gastrocnemius in three
liteal surface of the femur, lateral extension of the linea cases (5.77%); and from the lateral supracondylar line, cap-
aspera, lateral or medial femoral condyles, the intercondy- sule of the knee joint, lateral head of gastrocnemius, and
lar fossa, the posterior oblique ligament, the knee joint cap- fibular collateral ligament in seven cases (13.46%).
sule, the fascia of the leg, the lateral head of gastrocnemius, Three types of insertions were noted: into the flexor reti-
the soleal line of the tibia, or the proximal fibula (Wood naculum in 15 cases (28.85%), into the calcaneus in 19 cases
1868; Macalister 1875; Knott 1883b; Mori 1964; Bergman (36.54%), and into the calcaneal tendon in 14 cases (26.92%).
et al. 1988; Rana et al. 2006; Nayak et al. 2010; Kwinter et Using MRI, Herzog (2011) found accessory plantaris
al. 2010; Upasna and Kumar 2011; Lambert 2016). muscles in 63 of 1,000 (6.3%) symptomatic patients. In 62
Kotian et al. (2013) report a case where the plantaris cases, the normal plantaris and accessory plantaris had
muscle originates normally but bifurcates after about 1 cm merged origins. One accessory plantaris muscle originated
from its origin. The inferior belly passed posterior to the with the lateral head of the gastrocnemius muscle. The
popliteal neurovasculature and ended in a tendon that accessory plantaris muscles had insertions into the iliotibial
joined with the tendon of the superior belly near the origin tract in 5 out of the 63 cases (7.9%), the lateral patellar reti-
of soleus. The superior belly ran anterior to the popliteal naculum in 15 cases (23.8%), and the iliotibial band in 43
neurovasculature. The common tendon of the two bellies cases (68.3%) (Herzog 2011). Ashaolu et al. (2014) studied
inserted onto the calcaneus. Kalniev et al. (2014) report an gastrocnemius and plantaris in 60 legs from 30 Nigerian
accessory plantaris tendon that originated from soleus and cadavers. In the legs with a two-headed gastrocnemius (21
merged with the tendon of the main belly. The two inserted legs), there was 100% simultaneous occurrence of plantaris.
together onto the calcaneal tendon. In the legs with multi-headed gastrocnemius (39 legs), there
was 90% simultaneous occurrence of plantaris. Plantaris
Prevalence was thus present in 56 out of 60 legs (93.3%).
Knott (1883b) found plantaris absent in 3 out of 40 subjects Olewnik et al. (2018b) studied plantaris in 130 cadav-
(7.5%). Sato (1970) found that plantaris was absent in 43 eric lower limbs. Plantaris was absent in 14 limbs (10.8%).
out of 406 limbs (10.6%) from Kyushu Japanese males and In 51 out of the 116 cases (44%) in which the muscle was
in 36 out of 264 limbs (13.6%) from females. Moss (1988) present, plantaris had a wide insertion onto the calca-
found that plantaris was absent in 11 out of 150 cadavers neus medial to the Achilles tendon. In 26 cases (22.4%),
(7.3%), and in only four bodies (2.7%) the absence was the insertion onto the calcaneus blended with that of the
bilateral. Saxena and Bareither (2000) found that plantaris Achilles tendon. In eight cases (6.9%), the insertion was
was absent in 1 out of 40 cadaveric specimens (2.5%). In into the calcaneus anterior to the Achilles tendon. In four
an MRI study of 1,039 knees, Koplas et al. (2009) found cases (3.4%), the insertion was into the deep crural fas-
that plantaris originated from the lateral retinaculum in 19 cia. In 21 cases (18.1%), there was a wide insertion onto
knees (1.8%). the posterior and medial aspects of the Achilles tendon.
Mori (1964) found that plantaris was absent in 4% of In six cases (5.2%), there was an insertion into the flexor
cases. Mori (1964) also found that an accessory head arose retinaculum.
from the capsule of the knee joint in 16% of cases and from
the upper part of the tibia in 2.5% of cases. Daseler and Anomalies
Anson (1943) found that plantaris was absent in 50 out of Description
750 lower limbs (6.7%). Insertion of plantaris was examined Plantaris is often reduced or absent as an anomaly (Windle
in 150 cases. In 71 cases (47.3%), plantaris inserted into the 1893; Bersu et al. 1976; Pettersen et al. 1979; Aziz 1980;
medial aspect of the calcaneus medial to the calcaneal ten- Mieden 1982; Urban and Bersu 1897; Smith et al. 2015).
don. In 48 cases (32%), plantaris inserted into the calcaneus Bersu et al. (1976) describe a male infant with Hanhart
anterior to the calcaneal tendon. In 23 cases (15.3%), the syndrome. The femora of this infant were normally devel-
insertion joined blended with the insertion of the calca- oped, but distal secondary ossification centers were absent.
neal tendon. In seven cases (4.7%), plantaris inserted into The left leg stump had a patella and a small rudiment of
the medial border of the calcaneal tendon. Freeman et al. the proximal tibia but no fibular rudiment. The right leg
(2008) studied plantaris in 46 cadaveric knees. Plantaris stump was less developed and had a patella, smaller tibial
was absent in six knees (13.04%). In nine knees (19.57%), rudiment, and no fibular rudiment. There were no muscle
plantaris interdigitated with the lateral head of gastrocne- masses suggestive of plantaris at the knee.
mius. In five knees (10.87%), there was a strong fibrous Windle (1893) noted that plantaris was absent in some
extension of plantaris that went to the patella. anencephalic fetuses. Mieden (1982) describes an infant
Nayak et al. (2010) studied the origin and insertion with median cleft lip, hypotelorism, and alobar holopros-
of plantaris in 52 lower limbs from 26 cadaveric males. encephaly. Plantaris was absent on the right side (Mieden
Plantaris was completely absent in four limbs (7.69%). 1982). In one male neonate with trisomy 13, plantaris
Three types of origins were noted: an origin from the lat- lacked a tendon on the right side (Pettersen et al. (1979). In
eral supracondylar line, capsule of the knee joint, and lateral one female fetus, plantaris was hypoplastic on the left side
head of gastrocnemius in 38 cases (73.08%); from capsule (Pettersen et al. 1979). In a female neonate with trisomy 13,
plantaris was absent bilaterally (Aziz 1980). In one fetus Variations

with mosaic trisomy 18, found that plantaris was absent Description
bilaterally (Urban and Bersu 1987).
Popliteus may have connections to the posterior capsule of
Prevalence the knee joint, the oblique popliteal ligament, the posterior
cruciate ligament, and/or the posterior meniscofemoral liga-
Among the limbs of ten anencephalic fetuses, Windle
ment (Last 1950; Fetto et al. 1977; Feipel et al. 2003; Lambert
(1893) noted that plantaris was absent unilaterally in two
2016). The popliteal tendon near the lateral meniscus may
cases (20%). In their literature review, Smith et al. (2015)
be bifurcated (Burman 1968; Perez Carro et al. 1999; Leal-
found that plantaris was absent in 8 out of 20 individuals
Blanquet et al. 2009; Lambert 2016). Doral et al. (2006) report
with trisomy 13 (40%) and in 1 out of 17 individuals with
a popliteus tendon comprised of three bundles. Popliteus may
trisomy 18 (5.9%).
be absent or doubled (Macalister 1875; Knott 1883b; Le
Clinical Implications Double 1897; Lambert 2016).
An accessory slip or head of popliteus may originate
An extension of plantaris to the patella could have a role in
from the lateral femoral condyle, from the sesamoid bone
patellofemoral pain syndrome (Freeman et al. 2008). A split
in the lateral head of gastrocnemius, from the posterior
plantaris that surrounds the tibial nerve, popliteal vessels,
cruciate ligament, or from the posterior knee joint capsule
and nerve to soleus may lead to the entrapment of these
(Macalister 1875; Knott 1883a; Le Double 1897; Bergman
structures (Kotian et al. 2013). An insertion of the plantaris
et al. 1988; Lambert 2016; Standring 2016). It can insert
tendon into the flexor retinaculum may affect the tendinop-
with popliteus into the tibia and may be situated in front
athy of tibialis posterior and an insertion that blends with
of or behind the vessels in the popliteal fossa (Macalister
the Achilles tendon may increase the risk of Achilles tendi-
1875; Bergman et al. 1988; Lambert 2016). Accessory slips
nopathy (Olewnik et al. 2018b).
in this region may also extend to the posterior wall of the
capsule of the knee joint, the fibular head, or to the tibia near
Popliteus (Figures 5.6 and 5.7) the insertion of the posterior cruciate ligament (Bergman
et al. 1988). Using MRI, both Duc et al. (2004) and Hahn et
See also: Peroneotibialis
al. (2019) found accessory popliteus muscles with an origin
Synonyms from the lateral head of gastrocnemius and insertion into
An accessory popliteus may be referred to as popliteus the posteromedial articular capsule of the knee joint. An
biceps (Gruber), proximal popliteus, or popliteus geminus accessory slip of popliteus may present as popliteus minor,
(Fabrice) (Lambert 2016). a muscle situated medial to plantaris that extends from the
posterior surface of the lateral condyle of the tibia to the
Typical Presentation oblique popliteal ligament (Standring 2016). Knott (1883b)
describes popliteus minor as an accessory bundle that orig-
Description
inates from the lateral tendon of gastrocnemius.
Popliteus originates via a tendon from the lateral condyle of
the femur with supplementary fibers from the arcuate pop- Prevalence
liteal ligament and the lateral meniscus (Standring 2016). It
Gruber (1875b) found an accessory popliteus in 11 out of
inserts onto the posterior surface of the tibia just above the
695 cases (1.6%). In an MRI study of 1,039 knees, Koplas
soleal line (Standring 2016).
et al. (2009) found an accessory popliteal muscle in three
Innervation patients (0.3%). In a sample of 42 knees, Feipel et al. (2003)
found a fibular attachment of popliteus in 98% of cases,
Popliteus is innervated by the tibial nerve (Standring 2016).
at least one attachment to the lateral meniscus in 95% of
Comparative Anatomy cases, and the arcuate ligament in 90% of cases. Additional
attachments include the posterior knee joint capsule in 57%
Popliteus has a similar typical presentation in the apes,
of cases, the oblique popliteal ligament in 79% of cases, the
extending from the lateral condyle of the femur to the pos-
posterior cruciate ligament in 5% of cases, and the menis-
terior surface of the tibia (Champneys 1872; Hepburn 1892;
cofemoral ligament in 33% of cases. In a series of 1,569
Beddard 1893; Sonntag 1924; Raven 1950; Miller 1952;
knee arthroscopies, Leal-Blanquet et al. (2009) found six
Gibbs 1999; Ferrero 2011; Ferrero et al. 2012; Diogo et al.
bifurcated popliteus tendons (0.4%).
2010, 2012, 2013a,b, 2017). An origin from the fibular head
and adjacent knee joint capsule is present in orangutans and
Anomalies
bonobos (Hepburn 1892; Diogo et al. 2013b, 2017). An ori-
gin from the knee joint capsule is also found in gorillas and Description
common chimpanzees (Hepburn 1892; Raven 1950; Gibbs Popliteus was absent on the right side of the trisomy 18
1999). The muscle in orangutans also has an origin from the cyclopic fetus dissected by Smith et al. (2015). Bersu et al.
oblique popliteal ligament and the lateral meniscus (Ferrero (1976) describe a male infant with Hanhart syndrome. The
2011; Ferrero et al. 2012; Diogo et al. 2013b). femora of this infant were normally developed but distal
secondary ossification centers were absent. The left leg Flexor digitorum longus (Figure 5.7)
stump had a patella and a small rudiment of the proximal
tibia but no fibular rudiment. The right leg stump was less Synonyms
developed and had a patella, smaller tibial rudiment, and no N/A
fibular rudiment. There were no muscle masses suggestive
of popliteus at the knee. Typical Presentation
Description
Prevalence Flexor digitorum longus originates from the tibia, attach-
N/A ing from just below the soleal line to the distal end of the
tibia (Standring 2016). It sends four tendons to digits two
Clinical Implications through five that pass deep to the tendons of flexor digi-
Accessory attachments of popliteus may help to protect the torum brevis (Standring 2016). These tendons insert into
lateral meniscus and the tibiofemoral joint from degenera- the bases of the distal phalanges of digits two through five
tive disorders and cartilage alterations (Feipel et al. 2003). (Standring 2016). These tendons also receive contributions
A popliteus tendon comprised of multiple bundles may be from the quadratus plantae and the tendons of flexor hal-
mistaken for tendinosis (Doral et al. 2006). lucis longus (Standring 2016).
FIGURE 5.7 Flexor digitorum accessorius longus (a) and fibulocalcaneus internus (b) in medial view. Deep muscles of the posterior
lower leg in posterior view (c).
Innervation five from flexor digitorum brevis (Bergman et al. 1988). A

Flexor digitorum longus is innervated by the tibial nerve slip from flexor digitorum longus may pass to join the flexor
(Standring 2016). hallucis longus tendon to digit one (Bergman et al. 1988;
Lambert 2016; Lee and Hur 2017). An accessory head of
Comparative Anatomy flexor digitorum longus may be present and is referred to
as flexor accessorius longus or flexor digitorum accessorius
Flexor digitorum longus has a similar typical presentation
longus (see the entry for this muscle) (Standring 2016).
in the apes, extending from the posterior surface of the tibia
to the distal phalanges of the lateral digits, with some fusion Prevalence
with nearby muscles and frequent variation in which digits
In a study of 33 feet, Martin (1964) found no connection
receive tendons (Champneys 1872; Hepburn 1892; Beddard
between the tendons of flexor hallucis longus and flexor
1893; Sonntag 1924; Raven 1950; Miller 1952; Lewis 1962;
digitorum longus in two cases (6.1%) and two other cases
Gibbs 1999; Diogo et al. 2010, 2012, 2013a,b, 2017). In
(6.1%) where the tendons gave slips to each other. In a
gibbons, the tendons to digits two, four, and five are typi-
study of 16 cadaveric feet, O’Sullivan et al. (2005) identi-
cally present while the tendon to digit three is often absent
fied three patterns of connections between the tendons of
(Hepburn 1892; Beddard 1893; Lewis 1962; Diogo et al.
flexor digitorum longus and flexor hallucis longus. In 11
2012). In orangutans, digits two, four, and five are typically
feet (68.75%), tendinous fibers passed from the flexor hal-
supplied by the tendons (Gibbs 1999; Diogo et al. 2013b).
lucis longus tendon to the flexor digitorum longus tendon.
In gorillas, digits two and five are always supplied by the
In 2 feet (12.5%), tendinous fibers passed from the flexor
tendons, and digits three and four are supplied in half of
digitorum longus tendon to the flexor hallucis longus ten-
all cases (Gibbs 1999). In common chimpanzees and bono-
don. In 3 feet (18.75%), tendinous fibers passed from the
bos, digits two and five are almost always supplied by the
flexor digitorum longus tendon to the flexor hallucis longus
tendons while the tendons to digits three and four are typi-
tendon and an additional slip passed from the flexor hallucis
cally absent (Champneys 1872; Gibbs 1999; Hepburn 1892;
longus tendon to the flexor digitorum longus tendon.
Beddard 1893; Dwight 1895; MacDowell 1910; Miller 1952;
In a study of 24 cadaveric legs, LaRue and Anctil (2006)
Lewis 1962; Diogo et al. 2013a, 2017).
found three patterns of connections between the tendons
of flexor digitorum longus and flexor hallucis longus. In 10
Variations feet (41.7%), a tendinous slip passed from the flexor hal-
Description lucis longus tendon to the flexor digitorum longus tendon.
The lateral portion of flexor digitorum longus or the entire In ten other feet (41.7%), a tendinous slip passed from the
muscle may be absent (Bergman et al. 1988). Accessory flexor hallucis longus tendon to the flexor digitorum longus
slips from the fibula, tibia, tibialis anterior, tibialis pos- tendon, and another slip passed from the flexor digitorum
terior, or the crural fascia may join the muscle above the longus tendon to the flexor hallucis longus tendon. In 4 feet
ankle (Macalister 1875; Le Double 1897; Bergman et al. (16.7%), there was no attachment of the tendons.
1988; Lambert 2016). A slip may originate from the fascia Athavale et al. (2012a) studied the connections between
over flexor digitorum longus and merge with the tendon of the long flexor tendons in sample of 47 lower limbs. In 19
flexor hallucis longus in the foot (Bergman et al. 1988). limbs (40.4%), flexor hallucis longus sent a slip to the main
The slips that connect the tendon(s) of flexor digitorum tendon of flexor digitorum longus. In 21 limbs (44.7%), the
longus to the tendon of flexor hallucis longus frequently slip from flexor hallucis longus contributed to the digital
vary (Macalister 1875; Knott 1883b; Martin 1964; Bergman tendons of flexor digitorum longus, going to the tendon to
et al. 1988; Lambert 2016; Standring 2016). The variable digit two in two cases (4.3%), the tendons to digits two and
blending of the flexor digitorum longus and flexor hallucis three in 14 cases (29.8%), and the tendons to digits two,
longus is referred to as the chiasma plantare or the knot of three, and four in five cases (10.6%). In two cases (4.3%),
Henry (e.g., O’Sullivan et al. 2005; LaRue and Anctil 2006; a slip from flexor digitorum longus went to flexor hallucis
Plaass et al. 2013; Pretterklieber 2017; Beger et al. 2018b; longus. In five cases (10.6%), there were no connections
Elvan et al. 2019). This blending often involves quadratus between the tendons of these two muscles.
plantae (Pretterklieber 2017). There may also be no connec- Plaass et al. (2013) studied the connections between the
tions between the tendons of the long flexors (Knott 1883b; long flexor tendons in sample of 60 feet from 30 cadavers.
Martin 1964; LaRue and Anctil 2006). See prevalence sec- In 40 feet (67%) there was a proximal slip from flexor hal-
tion below for more information on the anatomical relation- lucis longus to the distal flexor digitorum longus only. Out of
ships of these tendons. these 40 cases, there was a connection to digit two in 18 feet
Some insertion tendons of flexor digitorum longus may (45%), a connection to digits two and three in 18 feet (45%),
be absent or otherwise vary (Macalister 1875; Knott 1883b; and a connection to digits two, three, and four in 4 feet (10%).
Bergman et al. 1988; Lambert 2016). The tendon to digit In 2 feet out of the 60 feet (3%), there was a proximal slip
two may be absent and replaced by a tendon from flexor passing from flexor digitorum longus to the distal flexor hal-
hallucis longus (Bergman et al. 1988; Lambert 2016). A slip lucis longus only. In 18 out of 60 feet (30%), there was a slip
from flexor digitorum longus may replace the tendon to digit from flexor hallucis longus to flexor digitorum longus and
additional slips between the proximal flexor digitorum lon- the tendon of flexor digitorum longus. Hootnick et al.
gus and distal flexor hallucis longus. Out of these 18 cases, (1987) describe an individual that had a right limb with con-
there was a connection to digit two in three cases (17%) and a genital tibial aplasia, talocalcaneal synchondrosis, and an
connection to digits two and three in 15 cases (83%). adducted foot with five toes. In this limb, flexor digitorum
Pretterklieber (2017) studied the chiasma plantare in longus originated from the interosseus membrane, became
100 feet from 50 cadavers and conducted a literature review tendinous halfway along its length, and inserted into the
of previous research. In 61 specimens (61%), the tendon of common flexor tendon sheet in the foot.
flexor hallucis longus sent slips to the tendon of flexor digi-
torum longus. In 39 specimens (39%), the tendons of both Prevalence
muscles exchanged slips with each other. In a review of 14 N/A
studies, including the data from Martin (1964), O’Sullivan
et al. (2005), LaRue and Anctil (2006), and Plaass et al. Clinical Implications
(2013), Pretterklieber (2017) found that out of a total of 754 Knowledge of the variation in tendinous connections
cases, slips from flexor hallucis longus to flexor digitorum between flexor digitorum longus and flexor hallucis longus
longus only were present in 564 cases (74.8%), slips from is important for planning tendon transfer to treat tibialis
flexor digitorum longus to flexor hallucis longus only were posterior tendon deficiency or dysfunction (O’Sullivan et al.
present in four cases (0.5%), slips exchanged between both 2005; Plaass et al. 2013).
tendons were found in 172 cases (22.8%), and no connec-
tions between the tendons occurred in 14 cases (1.9%). Flexor digitorum accessorius longus (Figure 5.7)
Pretterklieber (2017) also examined the contributions of
flexor digitorum longus, flexor hallucis longus, and quadra- Synonyms
tus plantae to the flexor tendons. Flexor digitorum longus This muscle may also be referred to as flexor accesso-
contributed to the first tendon in 39 cases (39%), to the sec- rius longus (Standring 2016), accessorius ad accessorium
ond tendon in 98 cases (98%), and to the third, fourth, and (Turner), accessorius ad flexor accessorium, accessorius
fifth tendons in all 100 cases (100%). Elvan et al. (2019) ad quadratum plantae, accessorius longus ad flexor digi-
studied the relationships between the tendons of flexor hal- torum longum, accessorius flexori hallucis longi superior
lucis longus and flexor digitorum longus in 56 feet from (Reinhart), fibulocalcaneus medialis, flexor accessorius
28 fetuses. In 48 feet (85.7%), there was a slip from flexor digiti longus, flexor accessorius longus digitorum pedis
hallucis longus to the chiasma plantare. In 8 feet (14.3%), (Hallett), and peroneocalcaneus internus (Macalister)
there were cross-connections between the tendons of flexor (Bergman et al. 1988).
hallucis longus and flexor digitorum longus.
Lee and Hur (2017) found a tendinous slip of flexor digito- Typical Presentation
rum longus going to digit one in 2 out of 66 cadaveric speci- This muscle is only present as a variation or anomaly.
mens (3%). In a study of 20 feet from 10 cadavers, Beger
et al. (2018b) found five patterns of connections between the Comparative Anatomy
tendons of flexor hallucis longus and flexor digitorum lon- The flexor digitorum accessorius longus muscle may rep-
gus. In 15 feet (75%), there was one slip from flexor hallucis resent a remnant of the higher, tibial sided (medial) origin
longus to flexor digitorum longus. In two feet (10%), each of quadratus plantae (flexor accessorius) present in some
tendon sent one slip to the other tendon. In one foot (5%), lower mammals (Lewis 1962; Jaijesh et al. 2005, 2006).
flexor hallucis longus sent two slips to flexor digitorum lon- This variant is not found in apes (Jaijesh et al. 2005, 2006).
gus. In one foot (5%), flexor hallucis longus sent two slips to
flexor digitorum longus and flexor digitorum longus sent one Variations
slip to flexor hallucis longus. In one foot (5%), flexor digito- Description
rum longus sent two slips to flexor hallucis longus and flexor Flexor digitorum accessorius longus can originate from
hallucis longus sent one slip to flexor digitorum longus. any structure or combination of structures in the posterior
compartment of the lower leg, with recorded origins includ-
Anomalies ing the tibia, fibula, both the tibia and fibula, calcaneus, the
Description deep fascia of the posterior compartment, posterior crural
In a female neonate with trisomy 13, Aziz (1980) found that intermuscular septum, flexor retinaculum, flexor digito-
flexor digitorum longus on the left side had a distally shifted rum longus, flexor hallucis longus, and/or tibialis posterior
origin. In a male fetus with triploidy studied by Moen et al. (Wood 1868; Hallett 1848; Macalister 1875; Driver and
(1984), flexor digitorum longus had an extra belly with a Denison 1914; Nathan et al. 1975; Bergman et al. 1988;
tendon attaching to the tendon of flexor hallucis longus on Sammarco and Conti 1994; Peterson et al. 1995; Cheung
the left side. On the right foot of a male neonate with Meckel et al. 1999; Gümüşalan and Kalaycioğlu 2000; Sookur et
syndrome, Pettersen (1984) found that flexor digiti minimi al. 2008; Bowers et al. 2009; Upasna et al. 2011; Standring
brevis attached to the fifth digit and received a slip from 2016; Lambert 2016). It can originate via two distinct heads,
flexor digitorum brevis that did not split to accommodate described as either a long and short head or medial and lateral
head (Driver and Denison 1914; Nathan et al. 1975; Pác and posterior compartment and six muscles (54.5%) arose from
Malinovský 1985; Gümüşalan and Kalaycioğlu 2000). the fibula. The five tendons that had origins from the tibia
Flexor digitorum accessorius longus passes through the inserted onto quadratus plantae with two of them having an
tarsal tunnel to enter the foot (Bergman et al. 1988; Cheung additional attachment to flexor digitorum longus. Three of
et al. 1999; Sookur et al. 2008; Duran-Stanton and Bui- the tendons with origins from the fibula inserted onto qua-
Mansfield 2010; Lambert 2016). It often has fleshy, muscular dratus plantae, with the remaining three tendons inserting
fibers as it passes through the tarsal tunnel (Nathan et al. 1975; onto flexor digitorum longus.
Cheung et al. 1999; Sookur et al. 2008; Bowers et al. 2009). In a sample of 100 ankle MRI examinations from asymp-
It may insert into the main tendon of flexor digitorum tomatic patients, Cheung et al. (1999) found flexor digitorum
longus and/or into quadratus plantae (Hallett 1848; Wood accessorius longus in six cases (6%). These authors combined
1864, 1868; Bergman et al. 1988; Sammarco and Conti data from these 6 cases and 14 other cases from symptomatic
1994; Peterson et al. 1995; Cheung et al. 1999; Gümüşalan patients to study flexor digitorum accessorius longus in a total
and Kalaycioğlu 2000; Sookur et al. 2008; Bowers et al. sample of 20 cases. Cheung et al. (1999) found that the muscle
2009; Georgiev et al. 2009; Upasna et al. 2011; Lambert arose from flexor hallucis longus in 8 out of 20 cases (40%),
2016; Standring 2016). It may also insert into the flexor ten- from the flexor retinaculum in 6 cases (30%), and the other 6
dons going to digit two (Georgiev et al. 2009; Upasna et al. cases (30%) had indeterminate origins.
2011), those going to digits two and three (Knott 1883b), or In a sample of 200 legs from 100 cadavers, Nathan et al.
those going to digits three and four (Driver and Denison (1975) found flexor digitorum accessorius longus in 14 legs
1914). Pác and Malinovský (1985) note that the tendon may from 12 cadavers (7%). A double origin of the muscle was
join the tendons of flexor digitorum longus to digits two, found in 8 out of the 14 cases (57.1%). Out of a sample of 60
three, and four, and may receive a tendinous slip from flexor cadavers, Gümüşalan and Kalaycioğlu (2000) found flexor
hallucis longus. Sookur et al. (2008) state that its tendon digitorum accessorius longus with two heads bilaterally in
splits into four slips and sends a tendon to the distal phalan- only one specimen (1.7%).
ges of digits two through five. In a sample of 44 feet from 37 patients who underwent
This accessory muscle may either be separate or attached operations to treat tarsal tunnel syndrome, Sammarco and
to flexor digitorum longus (Macalister 1875). Hallett (1848) Conti (1994) found flexor digitorum accessorius longus in 6
notes that quadratus plantae may be unusually large when feet (13.6%) from five patients. In a sample of 49 feet from
flexor digitorum accessorius longus is present. The pres- 41 patients who underwent operations to treat tarsal tunnel
ence of flexor digitorum accessorius longus may also be syndrome, Kinoshita et al. (2003) found flexor digitorum
associated with the absence of quadratus plantae (Pác and accessorius longus in 6 feet (12.2%) from six patients.
Malinovský 1985). It may send slips to flexor hallucis lon-
gus or tibialis anterior (Standring 2016). Flexor digitorum Anomalies
accessorius longus may present as flexor digiti secundi pro-
Description
prius, which originates from the posterior surface of the
tibia (Macalister 1875; Knott 1883b). Flexor digitorum accessorius longus muscles are sometimes
found in individuals born with clubfoot (talipes equin-
Innervation ovarus) (Sodre et al. 1994). The muscle was present in the
Flexor digitorum accessorius longus is innervated by the right club foot of an infant with Nager syndrome described
tibial nerve (Driver and Denison 1914; Upasna et al. 2011). by Kubota et al. (2001). It originated from the fascia over
flexor hallucis longus and the deep fascia of the lower leg
Prevalence (Kubota et al. 2001).
Wood (1868) reports that the muscle was found in 4 out Urban and Bersu (1987) found extra muscle slips extend-
of 68 male subjects (5.9%) and 1 out of 34 female subjects ing between the tibia and quadratus plantae on the right
(2.9%). Driver and Denison (1914) found four flexor digi- side of a fetus with full trisomy 18 and bilaterally in a fetus
torum accessorius longus muscles from two cadavers in a with mosaic trisomy 18. Bersu (1980) found a muscle that
sample of 50 limbs from 25 cadavers (8%). In a sample of extended between the fibula and quadratus plantae bilater-
36 legs from 18 cadavers, Lewis (1962) found the muscle ally in a child with trisomy 21.
in three legs from two cadavers (8.3%). In a sample of 52
legs from 26 cadavers, Canter and Siesel (1997) found the Prevalence
muscle in two legs (3.8%) from two cadavers. Upasna et al. Flexor digitorum accessorius longus muscles were found in 4
(2011) found this muscle in 1 out of 60 legs (1.7%). In a sam- out of 72 patients with club foot (5.6%) (Sodre et al. 1994). In
ple of 47 lower limbs, Athavale et al. (2012a) found flexor their literature review, Smith et al. (2015) found that the extra
digitorum accessorius longus in two cases (4.3%). muscle slips found by Urban and Bersu (1987) were only pres-
In a sample of 136 lower limbs from 68 cadavers, Peterson ent in the two fetuses examined by them and were not present
et al. (1995) found flexor digitorum accessorius longus in 11 in 15 other individuals with trisomy 18, yielding a prevalence
lower limbs (8.1%) from nine cadavers. Five out of the 11 of 11.8%. The muscle described by Bersu (1980) was found in
muscles (45.5%) arose from the tibia and deep fascia of the one out of five individuals with trisomy 21 (20%).
Clinical Implications Typical Presentation

Flexor digitorum accessorius longus can compress the This muscle is only present as a variation or anomaly.
tibial nerve or its branches and cause tarsal tunnel syn-
drome (Nathan et al. 1975; Sammarco and Stephens Comparative Anatomy
1990; Sammarco and Conti 1994; Canter and Siesel 1997; N/A
Kinoshita et al. 2003; Molloy et al. 2015). The presence
of this muscle may also cause flexor hallucis syndrome Variations
(Eberle et al. 2002). Description
Fibulocalcaneus internus typically originates from the
distal third of the fibula below the origin of flexor hallu-
Fibulocalcaneus internus (Figure 5.7) cis longus (Macalister 1875; Le Double 1897; Knott 1883a;
See also: Flexor digitorum accessorius longus Mellado et al. 1997; Sookur et al. 2008; Lambert et al 2011a;
Lambert 2016). It may also take origin from the posterior
Synonyms intermuscular septum and/or flexor hallucis longus (Perkins
This muscle may also be referred to as peroneocalcaneus 1914; Lambert et al. 2011a). Its origin may interdigitate with
internus (Macalister 1875) or pronator pedis (Knott 1883a). flexor hallucis longus (Mellado et al. 1997; Lambert et al.
Peroneocalcaneus medialis or fibulocalcaneus medialis may 2011a). The muscle becomes tendinous before entering the
be used as synonyms for this muscle. Although Lambert tarsal tunnel (Mellado et al. 1997; Howe and Murthy 2012).
et al. (2011a) note that the description of fibulocalcaneus The muscle courses lateral to flexor hallucis longus,
medialis by Jackson (1921) best matches descriptions of passes deep to the flexor retinaculum, and inserts via a
flexor digitorum accessorius longus, the lateral origins and tendon onto the inferomedial aspect of the calcaneus, typi-
point of insertion of the fibulocalcaneus medialis muscles cally into a small tubercle below the sustentaculum tali
described by Pettersen (1979) seem more aligned with (Macalister 1875; Le Double 1897; Perkins 1914; Mellado
descriptions of fibulocalcaneus internus. et al. 1997; Sookur et al. 2008; Lambert et al 2011a; Howe
Fibulocalcaneus internus is listed as a synonym of flexor and Murthy 2012; Lambert 2016). It may insert directly
digitorum accessorius longus by Bergman et al. (1988). into the sustentaculum tali (Knott 1883a; Best et al. 2005;
However, some authors note that this synonymy is errone- Seipel et al. 2005; Lambert 2016). Lambert et al. (2011a)
ous as the two muscles are often confused with one another observe that the tendon inserts distal to the calcaneal coro-
since they both pass deep to the flexor retinaculum (Mellado noid fossa.
et al. 1997; Lambert et al. 2011a; Lambert 2016). The two
muscles may be distinguished by some key anatomical Innervation
differences. First, Nathan et al. (1975) suggest that flexor Fibulocalcaneus internus is innervated by the branch of the
digitorum accessorius longus is the only muscle that retains tibial nerve that supplies the distal muscle belly of flexor
fleshy fibers within the tarsal tunnel, while only the ten- hallucis longus (Lambert et al. 2011a).
dons of other muscles pass through the flexor retinaculum.
Prevalence
Second, flexor digitorum accessorius longus has a variable
origin from the posterior compartment of the lower leg Mellado et al. (1997) found only one case of fibulocalca-
while fibulocalcaneus internus often originates from the neus internus in a sample of MR images from 100 asymp-
distal fibula (Lambert et al. 2011a; Lambert 2016). Third, tomatic individuals (1%).
while flexor digitorum accessorius longus typically inserts
Anomalies
onto quadratus plantae and/or the flexor digitorum longus
tendon, fibulocalcaneus internus inserts onto the medial Description
aspect of the calcaneus and is separated from quadratus Ramirez-Castro and Bersu (1978) report the bilateral pres-
plantae (Peterson et al. 1995; Mellado et al. 1997; Cheung ence of “peroneocalcaneus medialis” in three neonates
et al. 1999; Sookur et al. 2008; Lambert et al. 2011a; Lambert with trisomy 18. These muscles were each innervated by a
2016). Another way to distinguish the muscles is via their branch from the tibial nerve. In all three cases, the muscles
relationships with flexor hallucis longus; flexor digitorum originated from the posterior aspect of the distal third of the
accessorius longus is typically situated medial and posterior fibula and the corresponding interosseous membrane. The
to flexor hallucis longus within the tarsal tunnel while fibu- muscles traveled with the tendon of flexor hallucis longus
localcaneus internus typically lies lateral to flexor hallucis and inserted into the plantar aspect of the calcaneus. In the
longus (Lambert 2016; Lambert et al. 2011a). Lastly, flexor foot, the peroneocalcaneus medialis muscles sent tendons to
digitorum accessorius longus is closely associated with the nearby structures. In one case, the tendons merged with the
neurovascular structures in the tarsal tunnel while fibulo- medial head of quadratus plantae. In the other two cases,
calcaneus internus is typically separated from these struc- the tendons extended laterally from between the origins of
tures by flexor hallucis longus (Mellado et al. 1997; Sookur flexor digitorum brevis and quadratus plantae to insert onto
et al. 2008; Lambert et al. 2011a; Lambert 2016). the lateral margin of the calcaneus.
Pettersen (1979) reports the presence of “fibulocalcaneus Comparative Anatomy

medialis” on both sides of a child with trisomy 13q. On the Flexor hallucis longus has a similar typical presentation in
left side, the muscle originated in the lower third of the pos- most apes, extending from the interosseous membrane, pos-
terior lower leg from the fascia of flexor hallucis longus, the terior crural intermuscular septum, and/or the fibula to the
fascia of the peroneal muscles, and the lateral intermuscular base of the distal phalanx of digit one (Champneys 1872;
septum. Its tendon passed under the flexor retinaculum, and it Hepburn 1892; Beddard 1893; Sonntag 1924; Raven 1950;
inserted into the deep surface of flexor digitorum brevis near Miller 1952; Lewis 1962; Gibbs 1999; Diogo et al. 2010,
its origin. On the right side, the muscle had a similar insertion 2012, 2013a,b, 2017). In orangutans, the tendon inserts onto
but originated from the superficial aspects of fibularis longus digits three and four only, with no insertion onto digit one,
and brevis. Pettersen et al. (1979) also found “fibulocalcaneus and there may be an origin from the lateral femoral condyle
medialis” bilaterally in a fetus with trisomy 13. The muscle (Chapman 1880; Hepburn 1892; Beddard 1893; Sonntag
originated from the back of the fibula just above the calca- 1924; Gibbs 1999; Diogo et al. 2013b).
neus. The muscle ran parallel and posterior to flexor hallucis In gibbons and gorillas, there are often additional inser-
longus. It inserted via a tendon onto the medial surface of the tions onto digits three and four, and sometimes insertions
calcaneus anterior to the tendocalcaneus. onto digits two and five (Chapman 1878; Hepburn 1892;
Hootnick et al. (1987) describe an individual that had a right Beddard 1893; Raven 1950; Lewis 1962; Gibbs 1999;
limb with congenital tibial aplasia, talocalcaneal synchondro- Diogo et al. 2010, 2012). In common chimpanzees and
sis, and an adducted foot with five toes. In this limb, an acces- bonobos, there are often additional insertions onto digits
sory muscle originated from the distal fibula and interosseous three and four, and sometimes an insertion onto digit two
membrane and inserted onto the superior surface of the calca- (Champneys 1872; Beddard 1893; Miller 1952; Diogo et al.
neus, medial to the insertion of the calcaneal tendon. 2013a, 2017).
In their literature review, Smith et al. (2015) found that Description
peroneocalcaneus medialis was only present in 3 out of 17
Flexor hallucis longus may be doubled, with both tendons
individuals with trisomy 18 (17.6%) and 2 out of 20 indi-
ending on the distal phalanx of digit one (Bergman et al.
viduals with trisomy 13 (10%).
1988). A slip may originate from the fascia over flexor
Clinical Implications digitorum longus and merge with the tendon of flexor hal-
lucis longus in the foot (Bergman et al. 1988). The slips
If fibulocalcaneus internus shifts flexor hallucis longus
that connect the tendon(s) of flexor digitorum longus to the
medially, it may indirectly lead to compression or entrap-
tendon of flexor hallucis longus frequently vary (Macalister
ment of the neurovascular bundle via the displaced flexor
1875; Knott 1883b; Martin 1964; Bergman et al. 1988;
hallucis longus (Mellado et al. 1997). The presence of
Lambert 2016; Standring 2016). The variable blending of
fibulocalcaneus internus has been associated with pain,
the flexor digitorum longus and flexor hallucis longus is
posterior ankle impingement, and flexor hallucis longus
referred to as the chiasma plantare or the knot of Henry
tenosynovitis (Best et al. 2005; Seipet et al. 2005). The
(e.g., O’Sullivan et al. 2005; LaRue and Anctil 2006; Plaass
presence of this muscle may also contribute to tarsal tunnel
et al. 2013; Pretterklieber 2017; Beger et al. 2018b; Elvan
syndrome (Duran-Stanton and Bui-Mansfield 2010).
et al. 2019). This blending often involves quadratus plan-
tae (Pretterklieber 2017). There may also be no connec-
Flexor hallucis longus (Figure 5.7) tions between the tendons of the long flexors (Knott 1883b;
Martin 1964; LaRue and Anctil 2006).
Synonyms When the tendon of flexor hallucis longus provides
N/A slips to the other toes (which occurs in most individuals),
it frequently supplies the second and third toes and some-
times extends to the fourth toe (Knott 1883b; Mori 1964;
Description Bergman et al. 1988; Standring 2016). It rarely contributes
Flexor hallucis longus originates from the fibula, the inter- to the tendon going to digit five (Mori 1964; Bergman et al.
osseous membrane, the posterior crural intermuscular sep- 1988; Pretterklieber 2017). Flexor hallucis longus is associ-
tum, and from the tibialis posterior fascia (Standring 2016). ated with the supernumerary muscle fibulotarsalis, which
Its tendon enters the foot, sends slips to the tendon of flexor extends from the fibula to the plantar aspects of the calca-
digitorum longus, and inserts onto the base of the distal neus and navicular (Bergman et al. 1988).
phalanx of digit one (Standring 2016).
Prevalence
Innervation Mori (1964) found that flexor hallucis longus sent slips to
Flexor hallucis longus is innervated by the tibial nerve digits one and two in 8% of cases; to digits one, two, and
(Standring 2016). three in 64% of cases; to digits one through four in 26%
of cases; and to all five digits in 2% of cases. Athavale et on the right side of the mosaic trisomy 18 fetus. Hootnick
al. (2012a) found that slips of flexor hallucis longus gave et al. (1987) describe an individual that had a right limb
rise to the first lumbrical muscle in 18 cases (38.3%) and to with congenital tibial aplasia, talocalcaneal synchondrosis,
other lumbrical muscles in nine cases (19.15%). Beger et al. and an adducted foot with five toes. In this limb, flexor hal-
(2018b) classified the contributions of flexor hallucis longus lucis longus was narrow and cord like and inserted onto
to the flexor tendons of the toes. In 5 feet (25%), flexor hal- the calcaneus.
lucis longus sent a slip to the second digit. In 12 feet (60%),
flexor hallucis longus sent slips to digits two and three. In Prevalence
3 feet (15%), flexor hallucis longus sent slips to digits two, In their literature review, Smith et al. (2015) found that
three, and four. the anomalies described by Urban and Bersu (1987) were
Pretterklieber (2017) found that the tendon of flexor hal- only present in the two specimens described by these
lucis longus sent a slip to digit two in 45 cases (45%), to authors out of a sample of 17 individuals with trisomy
digits two and three in 46 cases (46%), and to the second 18 (11.8%).
through fourth digits in 5 cases (5%). Four cases showed
other insertions, including slips to the second and fourth Clinical Implications
digits, slips to the third and fourth digits, a slip to only Knowledge of the variation in tendinous connections
digit three, and a slip to only digit four. In a review of between flexor digitorum longus and flexor hallucis longus
nine studies, including the data from Plaass et al. (2013), is important for planning tendon transfer to treat tibialis
Pretterklieber (2017) reports that out of 637 total cases, posterior tendon deficiency or dysfunction (O’Sullivan et
flexor hallucis longus sent a slip to digit two only in 223 al. 2005; Plaass et al. 2013).
cases (35%), digits two and three in 334 cases (52.4%), dig-
its two through four in 75 cases (11.8%), digits two through
Tibialis posterior (Figure 5.7)
five in one case (0.16%), and other patterns of insertion in
four cases (0.6%). Pretterklieber (2017) also examined the Synonyms
contributions of flexor digitorum longus, flexor hallucis This muscle may also be referred to as tibialis posticus
longus, and quadratus plantae to the flexor tendons. Flexor (Macalister 1875; Knott 1883b).
hallucis longus contributed to the first tendon in 100 cases
(100%), to the second tendon in 97 cases (97%), to the third Typical Presentation
tendon in 53 cases (53%), to the fourth tendon in eight cases Description
(8%), and never to the fifth tendon. Tibialis posterior originates via medial and lateral parts
Elvan et al. (2019) studied the relationships between (Standring 2016). The medial part originates from the
the tendons of flexor hallucis longus and flexor digito- interosseous membrane and from the posterior tibial sur-
rum longus in 56 feet from 28 fetuses. In 48 feet (85.7%), face (Standring 2016). The lateral part originates from the
there was a slip from flexor hallucis longus to the chiasma posterior fibular surface (Standring 2016). When it reaches
plantare. In 8 feet (14.3%), there were cross connections the foot, the tendon of tibialis posterior typically divides
between the tendons of flexor hallucis longus and flexor into two portions (Standring 2016). The superficial portion
digitorum longus. See the entry for flexor digitorum lon- has insertions onto the navicular, medial cuneiform, and
gus for more prevalence information regarding the vari- the sustentaculum tali (Standring 2016). The deeper por-
able the anatomical relationships between the tendons of tion has insertions onto the intermediate cuneiform and the
these muscles. bases of metatarsals two through four (Standring 2016).
Flexor hallucis brevis partially originates from this portion
Anomalies
of the tendon (Standring 2016).
Description
Mieden (1982) describes two male fetuses with cyclopia Innervation
and alobar holoprosencephaly. On the right side of one Tibialis posterior is innervated by the tibial nerve (Standring
fetus, the tendon of flexor hallucis longus was divided, one 2016).
tendon having the typical attachment and the other attach-
ing to the tendon of flexor digitorum brevis going to digit Comparative Anatomy
two. In a male fetus with triploidy studied by Moen et al. Tibialis posterior has a similar typical presentation in the
(1984), flexor hallucis longus sent tendons to all phalanges apes, originating from the tibia, fibula, and interosseous
on the left side. In one fetus with full trisomy 18, Urban membrane and typically inserting into the navicular and
and Bersu (1987) found that flexor hallucis longus sent a metatarsals two through four, with occasional attachments
tendon to the navicular. In another fetus with mosaic tri- to the cuneiforms and tendon of fibularis longus (Champneys
somy 18, flexor hallucis longus sent a tendon to the cal- 1872; Hepburn 1892; Beddard 1893; Sonntag 1924; Raven
caneus. Slips from this muscle to flexor digitorum longus 1950; Miller 1952; Lewis 1964; Gibbs 1999; Vereecke et al.
were present bilaterally in the full trisomy 18 fetus and 2005; Diogo et al. 2010, 2012, 2013a,b, 2017).
Variations Bloome et al. (2003) studied the insertions of the tibialis

Description posterior tendon in 11 feet from 10 cadavers. Each tendon
had three distinct bands and their insertions included the
Tibialis posterior may be absent (Macalister 1875; Le
base of the fifth metatarsal in 7 out of 11 cases (63.6%),
Double 1897; Lambert 2016). The tendon may be split
the spring ligament in four cases (36.4%), fibularis longus
into three distinct bands (Macalister 1875; Lewis 1964;
tendon in four cases (36.4%), and flexor hallucis brevis in
Bloome et al. 2003; Lambert 2016). The number of slips
nine cases (81.2%). Slips to abductor hallucis were found in
that arise from the tendon varies (Mori 1964; Bloome
five cases (45.5%).
et al. 2003; Standring 2016). Its tendon may send slips to the
tendon of fibularis longus, tendon of flexor digitorum bre- Anomalies
vis, flexor hallucis brevis, abductor hallucis, adductor hallu-
cis, the plantar aponeurosis, the spring ligament, capsule of Description
the naviculocuneiform joint, cuboid, lateral cuneiform, or Sodre et al. (1994) note that agenesis of tibialis posterior
base of the fifth metatarsal (Macalister 1875; Knott 1883b; may be found in individuals born with clubfoot (talipes
Lewis 1964; Mori 1964; Gunal et al. 1994; Lohrmann et al. equinovarus). Anomalous insertions of the tibialis pos-
1997; Bloome et al. 2003; Raheja et al. 2005; Lambert 2016; terior tendon are found in cases of congenital metatarsus
Standring 2016). The attachment onto the medial cuneiform varus (Browne and Paton 1979). Hootnick et al. (1987)
may be missing (Bergman et al. 1988). describe an individual that had a right limb with con-
Tibialis posterior is associated with three rare super- genital tibial aplasia, talocalcaneal synchondrosis, and an
numerary muscles. Tensor capsuli tibiotarsalis anterior is adducted foot with five toes. In this limb, tibialis posterior
considered a variant of tibialis secundus and inserts onto arose with flexor hallucis longus and inserted by a short
the anterior wall of the ankle joint capsule (Bergman et al. tendon onto the common flexor tendon sheet in the foot.
1988). Macalister (1875) notes that it originates from the
Prevalence
lower half of the lateral surface of the tibia below flexor
digitorum longus. Tibialis secundus (Bahnsen) or acces- Agenesis of tibialis posterior was found in 1 out of 72 patients
sory tibialis posterior extends from the posterior surface born with clubfoot (1.4%) (Sodre et al. 1994). Browne and
of the tibia, below tibialis posterior, to the ankle joint cap- Paton (1979) operated on 15 feet from ten children to treat
sule or the flexor retinaculum (Bahnsen 1868; Macalister congenital metatarsus varus and found anomalous inser-
1875; Knott 1883b; Le Double 1897; Bergman et al. 1988; tions of the tibialis posterior tendon in 14 feet (93.3%).
Lambert 2016; Standring 2016). Accessory tibialis muscles
may also have insertions into the tarsals, second metatarsal,
and tendons in the foot (Le Double 1897; Lambert 2016). Variations in the insertion of the tibialis posterior tendon
Tibiotarsalis or tensor fasciae plantaris (also described may lead to hallux valgus (Bozant et al. 1994; Gunal et al.
under the entry for soleus) extends from the soleal line 1994). Accessory or duplicated tibialis posterior tendons
of the tibia to insert into the plantar fascia (Wood 1864; can cause posterior tibial tenosynovitis (Ghormley and
Macalister 1875; Anderson 1880; Knott 1883b; Bergman et Spear 1953).
al. 1988; Lambert 2016).
ANTERIOR AND LATERAL MUSCLES
Prevalence
OF THE LOWER LEG
Mori (1964) found ten different patterns of insertion of the
tibialis posterior tendon. In 11% of cases, the tendon inserted Tibialis anterior (Figure 5.8)
onto the navicular, medial cuneiform, and intermediate cune-
iform. In 31% of cases, the tendon inserted onto the navicu- Synonyms
lar, intermediate cuneiform, and lateral cuneiform. In 1% of This muscle may be referred to as tibialis anticus (Macalister
cases, the tendon inserted onto the navicular, medial cunei- 1875; Knott 1883b).
form, and first metatarsal. In 2% of cases, the tendon inserted
onto the navicular, medial cuneiform, and second metatarsal. Typical Presentation
In 6% of cases, the tendon inserted onto the navicular, medial Description
cuneiform, and third metatarsal. In 6% of cases, the tendon Tibialis anterior originates from the tibia via the lateral
inserted onto the navicular, medial cuneiform, and fourth condyle and proximal shaft and from the interosseous
metatarsal. In 14% of cases, the tendon inserted onto the membrane (Standring 2016). It inserts via a tendon onto
navicular, medial cuneiform, and cuboid. In 10% of cases, the medial cuneiform and base of the hallucal metatarsal
the tendon inserted onto the navicular, medial cuneiform, and (Standring 2016).
fascia of flexor hallucis brevis. In 10% of cases, the tendon
inserted onto the navicular, medial cuneiform, and tendon Innervation
of fibularis longus. In 5% of cases, the tendon inserted onto Tibialis anterior is innervated by the deep fibular nerve
the navicular, medial cuneiform, and the plantar aponeurosis. (Standring 2016).
FIGURE 5.8 Anterior muscles of the lower leg in anterior view.
Comparative Anatomy muscle, abductor hallucis longus (Gibbs 1999; Diogo et al.
Tibialis anterior has a similar typical presentation in the 2013a, 2017). There may be an origin from extensor digi-
apes, extending from the tibia to the medial cuneiform and torum longus in orangutans (Sonntag 1924; Gibbs 1999;
first metatarsal (Hepburn 1892; Beddard 1893; MacDowell Diogo et al. 2013b).
1910; Sonntag 1924; Raven 1950; Miller 1952; Lewis 1966;
Gibbs 1999; Vereecke et al. 2005; Ferrero 2011; Ferrero Variations
et al. 2012; Diogo et al. 2010, 2012, 2013a,b, 2017). The Description
muscle is divided into two bellies in gibbons, common Additional fibers of origin may arise from the fibula (Knott
chimpanzees, and bonobos (Hepburn 1892; Beddard 1893; 1883b). The origin may also be shifted proximally to the
MacDowell 1910; Miller 1952; Lewis 1966; Diogo et al. lateral condyle of the femur (Le Double 1897; Lambert
2017). The insertion onto the first metatarsal in common 2016). The insertion onto the hallucal metatarsal, or more
chimpanzees and bonobos may be considered a separate infrequently the insertion onto the medial cuneiform, may
be absent (Hallisy 1930; Arthornthurasook and Gaew-Im and base of the proximal phalanx of digit one. In seven cases
1990; Brenner 2002; Lambert 2016; Olewnik et al. 2019a). (2.4%), the tendon had an extra slip that inserted onto the
Tibialis anterior may have distal attachments to the head of medial side of the base of the proximal phalanx of digit one.
the first metatarsal, base of the proximal phalanx of digit In two cases (0.7%), a small muscular belly was present that
one, talus, navicular, calcaneus, extensor retinaculum, apo- sent a tendon to the base of the proximal phalanx of digit one.
neuroses of the foot, or the ankle joint capsule (Macalister Mori (1964) found that in 78% of cases, an undivided
1875; Knott 1883b; Hallisy 1930; Mori 1964; Bergman et al. tendon of tibialis anterior coursed over the anterior surface
1988; Brenner 2002; Lambert 2016; Standring 2016). Ikiz of the tibia to the medial aspect of the foot and inserted
and Üçerler (2005) note an insertion of the tibialis anterior into the medial surface of the medial cuneiform and into
tendon onto the lateral side of the foot, with attachments the base of the first metatarsal. In 20% of cases, the tendon
onto the fifth metatarsal and cuboid. divided into two slips, one inserting onto the medial cunei-
The entire muscle, or more often just the tendon, may form and the other into the first metatarsal. In 2% of cases,
be divided (Macalister 1875; Hallisy 1930; Bergman et al. the slip was divided into three portions and inserted onto
1988; Lambert 2016). When the tendon of tibialis anterior is the medial cuneiform, first metatarsal, and navicular.
divided into two slips, one slip goes to the medial cuneiform Arthornthurasook and Gaew-Im (1990) studied the inser-
and the other goes to the first metatarsal (Macalister 1875; tion of tibialis anterior in 44 feet from 22 cadavers. In 25
Mori 1964; Bergman et al. 1988). It may also be divided cases (56.8%), the tendon divided into two equal slips, one
into three slips (Mori 1964; Olewnik et al. 2019a). going to the medial cuneiform and the other to the base of
A deep slip of muscle that extends from the distal tibia the first metatarsal. In 12 cases (27.3%), the tendon sent a
and interosseous membrane to the neck of the talus may large slip to the medial cuneiform and a small slip to the
be referred to as tibioastragalus anticus (Gruber) or ante- base of the first metatarsal. In seven cases (15.9%), the ten-
rior tibiotalus (Bergman et al. 1988; Berkowitz et al. 2016; don inserted onto the medial cuneiform only. Brenner (2002)
Lambert 2016). A separate slip extending from the distal examined the insertion of tibialis anterior in 156 feet. In two
tibia to the annular ligament and/or dorsal fascia of the cases (1.3%), there was an insertion only onto the medial
foot may be referred to as tibiofascialis anticus (Wood) cuneiform, and in two other cases (1.3%), there was an inser-
or tibialis anticus accessorius (s. profundus) (Macalister tion only onto the first metatarsal. In one case (0.6%), there
1875; Knott 1883b; Bergman et al. 1988; Lambert 2016). was an insertion onto the navicular and medial cuneiform.
It may present as a tendinous slip with an origin from tibi- Olewnik et al. (2019a) studied the insertion of tibialis
alis anterior instead of presenting as a separate structure anterior in 100 cadaveric lower limbs. In 31 cases (31%), the
(Macalister 1875; Knott 1883b; Bergman et al. 1988; Jain et tendon divided into two equal slips, one going to the medial
al. 2013; Lambert 2016). Although Knott (1883b) lists ten- cuneiform and the other to the base of the first metatarsal
sor fasciae dorsalis pedis (Krause) as a synonym of tibio- (Type I). In 24 cases (24%), the tendon sent a large slip to
fascialis anticus, Lambert (2016) describes it as a separate the medial cuneiform and a small slip to the base of the
slip that has the same insertion onto the inferior extensor first metatarsal (Type II). In 11 cases (11%), the tendon sent
retinaculum and dorsal fascia of the foot but with an origin a small slip to the medial cuneiform and a large slip to the
from the distal fibula. base of the first metatarsal (Type III). In two cases (2%), the
tendon trifurcated, sending one slip to the medial cuneiform
Prevalence and two slips to the first metatarsal (Type IV). In 32 cases
Macalister (1875) notes that Wood found tibiofascialis anti- (32%), the tendon inserted onto the medial cuneiform only
cus in 2 out of 36 cases (5.6%). Knott (1883b) found tibio- (Type V). Olewnik et al. (2019a) also studied the insertion
fascialis anticus in 2 out of 40 cases (5%). Out of 30 cases of tibialis anterior in 100 limbs from 50 volunteers using
(Knott 1883b), two (6.7%) sent a slip to the extensor retinac- ultrasound. Type I insertion was found in 20 limbs (20%),
ulum (“ligamentum cruciatum”), two (6.7%) sent a slip to Type II was found in 35 limbs (35%), Type III was found in
the head of the first metatarsal, and one (3.3%) sent a slip to 13 limbs (13%), Type IV in no limbs (0%), and Type V in
the base of the proximal phalanx of digit one. Knott (1883b) 20 limbs (20%). A sixth type was found in the remaining
notes that Kraus found a slip to the ligamentum cruciatum 12 limbs (12%), which demonstrated two slips of the tendon
in 6% of cases. that both inserted onto the medial cuneiform.
Hallisy (1930) studied tibialis anterior in 290 feet. In 261
feet (90%), the muscle had the typical insertions onto the
first metatarsal and medial cuneiform. In three cases (1%), Anomalies
the tendon split just prior to insertion. In two cases (0.7%), Description
the tendon inserted onto the first metatarsal only. In one case Macalister (1875) notes that in a case of clubfoot, tibialis
(0.3%), the tendon inserted onto the navicular, medial cunei- anterior had a split tendon. One part of the tendon attached
form, and first metatarsal. In three cases (1%), the tendon to the navicular and medial cuneiform and the other part
inserted normally and sent a slip to the shaft of the first meta- attached to the talus and calcaneus. Macalister (1875) also
tarsal just behind the head. In one case (0.3%), the tendon had notes that in a congenitally deformed limb described by
a slip with attachments into the head of the first metatarsal Ringhoffer, tibialis anterior inserted onto the plantar fascia.
Hootnick et al. (1987) describe an individual that had a 2008). The tendon may also have insertions onto the hallu-
right limb with congenital tibial aplasia, talocalcaneal cal metatarsal, metatarsophalangeal joint capsule, or both
synchondrosis, and an adducted foot with five toes. In this phalanges of digit one (Mori 1964; Hallisy 1930; Bergman
limb, tibialis anterior inserted onto a flexor tendon sheet on et al. 1988; Al-Saggaf 2003; Hill and Gerges 2008; Arora et
the plantar aspect of the foot. Tibialis anterior was absent al. 2011; Lambert 2016). Extensor hallucis longus may send
bilaterally in one neonate with trisomy 13 examined by a slip to digit two or join with the common extensor tendons
Colacino and Pettersen (1978). of the toes (Hallisy 1930; Bergman et al. 1988; Lambert
2016; Standring 2016). Extensor hallucis longus may be
Prevalence doubled or tripled, presenting with multiple bellies with
According to their literature review, Smith et al. (2015) separate tendons (Macalister 1875; Hallisy 1930; Bergman
found that tibialis anterior was absent in 1 out of 20 indi- et al. 1988; Lambert 2016).
viduals (5%) with trisomy 13. Extensor hallucis longus is also associated with mul-
tiple named accessory bellies or tendons. One of these
Clinical Implications accessory muscles is referred to as extensor ossis metatarsi
Tibioastragalus anticus may be useful for tendon transfer or hallucis or extensor ossis primi metatarsi (Macalister 1875;
a graft (Berkowitz et al. 2016). Lambert 2016). It may present as a slip of extensor hal-
lucis longus or as a distinct accessory muscle (Macalister
Extensor hallucis longus (Figure 5.8) 1875; Hallisy 1930; Lambert 2016). It may also present as
a slip from tibialis anterior or extensor digitorum longus
Synonyms (Macalister 1875; Lambert 2016). This muscle usually
This muscle may also be referred to as extensor proprius inserts onto the distal part of the first metatarsal (Hallisy
hallucis (Macalister 1875; Knott 1883b). 1930; Lambert 2016). It may also present only as a ten-
don originating from the annular ligament with no belly
Typical Presentation (Macalister 1875).
Description Another associated muscle is referred to as extensor
primi internodii hallucis (Wood) or extensor hallucis lon-
Extensor hallucis longus originates from the midportion
gus accessorius (s. minor) (Macalister 1875; Knott 1883b;
of the medial fibula and from the interosseous membrane
Lambert 2016). It usually presents as a slip originating
(Standring 2016). The muscle ends in a tendon that inserts
from extensor hallucis longus (Macalister 1875; Knott
onto the base of the distal hallucal phalanx (Standring
1883b; Hallisy 1930; Lambert 2016). It can have a sepa-
2016). The tendon often expands to insert onto the base of
rate origin above extensor hallucis longus with an inser-
the proximal hallucal phalanx (Standring 2016).
tion joining extensor hallucis brevis (Macalister 1875). It
Innervation may originate in the ankle from the tendon of extensor
hallucis longus (Macalister 1875; Knott 1883b). It may
Extensor hallucis longus is innervated by the deep fibular
also arise from extensor digitorum longus or tibialis ante-
rior (Macalister 1875; Knott 1883b; Hallisy 1930; Lambert
2016). Hallisy (1930) notes an origin from the tibia.
Comparative Anatomy
Extensor primi internodii hallucis typically inserts into
Extensor hallucis longus has a similar typical presenta- the proximal hallucal phalanx (Macalister 1875; Hallisy
tion in the apes, extending from the fibula and sometimes 1930; Lambert 2016).
the lateral tibial condyle or interosseous membrane to the Lambert (2016) describes extensor hallucis capsularis
distal hallucal phalanx (Hepburn 1892; Beddard 1893; as a synonym or variant of extensor primi internodii hal-
Sonntag 1924; Raven 1950; Miller 1952; Lewis 1966; Gibbs lucis. This muscle presents as a slip or accessory tendon
1999; Ferrero 2011; Ferrero et al. 2012; Diogo et al. 2010, of extensor hallucis longus that inserts into the capsule of
2012, 2013a,b, 2017). There may be an insertion onto the the first metatarsophalangeal joint (Tate and Pachnik 1976;
proximal hallucal phalanx in bonobos and gorillas (Raven Lundeen et al. 1983; Bibbo et al. 2004; Boyd et al. 2006;
1950; Miller 1952; Gibbs 1999; Vereecke et al. 2005; Diogo Bayer et al. 2014; Lambert 2016; Natsis et al. 2017). It may
et al. 2017). also originate from tibialis anterior (Bibbo et al. 2004;
Boyd et al. 2006).
Variations
Accessory extensor digiti secundus may refer to an
Description accessory slip that is situated between extensor hallucis lon-
Extensor hallucis longus may be joined with or send a gus and extensor digitorum longus (Tezer and Cicekcibasi
slip to extensor digitorum longus or its tendon (Macalister 2012). It may originate from the tendon of extensor hal-
1875; Bergman et al. 1988; Lambert 2016; Standring 2016). lucis longus (Tezer and Cicekcibasi 2012). The tendon of
It may also be joined with or send a slip to extensor hal- this muscle joins the tendon of extensor digitorum lon-
lucis brevis (Macalister 1875; Knott 1883b; Hallisy 1930; gus to insert onto the second digit (Tezer and Cicekcibasi
Bergman et al. 1988; Al-Saggaf 2003; Hill and Gerges 2012). The tendon of extensor digiti secundus may also
send a slip to the extensor hallucis longus tendon (Tezer and onto the proximal phalanx. In one case (0.3%), it presented
Cicekcibasi 2012). as a common insertion into the base of the proximal pha-
lanx with the extensor ossis metatarsi tendon from tibialis
Prevalence anterior.
Knott (1883b) found a slip extending from extensor hallucis Al-Saggaf (2003) studied the insertions of extensor hal-
longus to the base of the first metatarsal in 16 out of 40 lucis longus in 60 cadaveric lower limbs. In 39 limbs (65%),
cases (40%). Mori (1964) found that in 16% of cases, the the muscle inserted as a single tendon onto the distal hal-
tendon of extensor hallucis longus inserted onto the distal lucal phalanx. In 16 cases (26.7%), the muscle inserted as
phalanx of digit one without sending slips to other struc- two tendons. In nine of these 16 cases (15% of all limbs),
tures. In 78% of cases, the tendon sent a slip to the proximal the accessory tendon inserted onto the base of the proxi-
phalanx of digit one. In 2% of cases, the tendon divided into mal hallucal phalanx, distal to the insertion of extensor
two slips that both inserted onto the distal phalanx of digit hallucis brevis. In three cases (5% of all limbs), the acces-
one. In 4% of cases, the tendon sent a slip to the base of the sory tendon inserted with extensor hallucis brevis onto the
hallucal metatarsal. base of the proximal hallucal phalanx. In two cases (3.3%
Hallisy (1930) found that in 212 out of 290 feet (73.1%) of all limbs), the accessory tendon inserted onto the base
extensor hallucis longus had a single typical insertion onto of the proximal hallucal phalanx, medial to the insertion of
the base of the distal hallucal phalanx. In 70 feet (24.1%), extensor hallucis brevis. In two cases (3.3% of all limbs),
the tendon sent a slip to the base of the proximal hallucal the accessory tendon joined the tendon of extensor hallu-
phalanx. In two cases (0.7%), the tendon divided into three cis brevis, forming a common tendon that inserted onto the
slips, one going to the base of the distal phalanx of digit one base of the proximal hallucal phalanx. In five cases (8.3%),
and the other two inserting onto the base of the proximal the muscle had three tendons, with all accessory tendons
phalanx. In two cases (0.7%), the tendon sent a slip to the inserting onto the first metatarsophalangeal joint.
distal part of the dorsal surface of the first metatarsal. In In a sample of 32 cadaveric feet, Bibbo et al. (2004) found
three cases (1%), the tendon sent a slip to the most medial extensor hallucis capsularis tendons in 26 feet (81.3%). The
tendon of extensor digitorum brevis (extensor hallucis bre- tendons originated from extensor hallucis longus in 24 feet
vis). In one case (0.3%), the muscle had both the normal (92.3%) and from tibialis anterior in 2 feet (7.7%). In two
tendon and a tendon that split from the normal tendon at cases (7.7%), a small muscle belly that was separate from
the ankle joint to join with the extensor digitorum longus extensor hallucis longus and tibialis anterior gave rise to the
tendon going to digit two. tendon. All tendons inserted onto the first metatarsophalan-
Hallisy (1930) found slips corresponding to extensor geal joint capsule.
ossis primi metatarsi in 6 out of 290 feet (2.1%). In 3 feet In a sample of 81 cadaveric feet, Boyd et al. (2006) found
(1%), it presented as a slip from the tendon of tibialis ante- 73 extensor hallucis capsularis tendons in 71 feet (87.7%). Out
rior that inserted onto the shaft of the first metatarsal just of the 73 tendons, 23 arose from the extensor hallucis lon-
behind the head. In one case (0.3%), it presented as a sepa- gus muscle (31.5%), 45 arose from a bifurcation point off of
rate tendon of tibialis anterior that inserted onto the head extensor hallucis longus (61.6%), two arose from the tibialis
of the first metatarsal and base of the proximal hallucal anterior tendon (2.7%), one arose from the extensor hallucis
phalanx. In 2 feet (0.7%), it presented as a slip from exten- brevis tendon (1.4%), and two tendons had unidentified ori-
sor hallucis longus and inserted onto the distal part of the gins (2.7%). Seventy-two tendons inserted into the metatarso-
dorsal surface of the first metatarsal. phalangeal joint capsule of digit one (98.6%) and one inserted
Hallisy (1930) found slips corresponding to extensor into the base of the proximal hallucal phalanx (1.4%).
primi internodii hallucis in 85 out of 290 feet (29.3%). In Arora et al. (2011) studied extensor hallucis longus in
72 feet (24.8%), extensor primi internodii hallucis pre- 60 lower limbs from 30 cadavers. In 54 limbs (90%), the
sented as a secondary insertion of the extensor hallucis lon- muscle inserted normally but in six limbs (10%) the mus-
gus tendon into the base of the proximal phalanx of digit cle had doubled tendons. In four limbs (6.7%), the lateral
one. In seven cases (2.4%), it presented as a tendon from tendon inserted onto the base of the distal hallucal pha-
tibialis anterior that inserted onto the medial aspect of the lanx and the medial tendon inserted onto the base of the
proximal phalanx. In two cases (0.7%), it presented as a proximal hallucal phalanx. In two limbs (3.3%), the lateral
small muscular belly originating from tibialis anterior and tendon inserted onto the base of the distal hallucal phalanx
inserting onto the medial aspect of the base of the proxi- and the medial tendon inserted onto the head of the first
mal phalanx. In one case (0.3%), it presented as a muscu- metatarsal.
lar belly from the lateral aspect of the middle third of the In a sample of 60 patients who underwent operative cor-
tibia with an insertion onto the medial aspect of the base rection of hallux valgus and interphalangeus, Bayer et al.
of the proximal phalanx. In one case (0.3%), it originated (2014) found extensor hallucis capsularis tendons in 59 cases
as a tendon with a separate muscular belly from extensor (98.3%). In a sample of 98 cadaveric feet, Natsis et al. (2017)
hallucis longus in the distal leg and inserted onto the proxi- found accessory tendons of extensor hallucis longus in 26
mal phalanx. In one case (0.3%), it originated as a tendon feet (26.5%). In all cases, the accessory tendon inserted onto
from extensor hallucis longus in the distal leg and inserted the metatarsophalangeal joint capsule of digit one.

Windle (1893) found an extensor primi internodii hallucis Extensor digitorum longus originates from the lateral tibial
on the left side of one female fetus with anencephaly. In condyle, the medial fibular surface, and the interosseous
a male fetus with anencephaly, Windle (1893) found that membrane (Standring 2016). The muscle ends in a tendon
extensor hallucis longus inserted via two tendons. Bersu that divides into four tendinous slips that go to digits two
et al. (1976) describe two infants with Hanhart syndrome. through five (Standring 2016). The tendons to digits two
In the infant with an intact right foot, the tendon of extensor through four receive a contribution from extensor digitorum
hallucis longus fused with the tendon of extensor digitorum brevis (Standring 2016). Fibers from the lumbricals and
longus. interossei also contribute to form the dorsal digital expan-
Aziz (1979) described the bilateral presence of extensions of each digit (Standring 2016). The dorsal digital
sor primi internodii hallucis in one female neonate with expansions divide into three parts, with one slip inserting
trisomy 18. On the right side, the muscle arose from the onto the base of the intermediate phalanx and two slips that
anterior surface of the distal tibia (the text says proximal unite on the dorsal aspect of the intermediate phalanx to
tibia, but the illustration in the paper shows an origin from insert onto the base of the distal phalanx (Standring 2016).
the distal tibia). The muscle coursed over the ankle joint
medial to extensor hallucis longus and then split into three Innervation
tendons. One tendon inserted onto the medial surface of the Extensor digitorum longus is innervated by the deep fibular
proximal hallucal phalanx. The second tendon inserted on nerve (Standring 2016).
the dorsal surface of the proximal hallucal phalanx. The
third tendon inserted with the tendon of extensor hallucis Comparative Anatomy
longus onto the distal hallucal phalanx. On the left side, the Extensor digitorum longus has a similar typical presentation
muscle had one tendon that inserted onto the medial surface in the apes, extending from the tibia, fibula, and associated
of the proximal hallucal phalanx. fascial and aponeurotic structures to the dorsal aponeu-
Hootnick et al. (1987) describe an individual that had roses of digits two through five (Hepburn 1892; Beddard
a right limb with congenital tibial aplasia, talocalcaneal 1893; MacDowell 1910; Sonntag 1924; Raven 1950; Miller
synchondrosis, and an adducted foot with five toes. A ten- 1952; Lewis 1966; Gibbs 1999; Vereecke et al. 2005; Diogo
dinous band replaced the tibia. In this limb, extensor hallu- et al. 2010, 2012, 2013a,b, 2017). In orangutans, the origin
cis longus originated from the medial side of the tendinous may be fused with tibialis anterior and the tendon to digit
band. Its tendon divided into a main tendon with its usual two may be reduced or absent (Beddard 1893; Gibbs 1999;
insertion and two accessory tendons. One accessory ten- Ferrero 2011; Ferrero et al. 2012; Diogo et al. 2013b). The
don inserted onto connective tissue near the talus, and the tendon to digit two may also be reduced or absent in com-
second accessory tendon traveled with the tibialis ante- mon chimpanzees (Beddard 1893; Gibbs 1999). Extensor
rior tendon to insert onto the common flexor tendon sheet. digitorum longus may send extra tendons to digit five in
Extensor hallucis longus also sent a slip to the medial side bonobos (Diogo et al. 2017).
of the extensor expansion.
Variations
In their literature review, Smith et al. (2015) found that The terminal tendons, especially those going to digits two and
extensor primi internodii hallucis was only present in 1 out five, may be doubled (Macalister 1875; Bergman et al. 1988;
of 17 individuals with trisomy 18 (5.9%). Lambert 2016; Standring 2016). The tendons going to digits
three, four, or five may be reduced or absent, and in some
Clinical Implications cases replaced by tendons from other muscles (Macalister
Accessory tendons of extensor hallucis longus— 1875; Stevenson 1921; Sakuma et al. 2004; Plochocki and
particularly extensor hallucis capsularis tendons—are Bodeen 2010; Sirasanagandla et al. 2014; Lambert 2016).
often associated with the presence of, but not necessar- The tendons of extensor digitorum longus may send acces-
ily the severity of, hallux valgus (Al-Saggaf 2003; Bibbo sory slips to various metatarsals and proximal phalanges,
et al. 2004; Bayer et al. 2014; Natsis et al. 2017). The role of digit one, extensor digitorum brevis, extensor hallucis lon-
these accessory attachments in the development of hallux gus, extensor hallucis brevis, or the interossei (Macalister
valgus is thus unclear. 1875; Knott 1883b; Bergman et al. 1988; Stevens et al. 1993;
Lambert 2016; Standring 2016). The tendon going to digit
Extensor digitorum longus (Figure 5.8) five may receive contributions from, or join with, the tendon
of fibularis tertius (Macalister 1875; Bergman et al. 1988;
Synonyms Stevens et al. 1993; Lambert 2016). Each tendon may have
N/A its own muscular belly (Macalister 1875).
Prevalence Hootnick et al. (1987) describe an individual that had a

In a sample of 30 cases, Knott (1883b) found slips to the right limb with congenital tibial aplasia, talocalcaneal syn-
medial most tendon of extensor digitorum longus that chondrosis, and an adducted foot with five toes. In this limb,
attached to the proximal phalanx of digit one in two cases extensor digitorum longus became tendinous just above the
(6.7%). Slips to the base of the proximal phalanx of digit ankle joint and entered the dorsal surface of the foot at a
two were found in two cases (6.7%). A slip from the tendon right angle as it coursed under the extensor retinaculum. Its
to digit five inserted onto the middle of the fifth metatarsal tendon split into four slips that entered the extensor expan-
in one case (3.3%) and onto the base of the fifth metatarsal sions of digits two through five. The lateral most slip con-
in two cases (6.7%). In one case (3.3%), slips from the ten- tributed to the fibularis tertius tendon.
dons to digits three and four inserted onto the bases of the In the fetus with trisomy 18 and cyclopia dissected by
fourth and fifth metatarsals. Smith et al. (2015), the right extensor digitorum longus
Mori (1964) found that in 34% of cases, the distal por- exchanged a slip with the tendon of fibularis tertius at the
tion of extensor digitorum longus was divided into two lay- fourth digit and did not send a tendon to digit five. On the
ers, and the tendons to digits two and three arose from the right side of a fetus with craniorachischisis dissected by
superficial layer while the tendons to digits four and five Alghamdi et al. (2017), the tendon of extensor digitorum
arose from the deep layer. In 40% of cases, the distal part of longus going to digit three sent a slip to the tendon going
the muscle was divided into two layers, and the superficial to digit four. In the female fetus with trisomy 18 dissected
layer sent tendons to digits one, two, and three while the by Alghamdi et al. (2018), extensor digitorum longus was
deep layer sent a tendon to digit five. In 26% of cases, the proximally fused with fibularis tertius bilaterally.
distal part of the muscle was not divided and sent tendons
to digits two through five. Prevalence
Stevens et al. (1993) studied the relationships between N/A
fibularis tertius and extensor digitorum longus in 40 cadav-
eric lower limbs. The origin of fibularis tertius was continu-
ous with the origin of extensor digitorum longus in 33 limbs Understanding variations and potential absences of exten-
(82.5%). In six limbs (15%), the two muscles exchanged sor digitorum longus tendons is important when planning
fibrous bands. In four limbs (10%), the tendons of the fibu- to use these tendons for tendon grafts (Sakuma et al. 2004).
laris tertius and extensor digitorum longus were fused. In
one limb (2.5%), the tendon of extensor digitorum longus Fibularis tertius (Figure 5.8)
going to digit five originated from the muscle belly of fibu-
laris tertius. In two limbs (5%), extensor digitorum longus Synonyms
sent a slip to the fifth metatarsal. In one limb (2.5%), both This muscle may also be referred to as peroneus tertius
muscles gave slips that fused together and then inserted (Macalister 1875; Standring 2016).
onto the fifth metatarsal.
Description Fibularis tertius originates from the distal third of the
Macalister (1875) notes that in a deformed limb, Ringhoffer medial fibula, the interosseous membrane, and the anterior
found that extensor digitorum longus sent one tendon to intermuscular septum (Standring 2016). It inserts via a ten-
digit two, two tendons to digit four, and one tendon to digit don onto the base of the fifth metatarsal (Standring 2016).
five. Bersu et al. (1976) describe two infants with Hanhart
syndrome. In the infant with an intact right foot, the ten- Innervation
don of extensor hallucis longus fused with the tendon of Fibularis tertius is innervated by the deep fibular nerve
extensor digitorum longus. On the right side of a neonate (Standring 2016).
with trisomy 13, Pettersen et al. (1979) found that extensor
digitorum longus sent an extra tendon to the head of the Comparative Anatomy
fifth metatarsal. Fibularis tertius is variably present among the apes (Gibbs
Mieden (1982) describes an infant with median cleft lip, 1999; Diogo et al. 2010, 2012, 2013a,b, 2017). It seems to
hypotelorism, and alobar holoprosencephaly. On the left be absent in orangutans and present in about half of gib-
side, the tendons of extensor digitorum brevis and longus bons, 30% of gorillas, and 5% of common chimpanzees
were attached to the metatarsophalangeal joints (Mieden (Gibbs 1999; Diogo et al. 2010, 2012, 2013a,b, 2017). In
1982). On the left foot of a male neonate with Meckel syn- some bonobos, extensor digitorum longus sends an addi-
drome, Pettersen (1984) found that the extensor digitorum tional tendon to the fifth metatarsal (Diogo et al. 2017).
longus tendon to the second digit was rudimentary and the This extra slip may correspond to fibularis tertius (Diogo
tendons to digits three and four were absent. et al. 2017).
Variations slips to digit five and to the fourth metatarsal. Plochocki

Description and Bodeen (2010) found a supernumerary fibularis ter-
tius muscle distal to the main muscle. The supernumerary
Fibularis tertius may be enlarged, reduced, or absent
muscle arose from the medial margin and diaphysis of the
(Hallett 1848; Macalister 1875; Le Double 1897; Bergman
fibula and the interosseous membrane. It ended in a tendon
et al. 1988; Stevens et al. 1993; Chaney et al. 1996; Joshi
that passed anterior to the lateral malleolus and deep to the
et al. 2006; Rourke et al. 2007; Peddity and Velichety 2013;
extensor retinaculum and split into two slips that inserted
Lambert 2016; Standring 2016; Albay and Candan 2017;
into the calcaneus. It was supplied by the deep fibular nerve.
Olewnik 2019). Its origin may be shifted more proximally
on the fibula (Macalister 1875; Stevens et al. 1993; Joshi
et al. 2006; Rourke et al. 2007). Yildiz and Yalcin (2012) report Prevalence
an origin of fibularis tertius from extensor hallucis longus. Macalister (1875) notes that fibularis tertius may be absent
The origin of fibularis tertius is often continuous with in 10% of individuals. Chaney et al. (1996) found that
the origin of extensor digitorum longus (Stevens et al. fibularis tertius was absent in 25 out of 282 limbs (8.9%).
1993; Lambert 2016). The tendon of fibularis tertius may be Albay and Candan (2017) studied the fibular muscles in
connected or fused with the tendon of extensor digitorum 200 limbs from 100 fetuses. Fibularis tertius was absent in
longus going to digit five (Macalister 1875; Bergman et al. 40 limbs (20%). Krammer et al. (1979) found that peroneus
1988; Stevens et al. 1993; Joshi et al. 2006; Lambert 2016). tertius was present in 157 out of 169 lower limbs, yielding
Fibularis tertius may present as a tendinous band that arises an absence of 7.1% (12 limbs). Mori (1964) found that the
from extensor digitorum longus (Macalister 1875; Rourke tendon of fibularis tertius inserted onto the base of the fifth
et al. 2007; Peddity and Velichety 2013; Olewnik 2019). metatarsal in 68% of cases and onto the base of the fourth
Fibularis tertius may replace the tendon going to digit five metatarsal in 32% of cases.
when the tendon of extensor digitorum longus to this digit Stevens et al. (1993) studied fibularis tertius and its rela-
is absent (Macalister 1875; Stevenson 1921; Lambert 2016). tionships with extensor digitorum longus in 40 cadaveric
Fibularis tertius may send a slip to the digit five tendon of lower limbs. The origin of fibularis tertius was continuous
extensor digitorum brevis (Macalister 1875). with the origin of extensor digitorum longus in 33 limbs
Its insertion may expand to the shaft of the fifth meta- (82.5%). In six limbs (15%), the two muscles exchanged
tarsal (Joshi et al. 2006; Lambert 2016; Standring 2016; fibrous bands. In four limbs (10%), the tendons of the fibu-
Olewnik 2019). It may insert onto the extensor aponeu- laris tertius and extensor digitorum longus were fused. In
rosis or proximal phalanx of digit five (Macalister 1875; one limb (2.5%), the tendon of extensor digitorum longus
Krammer et al. 1979; Joshi et al. 2006; Jana and Roy 2011; going to digit five originated from the muscle belly of fibu-
Lambert 2016). The tendon of fibularis tertius may be dou- laris tertius. In one limb (2.5%), both muscles gave slips
bled or it may split into two or three parts prior to insertion that fused together and then inserted onto the fifth meta-
(Macalister 1875; Stevens et al. 1993; Raheja et al. 2005; tarsal. Fibularis tertius was absent in two limbs (5%). In
Joshi et al. 2006; Rourke et al. 2007; Lambert and Atsas 35 limbs (87.5%), fibularis tertius originated from the lower
2010; Jana and Roy 2011; Lambert 2016; Olewnik 2019). middle quarter of the fibula. In 37 limbs (92.5%), fibularis
It may attach onto or send a slip to the fourth metatarsal tertius had a single tendon, and in one limb (2.5%), it had
or insert entirely onto this bone in some cases (Macalister a double tendon. The insertion of fibularis tertius onto the
1875; Mori 1964; Bergman et al. 1988; Raheja et al. 2005; fifth metatarsal was bifurcated in two limbs (5%) and trifur-
Joshi et al. 2006; Rourke et al. 2007; Lambert 2016; cated in one limb (2.5%).
Standring 2016; Nayak 2017; Olewnik 2019). It may send an Joshi et al. (2006) studied fibularis tertius in 220 lower
accessory tendon to the fascia that covers the fourth inter- limbs from 110 cadavers. The muscle was absent in 14 right
osseus muscle (the fourth interosseous space) (Macalister limbs and nine left limbs, being absent in a total of 23 limbs
1875; Joshi et al. 2006; Lambert 2016; Olewnik 2019). (10.5%). Out of the 96 right limbs in which the muscle was
Fibularis tertius may connect with flexor digiti minimi bre- present, the muscle had a normal origin from the distal
vis or the tendon of fibularis brevis (Jana and Roy 2011; third of the fibula in 46 limbs (47.9%), an extensive origin
Olewnik 2019). Sirasanagandla et al. (2014) report a case in from the lower half of the fibula in 31 limbs (32.3%), and a
which the fibularis tertius tendon divided into three slips, very extensive origin from the lower three quarters of the
one that inserted on the base of the distal phalanx of digit fibula in 19 limbs (19.8%). Also out of these 96 right limbs,
five, another that inserted onto the base of the fifth meta- the muscle inserted onto the base of the fifth metatarsal
tarsal, and one that inserted into the fascia over the ankle in 44 limbs (45.8%), onto the base and shaft of the fifth
joint capsule. metatarsal in 11 limbs (11.5%), onto the base and shaft of
Fibularis tertius may be doubled (Macalister 1875; the fifth metatarsal and the fourth interosseous space in 22
Lambert and Atsas 2010; Plochocki and Bodeen 2010; limbs (22.9%), and had other insertions in 19 limbs (19.8%).
Lambert 2016). Macalister (1875) notes a case in which the Out of the 101 left limbs in which the muscle was pres-
posterior muscle of the doubled fibularis tertius sent two ent, the muscle had a normal origin from the distal third
tendons to the fifth metatarsal while the anterior muscle sent of the fibula in 55 limbs (54.5%), an extensive origin from
the lower half of the fibula in 29 limbs (28.7%), and a very an accessory slip going to the base of the fourth metatar-
extensive origin from the lower three quarters of the fibula sal. In two limbs (2.2%), the tendon was fused with a slip
in 17 limbs (16.8%). Also out of these 101 left limbs, the from the fibularis brevis tendon and the fourth interosseous
muscle inserted onto the base of the fifth metatarsal in 55 muscle originated from this fusion.
limbs (54.5%), onto the base and shaft of the fifth metatar- Several researchers have used clinical assessments to
sal in 13 limbs (12.8%), onto the base and shaft of the fifth visualize and palpate the fibularis tertius tendon. Witvrouw
metatarsal and the fourth interosseous space in 14 limbs et al. (2006) used palpation to determine the presence of
(13.9%), and had other insertions in 19 limbs (18.8%). fibularis tertius in a sample of 200 limbs from 100 individu-
Rourke et al. (2007) examined fibularis tertius in 82 als. These authors found that fibularis tertius was absent in
lower limbs from 41 cadavers. The muscle was absent in 37 limbs (18.5%).
five limbs (6.1%). In one limb (1.2%), the tendon was dou- Ramirez et al. (2010) used palpation to determine the
bled and inserted onto the shafts of the fourth and fifth presence of fibularis tertius in a sample of 336 limbs from
metatarsals. In all other limbs, the tendons had a single 168 individuals from Chile. These authors were able to
insertion onto the shafts of both these bones. Nayak (2017) identify fibularis tertius in 49.11% of limbs. Ashaolu et al.
studied fibularis tertius in 100 cadaveric limbs. Fibularis (2013) used palpation to determine the presence of fibularis
tertius was present in all limbs (100%). In all limbs (100%), tertius in a sample of 200 limbs from 100 individuals in
it originated from the distal fourth of the fibula. In 94 limbs Nigeria. These authors were able to identify fibularis tertius
(94%), it inserted onto the base of the fifth metatarsal. In in 125 limbs (62.5%). Potu et al. (2016) used palpation to
four limbs (4%), it inserted onto the fifth metatarsal up to determine the presence of fibularis tertius in a sample of
the metatarsophalangeal joint. In two limbs (2%), it inserted 390 legs from 195 Southeastern Indian individuals. These
near the base of the fourth metatarsal. authors were able to identify the muscle in 203 legs (52.1%).
Yammine and Erić (2017) conducted a literature review Salem et al. (2018) used palpation to determine the presence
of 35 studies regarding the anatomy of fibularis tertius. of fibularis tertius in a sample of 1,248 Arab patients. The
Out of 7,601 legs, they found that the muscle was present sample included 439 individuals from Bahrain, 208 indi-
in 93.2% of cadaveric legs and in 80% of legs examined in viduals from Saudi Arabia, 153 individuals from Kuwait,
clinical studies. Out of the 1,026 legs for which the origin of 198 individuals from Tunisia, and 250 individuals from
the muscle was recorded, it originated from the distal half Egypt. The authors were able to identify the presence of
of the fibula in 721 cases (70.3%), the distal third of the fib- fibularis tertius in 42% of the Bahraini individuals, 38.5%
ula in 133 cases (13%), and from extensor digitorum longus of Saudi individuals, 41.2% of Kuwaiti individuals, 67.7%
in 162 cases (15.8%). Out of the 1,248 limbs for which the of Tunisian individuals, and 52.8% of Egyptian individuals.
insertion was recorded, it inserted on the shaft of the fifth Palomo-López et al. (2019) used palpation to determine the
metatarsal in 152 cases (12.2%), the base of the fifth meta- presence of fibularis tertius in a sample of 481 individuals
tarsal in 252 cases (20.2%), both the base and shaft in 292 in Spain. These authors were able to identify the presence
cases (23.4%), both the fourth and fifth metatarsals in 423 of fibularis tertius in 184 participants (38.3%).
cases (33.9%), the fourth metatarsal in 38 cases (3%), and
Anomalies
onto the tendon of extensor digitorum longus in 24 cases
(2%). A slip to the head of the fifth metatarsal or to the base Description
of digit five was found in 67 cases (5.3%). On the left side of a female fetus with anencephaly, Windle
Olewnik (2019) studied fibularis tertius in a sample of (1893) noted that fibularis tertius formed the bulk of the
106 cadaveric lower limbs. Fibularis tertius was absent in muscle belly of extensor digitorum longus. This author also
15 limbs (14.2%). Out of the 91 limbs in which the muscle found that fibularis tertius was enlarged in a male fetus
was present, it originated from the distal half of the fib- with anencephaly. In a male infant with triploidy studied
ula and intermuscular septum in 61 limbs (67%) and the by Moen et al. (1984), this muscle was absent bilaterally.
distal third of the fibula and intermuscular septum in 20 In a male neonate with Meckel syndrome, Pettersen (1984)
limbs (22%). In ten limbs (11%), the muscle belly was miss- observed that peroneus tertius was absent on the right side.
ing and the fibularis tertius tendon arose from the extensor On the left side, this muscle was attached to the fifth and
digitorum longus tendon. In 41 limbs (45%), the tendon had sixth metatarsals.
a single insertion onto the shaft of the fifth metatarsal. In In four neonates with trisomy 13, Colacino and Pettersen
20 limbs (22%), the tendon had a single insertion onto the (1978) found that fibularis tertius was absent bilaterally in
base of the fifth metatarsal. In 15 limbs (16.5%), the tendon all four cases. Fibularis tertius was absent bilaterally in two
had a single, broad insertion onto the base and shaft of the neonates with trisomy 18 and three neonates with trisomy
fifth metatarsal and the fascia over the fourth interosseous 13 (Aziz 1979, 1980, 1981), and in a boy with trisomy 13q
space. In eight limbs (8.8%), the tendon had a split inser- described by Pettersen (1979). In the fetus with trisomy 18
tion, with one slip going to the base of the fifth metatarsal and cyclopia dissected by Smith et al. (2015), the right fibu-
and an accessory slip going to the shaft of this bone. In five laris tertius exchanged a slip with the tendon of extensor
lower limbs (5.5%), the tendon had a split insertion, with digitorum longus at the fourth digit, had a separate fleshy
one slip inserting onto the base of the fifth metatarsal and head, and had a distally shifted insertion. In the female
fetus with trisomy 18 dissected by Alghamdi et al. (2018), individuals (Lambert and Atsas 2010; Lambert et al. 2011b)
extensor digitorum longus was proximally fused with fibu- and unilaterally in one individual (Upadhyay and Amiras
laris tertius bilaterally. 2015).
In their literature review, Smith et al. (2015) found that N/A
fibularis tertius was absent in 12 out of 20 individuals with
trisomy 13 (60%) and absent in 2 out of 17 individuals with Clinical Implications
trisomy 18 (11.8%). Clinicians should consider the presence of anterior fibulo-
calcaneus as a potential cause of ankle pain (Lambert and
Clinical Implications Atsas 2010; Lambert et al. 2011b; Upadhyay and Amiras
Witvrouw et al. (2006) found that individuals who lack 2015).
fibularis tertius are not at higher risk for ankle ligament
injuries.
Fibularis longus (Figure 5.9)
See also: Fibularis quartus
Anterior fibulocalcaneus (Figure 5.8)
Synonyms Synonyms
N/A This muscle may also be referred to as peroneus longus
(Macalister 1875; Standring 2016).
This muscle is only present as a variation. Typical Presentation
Description
Comparative Anatomy Fibularis longus originates from the head and lateral sur-
N/A face of the fibula, and from the anterior and posterior inter-
muscular septa (Standring 2016). It ends in a tendon that
Variations inserts by one slip onto the base of the first metatarsal and
Description by another slip onto the medial cuneiform (Standring 2016).
Anterior fibulocalcaneus is an accessory fibular muscle
considered to be in the anterior compartment of the leg Innervation
(Lambert and Atsas 2010; Lambert et al. 2011b; Upadhyay Fibularis longus is innervated by the superficial fibular
and Amiras 2015; Lambert 2016). It has been found in asso- nerve (Standring 2016).
ciation with the doubling of fibularis tertius and its tendon
(Lambert and Atsas 2010), as well as the absence of fibu- Comparative Anatomy
laris tertius (Upadhyay and Amiras 2015). Fibularis longus has a similar typical presentation in the
Anterior fibulocalcaneus originates from the fibula, apes, extending from the proximal fibula to the first meta-
anterior intermuscular septum, and the fascia covering tarsal (Hepburn 1892; Beddard 1893; MacDowell 1910;
fibularis tertius (Lambert and Atsas 2010; Lambert et al. Sonntag 1924; Raven 1950; Miller 1952; Lewis 1966; Gibbs
2011b; Upadhyay and Amiras 2015; Lambert 2016). Its ori- 1999; Vereecke et al. 2005; Ferrero 2011; Ferrero et al.
gin on the fibula varies, with recorded origins ranging from 2012; Diogo et al. 2010, 2012, 2013a,b, 2017). It may have
the head to the distal half of the bone (Lambert and Atsas an additional origin from the lateral condyle of the tibia and
2010; Lambert et al. 2011b; Lambert 2016). It courses ante- have a fibrous connection to the fifth metatarsal in gibbons,
rior to the lateral malleolus and inserts onto the lateral sur- common chimpanzees, and bonobos (Miller 1952; Lewis
face of the calcaneus in the region of the peroneal trochlea 1966; Gibbs 1999; Diogo et al. 2013a, 2017). In orangutans,
(Lambert and Atsas 2010; Lambert et al. 2011b; Upadhyay it may be fused or otherwise connected with fibularis bre-
and Amiras 2015; Lambert 2016). Its tendon may send a slip vis, extensor digitorum longus, and flexor hallucis longus
to the capsule of the talocrural joint (Lambert and Atsas (Beddard 1893; Sonntag 1924; Gibbs 1999).
2010).
Variations
Anterior fibulocalcaneus is innervated by the deep fibular Some fibers from the lateral tibial condyle, and more rarely
nerve (Lambert et al. 2011b). the lateral femoral condyle, may contribute to the origin of
this muscle (Bergman et al. 1988; Standring 2016). Mehta
Prevalence et al. (2011a) report a case in which fibularis longus was
Prevalence is unknown but, to our knowledge, anterior divided into superficial and deep portions, with the super-
fibulocalcaneus has been only found bilaterally in three ficial belly inserting onto the first metatarsal and the deep
FIGURE 5.9 Lateral muscles of the lower leg in lateral view. Fibularis quartus is illustrated on the right side, and the other muscles
are illustrated on the left.
belly inserting onto the lateral aspect of the calcaneus. This Knott 1883b; Shyamsundar et al. 2012; Verma and Arora
variation was also associated with a split fibularis brevis. 2012; Lambert 2016; Standring 2016; Edama et al. 2020).
As many as four slips of insertion may be present Slips may insert onto the navicular or tendon of tibialis pos-
(Verma et al. 2011; Verma and Arora 2012). The inser- terior (Mori 1964). A slip may pass to the first interosse-
tion onto the medial cuneiform may be absent (Mori 1964; ous space (Jayakumari et al. 2006; Chhaparwal et al. 2015;
Shyamsundar et al. 2012; Chhaparwal et al. 2015; Edama et Edama et al. 2020). It may have an insertion onto the cal-
al. 2020) or doubled (Verma and Arora 2012). Its insertion caneus (Athavale et al. 2012b; Shyamsundar et al. 2012) or
may have an additional attachment onto the first metatar- intermediate cuneiform (Lambert 2016). Fibularis longus
sal (Macalister 1875; Knott 1883b; Jayakumari et al. 2006; may connect to the cuboid (Patil et al. 2007; Cromeens and
Lambert 2016). Accessory slips may pass to the base of the Reeves 2011; Athavale et al. 2012b; Lambert 2016; Edama
second, third, fourth, or fifth metatarsals (Macalister 1875; et al. 2020). It may also send a slip to the lateral malleolus
or lateral ligaments of the ankle (Macalister 1875). Via the the tendon flared into a triangular expansion and had a nor-
anterior or posterior frenular ligaments, the tendon of fibu- mal insertion. In 9 feet (23.7%), the tendon flared into a tri-
laris longus may have connections to flexor digiti minimi angular expansion and had a normal insertion but received
brevis, opponens digiti minimi, some interosseous muscles, a slip from tibialis posterior. In 1 foot (2.6%), the tendon
and the fifth metatarsal (Edama et al. 2020). sent a slip to flexor hallucis brevis, and in 3 feet (7.9%), the
In rare cases, fibularis longus may fuse with fibularis tendon sent a slip to the first interosseous space. In 4 feet
brevis (Macalister 1875; Knott 1883b; Jayakumari et al. (10.5%), the tendon inserted onto the first metatarsal only.
2006; Lambert 2016; Standring 2016). A slip may extend In 2 feet (5.3%), the oblique head of adductor hallucis arose
to adductor hallucis (Standring 2016) or flexor hallucis bre- from the tendon.
vis (Chhaparwal et al. 2015). Tibialis posterior may send Edama et al. (2020) studied the insertion of fibularis lon-
a slip or insert onto the tendon of fibularis longus (Mori gus in 104 feet from 52 Japanese cadavers. The fibularis
1964; Lohrmann et al. 1997; Bloome et al. 2003; Raheja longus tendon received a slip from the tibialis posterior ten-
et al. 2005; Chhaparwal et al. 2015; Lambert 2016; Edama don in 32 feet (30.8%). All feet exhibited an insertion into
et al. 2020). Flexor digiti minimi brevis, the oblique head the first metatarsal. Additional insertions were present onto
of adductor hallucis, and some interosseus muscles may the medial cuneiform in 21 feet (20.2%) and the first dorsal
originate from the fibularis longus tendon (Lamont 1908; interosseous muscle in 38 feet (36.5%). The anterior frenu-
Ochiltree 1912; Harbeson 1933, 1938; Bergman et al. 1988; lar ligament was present in 33 feet (31.7%), and it attached
Jayakumari et al. 2006; Verma et al. 2011; Chhaparwal to flexor digiti minimi brevis and opponens digiti minimi
et al. 2015; Lambert 2016; Edama et al. 2020). in 33 feet (31.7%), the second plantar interosseous muscle
in 13 feet (12.5%), the third plantar interosseous muscle in
Prevalence 29 feet (27.9%), the fourth dorsal interosseous muscle in 20
In a sample of 40 cases, Knott (1883b) found that fibularis feet (19.3%), and the fifth metatarsal in 9 feet (8.7%). The
longus was fused with fibularis brevis in three cases (7.5%). posterior frenular ligament was present in 6 feet (5.8%), and
In two cases (5%), a slip from the tendon went to the base it attached to the fourth metatarsal in 4 feet (3.9%), the fifth
of the fifth metatarsal. In one case (2.5%), a slip passed to metatarsal in 1 foot (1%), and the cuboid in 1 foot (1%).
the head of the first metatarsal. In one case each (2.5%), a
slip passed to the base of the third and fourth metatarsals. Anomalies
Mori (1964) found that fibularis longus inserted onto Description
the first metatarsal in 64% of cases, the first metatarsal and Macalister (1875) notes that in a case of clubfoot, fibularis
medial cuneiform in 28% of cases, the first metatarsal and longus inserted via three tendons onto the first, third, and
navicular in 2% of cases, and the first metatarsal, medial fifth metatarsals. Hootnick et al. (1987) describe an indi-
cuneiform, and tendon of tibialis posterior in 4% of cases. vidual that had a right limb with congenital tibial aplasia,
Mori (1964) also found that in 10% of cases, the tendon of talocalcaneal synchondrosis, and an adducted foot with
tibialis posterior inserted onto the navicular, medial cunei- five toes. In this limb, fibularis longus became tendinous at
form, and tendon of fibularis longus. midleg. Its tendon inserted onto the distal end of the fibula.
Bloome et al. (2003) studied the insertions of the tibi- Its tendon sent a slip to the connective tissue of the groove
alis posterior tendon in 11 feet from 10 cadavers. Each ten- posterior to the lateral malleolus.
don had three distinct bands and their insertions included In a male fetus with triploidy studied by Moen et al.
the fibularis longus tendon in four cases (36.4%). Athavale (1984), fibularis longus had an extra slip attaching to the
et al. (2012b) studied the peroneal muscles in 92 cadaveric calcaneus on the left side. In a female fetus with trisomy
lower limbs. In five cases (5.4%), the fibularis longus ten- 18, the proximal two thirds of fibularis longus and fibu-
don split into two slips. The accessory slip inserted into laris brevis were fused bilaterally (Alghamdi et al. 2018).
the cuboid in one case, the peroneal trochlea in two cases, Pettersen et al. (1979) found extra slips of fibularis longus
and joined with the main tendon of fibularis longus in two in two male neonates with trisomy 13. On the left side of
cases. one specimen, a thin tendon originated from the muscle
Shyamsundar et al. (2012) studied the insertion of fibu- belly of fibularis longus and crossed the fibularis brevis
laris longus in 26 feet from 14 cadavers. All 26 feet had tendon to insert onto the posterolateral calcaneal surface.
an insertion onto the first metatarsal, but the insertion onto On the right side of the second specimen, a supernumerary
the medial cuneiform was absent in four specimens (15.4%). muscle belly originated from the anterior intermuscular
Slips passed to the base of the fourth/fifth metatarsals in 12 septum just anterior to fibularis longus. This belly became
feet (46.2%), base of the third metatarsal in 2 feet (7.7%), tendinous in the middle of the lower leg and crossed the
base of the second metatarsal in 8 feet (30.8%), and to the fibularis brevis tendon to insert onto the anterolateral cal-
calcaneus in 8 feet (30.8%). caneal surface. On the left side of this same specimen, two
Chhaparwal et al. (2015) studied the insertion of fibu- supernumerary muscles originated anterior to fibularis lon-
laris longus in 38 feet from 19 cadavers. In 15 feet (39.5%), gus. One inserted onto the posterior surface of the lateral
fibularis longus had a normal insertion onto the base of the malleolus, and the other inserted onto the inferior peroneal
first metatarsal and the medial cuneiform. In 4 feet (10.5%), retinaculum of the calcaneus.
Prevalence et al. 2013; Cecava and Campbell 2015). It may send a slip to
In their literature review, Smith et al. (2015) found that the the extensor tendon of digit five (Knott 1883b). It may send a
extra slips of fibularis longus found by Pettersen et al. (1979) slip to the distal phalanges of digits four or five (Mori 1964). A
were only present in the two neonates they described, out of slip from the fibularis brevis tendon to the metatarsal, dorsal
a sample of 20 individuals with trisomy 13 (10%). digital expansion, or phalanges of digit five may be designated
as fibularis digiti minimi (Cunningham and St. John Brooks
Clinical Implications 1889–1891; Le Double 1897; Macalister 1867b, 1875; Raheja
Mehta et al. (2011a) suggest that doubled peroneal muscles et al. 2005; Jadhav et al. 2013; Lambert 2016; Chaney et al.
and tendons could potentially contribute to ankle instability 2018; Olewnik et al. 2019b) (see the entry for this muscle).
and lateral ankle pain. In rare cases, fibularis brevis may fuse with fibularis lon-
gus (Macalister 1875; Knott 1883b; Jayakumari et al. 2006;
Lambert 2016; Standring 2016). It may have an attachment
Fibularis brevis (Figure 5.9) with flexor digiti minimi brevis or send a slip to fibularis
See also: Fibularis quartus tertius (Knott 1883b; Bergman et al. 1988; Lambert 2016;
Olewnik et al. 2019b). It may contribute to the origin of
Synonyms abductor digiti minimi (Macalister 1875; Lambert 2016)
This muscle may also be referred to as peroneus brevis or the fourth dorsal interosseous muscle (Olewnik et al.
(Macalister 1875; Standring 2016). 2019b). Mehta et al. (2011a) report a case in which fibularis
brevis was divided into superficial/lateral and deep/medial
portions, with the superficial/lateral belly inserting onto the
fifth metatarsal and the deep/medial belly inserting onto
Description the lateral aspect of the calcaneus. This variation was also
Fibularis brevis originates from the distal two-thirds of the associated with a split fibularis longus.
lateral fibula and from the anterior and posterior intermus-
cular septa (Standring 2016). Its tendon inserts onto the lat- Prevalence
eral surface of the base of the fifth metatarsal (Standring Knott (1883b) found that in 6 out of 40 cases (15%), fibu-
2016). laris brevis sent a slip to the extensor tendon of digit five. In
a review of 200 MRI examinations, Cecava and Campbell
Innervation (2015) found two cases (1%) of the fibularis brevis tendon
Fibularis brevis is innervated by the superficial fibular ending on the calcaneus. Mori (1964) found that fibularis
nerve (Standring 2016). brevis inserted onto the base of the fifth metatarsal in 47%
of cases, sent a slip to the distal part of the fifth metatarsal
Comparative Anatomy in 10% of cases, and sent a slip to the distal phalanx of
Fibularis brevis has a similar typical presentation in the the fourth digit in 5% of cases. Mori (1964) found that the
apes, extending from the distal fibula to the base of the tendon sent a slip to the distal phalanx of digit five in 20%
fifth metatarsal (Hepburn 1892; Beddard 1893; MacDowell of cases, with this slip having a separate muscular belly in
1910; Sonntag 1924; Raven 1950; Miller 1952; Lewis 1966; 8% of cases.
Gibbs 1999; Vereecke et al. 2005; Diogo et al. 2010, 2012, Olewnik et al. (2019b) studied the insertion of fibularis
2013a,b, 2017). It may have an insertion into the tendon of brevis in 102 lower limbs. In 72 cases (70.6%), the tendon
extensor digitorum brevis going to digit five in orangutans had a single insertion onto the base of the fifth metatarsal.
and common chimpanzees (Champneys 1872; Hepburn In 30 cases (29.4%), the tendon had a bifurcated attachment
1892; Beddard 1893; MacDowell 1910; Sonntag 1924; with the main tendon having a normal insertion onto the
Gibbs 1999; Ferrero 2011; Ferrero et al. 2012). In common fifth metatarsal. In 23 cases (22.5%), the accessory ten-
chimpanzees, it may have an insertion onto the proximal don inserted onto the dorsal surface of the base of the fifth
or intermediate phalanges of digit five (Champneys 1872; metatarsal. In five cases (4.9%), the accessory tendon sent a
MacDowell 1910; Diogo et al. 2013a). band to the dorsal surface of the base of the fifth metatarsal
and a band to the shaft of the fifth metatarsal. In two cases
Variations (2%), the accessory tendon sent a band to the dorsal surface
Description of the base of the fifth metatarsal while another band fused
The tendon of fibularis brevis may insert onto the dorsal sur- with fibularis tertius and gave origin to the fourth dorsal
face, shaft, and/or distal aspect of the fifth metatarsal (Knott interosseous muscle.
1883b; Mori 1964; Bergman et al. 1988; Lambert 2016;
Olewnik et al. 2019b; Rosser et al. 2019). The tendon may Anomalies
also insert into the fourth dorsal interosseous muscle (Knott Description
1883b; Bergman et al. 1988; Lambert 2016). The tendon may In a female fetus with trisomy 18, the proximal two-thirds
also be truncated and insert onto or near the peroneal trochlea of fibularis longus and fibularis brevis were fused bilater-
without extending into the foot (Athavale et al. 2012b; Lucas ally (Alghamdi et al. 2018).
Prevalence 1990; Sammarco and Brainard 1991; Cheung et al. 1997;

N/A Chepuri et al. 2001; Zammit and Singh 2003; Sookur et al.
2008; Athavale et al. 2012b; Clarkson et al. 2013; Lambert
Clinical Implications 2016; Standring 2016; Albay and Candan 2017; Mustafa
Mehta et al. (2011a) suggest that doubled peroneal mus- et al. 2017). This calcaneal insertion (peroneocalcaneus
cles and tendons could potentially contribute to ankle or fibulocalcaneus externum variant) is the most common
instability and lateral ankle pain. An insertion of the presentation of fibularis quartus (Hecker 1923; Cheung
fibularis brevis tendon onto the calcaneus may be misdi- et al. 1997; Saupe et al. 2007; Lambert 2016). Its insertion
agnosed as a tear of the tendon (Lucas et al. 2013; Cecava onto the calcaneus may be split into two tendons (Knott
and Campbell 2015). 1883b). It may have an attachment to flexor hallucis longus
(Macalister 1875). Fibularis quartus may be doubled (Hur
et al. 2015a).
Fibularis quartus (Figure 5.9) Fibularis quartus has several other variants that are named
for their insertions beyond the calcaneus. Fibularis digiti
See also: Fibularis longus, Fibularis brevis
minimi or fibularis digiti quinti refers to a variant of fibularis
Synonyms quartus that inserts onto the fifth digit (see the entry for this
This muscle may also be referred to as peroneus quartus muscle) (Macalister 1875; Bergman et al. 1988; Sookur et al.
(Macalister 1875; Bergman et al. 1988), peroneocalcaneus 2008; Lambert 2016; Olewnik et al. 2019b). Fibularis quartus
externum (Otto) (Donley and Leyes 2001; Sookur et al. may insert onto the cuboid, thus having the name peroneo-
2008; Lambert 2016), or fibulocalcaneus externus (Albay cuboideus (fibulocuboideus) (Chudzinski 1875; Macalister
and Candan 2017). Other synonyms include peroneus sex- 1875; Bergman et al. 1988; Donley and Leyes 2001; Sookur
tus, peroneus calcaneus externus, or peroneus medius (Le et al. 2008; Clarkson et al. 2013; Lambert 2016; Standring
Double 1897; Knott 1883a; Lambert 2016). 2016). Tubbs et al. (2008b) use the name peroneotalocalca-
neus to describe a muscle that originated from the anterior
intermuscular septum and fibularis longus and inserted onto
both the calcaneus and talus. Clarkson et al. (2013) use the
This muscle is only present as a variation or anomaly. name peroneocalcaneocuboideus to describe a muscle that
originated from fibularis brevis, the distal fibula, and the
Comparative Anatomy
posterior intermuscular septum and inserted into both the
Fibularis quartus appears to be a muscle variation unique calcaneus and the cuboid.
to humans (Hecker 1923; Athavale et al. 2012b; Lambert Fibularis accessorius (peroneoperoneolongus) is
2016). As such, muscles equivalent to fibularis quartus— regarded as a variant of fibularis quartus that takes origin
accessory fibular muscles in the lateral compartment that from either fibularis brevis or fibularis longus and inserts
insert onto the calcaneus—have not been found in the apes onto the fibularis longus tendon (Macalister 1875; Knott
(Gibbs 1999; Diogo et al. 2010; 2012, 2013a,b, 2017). 1883b; Hecker 1923; White et al. 1974; Bergman et al.
1988; Saupe et al. 2007; Sookur et al. 2008; Lambert 2016;
Variations Standring 2016). The muscle may also insert into the fibu-
Description laris brevis tendon (Athavale et al. 2012b; Hur et al. 2015a;
Fibularis quartus is an accessory muscle found in the lateral Lambert 2016).
compartment of the leg (Chaney et al. 1996; Sookur et al.
2008; Lambert 2016; Standring 2016). It typically origi- Innervation
nates from the posterior intermuscular septum, from the Fibularis quartus and its variants are innervated by the
bellies of fibularis brevis or fibularis longus, and/or from the superficial fibular nerve (Tubbs et al. 2008b; Athavale et al.
distal fibula (Wood 1867b; Macalister 1867b, 1875; Knott 2012b).
1883a; Hecker 1923; Bergman et al. 1988; Sobel et al. 1990;
Sammarco and Brainard 1991; Chaney et al. 1996; Chepuri Prevalence
et al. 2001; Donley and Leyes 2001; Zammit and Singh In a sample of 124 legs from 65 cadavers, Sobel et al. (1990)
2003; Sookur et al. 2008; Athavale et al. 2012b; Clarkson found fibularis quartus in 27 legs (21.8%). In 17 out of the
et al. 2013; Hur et al. 2015a; Lambert 2016; Standring 2016; 27 legs (63%), it originated from the belly of fibularis brevis
Albay and Candan 2017; Mustafa et al. 2017). and inserted into the peroneal trochlea. In 1 out of the 27
The tendon of fibularis quartus courses medial and pos- legs (3.7%), the muscle originated from fibularis brevis and
terior to the tendons of fibularis brevis and fibularis lon- inserted onto the fibularis longus tendon (fibularis accesso-
gus (Macalister 1867b, 1875; Cheung et al. 1997; Sookur rius). In another leg (3.7%), the muscle originated from the
et al. 2008). It most often inserts onto the lateral surface of fibularis brevis muscle and inserted onto its tendon. In one
the calcaneus, usually into the retrotrochlear eminence or leg (3.7%), the muscle originated from fibularis longus and
peroneal trochlea (Macalister 1867b, 1875; Wood 1867b; inserted into the fibularis longus tendon. In one leg (3.7%),
Knott 1883a; Hecker 1923; Bergman et al. 1988; Sobel et al. the muscle originated from fibularis longus and inserted
into fibularis brevis. In two legs (7.4%), the muscle pre- Cheung et al. (1997) studied a sample of 136 ankle MRI
sented as fibularis digiti minimi (see the entry for this mus- studies to determine the prevalence of fibularis quartus. This
cle). In two legs (7.4%), fibularis quartus originated from muscle was found in 14 cases (10.3%), inserting into the calca-
fibularis longus and inserted onto the peroneal trochlea. In neus in 11 of these cases. In a sample of 31 MR images from
three legs (11.1%), the muscle originated from fibularis bre- 27 patients that exhibited longitudinal splits of the fibularis
vis and inserted onto the lateral retinaculum. brevis tendon, Rosenberg et al. (1997) found fibularis quartus
Chaney et al. (1996) found that fibularis quartus was in two images (6.5%). Using sonography and MRI, Chepuri
present in 8 out of 269 limbs (3%). In a sample of 92 cadav- et al. (2001) found seven instances of fibularis quartus mus-
eric lower limbs, Athavale et al. (2012b) found fibularis cles in a sample of 32 patients (21.9%). Nascimento et al.
quartus in 20 limbs (21.7%). The tendon of fibularis quartus (2012) found 16 instances of fibularis quartus out of a sam-
inserted onto the retrotrochlear eminence in 15 out of the ple of 211 MR examinations (7.6%). Using MRI, Agha et al.
20 cases (75%) and into the peroneal trochlea in three cases (2018) found fibularis quartus in 15 out of 181 patients (8.3%).
(15%). In two cases (10%), it inserted onto the tendon of Yammine (2015c) conducted a comprehensive review
either fibularis longus or fibularis brevis. In a sample of 277 of fibularis quartus. This author found that out of a total
cadaveric legs, Clarkson et al. (2013) found fibularis quartus of 2,816 feet from 20 cadaveric studies, fibularis quartus
in 58 legs (20.9%), with 56 of these muscles presenting as has a “true” prevalence of 10.2%. Based on a total of 252
fibulocalcaneus externum, one of 58 these muscles present- cadavers from five cadaveric studies, fibularis quartus has a
ing as peroneocuboideus, and another muscle presenting as “crude” prevalence of 16.6%. Based on a total of 253 oper-
peroneocalcaneocuboideus. In a sample of Indian cadavers, ated ankles from six studies, fibularis quartus has a surgical
Prakash et al. (2011) found fibularis quartus in 3 out of 70 prevalence of 5.5%. Based on a total of 854 legs from 12
specimens (4.3%). In a sample of 20 Saudi Arabian cadav- studies, fibularis quartus has an MRI prevalence of 10.6%.
eric lower limbs, Mustafa et al. (2017) found fibularis quar- Albay and Candan (2017) studied the fibular muscles
tus in four limbs (20%). in 200 limbs from 100 fetuses. In four limbs (2%), “fibu-
In a sample of 80 lower limbs from 40 Korean cadavers, laris quartus” muscles were present, with an origin from
Hur et al. (2015a) found fibularis quartus in 13 specimens fibularis brevis and an insertion onto the lateral calcaneal
(16.3%). In two specimens, fibularis quartus was doubled, surface. In three limbs (1.5%), they found accessory fibular
thus yielding 15 instances of fibularis quartus within the muscles that originated from the fibula and inserted onto
sample. Fibularis quartus originated from fibularis brevis the lateral surface of the calcaneus. These authors deemed
in 12 cases (15%). The fibularis quartus muscle belly con- these muscles to be examples of “fibulocalcaneus externus.”
verged into a tendon in eight of these 12 cases, with that ten-
don inserting into the tendon of fibularis brevis in 3 cases Anomalies
(3.8%), the calcaneus in 2 cases (2.5%), the inferior pero- Description
neal retinaculum in 2 cases (2.5%), and the base of the fifth Pettersen (1979) found peroneus quartus bilaterally in a
metatarsal in 1 case (1.3%). The fibularis quartus originated boy with trisomy 13q. On the left side, it originated in the
from fibularis brevis as a tendon in four of these 12 cases, distal third of the leg with fibularis longus along the poste-
inserting onto the calcaneus in three of them (3.8%) and rior intermuscular septum. On the right side, it originated
into the periosteum of the retromalleolar grove in 1 case from the distal third of the fibula and along the posterome-
(1.3%). The remaining three cases (3.8%) did not arise from dial margin of fibularis brevis. On both sides, the tendon
fibularis brevis. In one of these cases, it extended from the inserted onto the peroneal trochlea. Pettersen et al. (1979)
fibula to the tendon of fibularis brevis. In the second case, found peroneus quartus bilaterally in two male neonates
it originated from the superior peroneal retinaculum and with trisomy 13. These muscles originated from the fibula
inserted onto the calcaneus. In the third case, a doubled between fibularis brevis and flexor hallucis longus and
fibularis quartus was present, with the tendon of one aris- inserted onto the calcaneus behind the tendons of fibularis
ing from the muscle belly of the other. brevis and fibularis longus.
Zammit and Singh (2003) searched for the presence of Mieden (1982) describes two male fetuses with cyclo-
fibularis quartus in 102 legs from 80 cadavers using dissec- pia and alobar holoprosencephaly. On the left side of one
tion, and in 80 symptomatic patients using MRI. Fibularis specimen, the authors state that peroneus digiti quinti was
quartus was present in six of the cadaveric legs (5.9%), with present, originating from the distal fibula and attaching to
two presenting as fibularis digiti quinti and one presenting the peroneal trochlea. Based on the insertion of this muscle,
as peroneocuboideus. Fibularis quartus was also present in it should be designated as fibularis quartus instead of the
six of the MRI images (7.5%). Saupe et al. (2007) used MRI variant that inserts onto digit five.
to study the peroneal region in 65 patients with asymptom-
atic ankles. Fibularis quartus was found in 11 ankles (17%), Prevalence
with ten of these muscles (91%) inserting onto the calcaneus In their literature review, Smith et al. (2015) found that fibu-
(peroneocalcaneus externum variant) and one muscle (9%) laris quartus was present in 3 out of 20 individuals with
inserting onto the fibularis longus tendon (peroneoperoneo- trisomy 13 (15%) and in 5 out of 17 individuals with trisomy
longus variant). 18 (29.4%).
Clinical Implications However, it is possible that the two are entirely separate
The presence of fibularis quartus and its variants may lead structures, due to differences in their shape, composition,
to lateral ankle pain, weakness, instability, and/or swelling and prevalence (Yammine 2015c).
(White et al. 1974; Buschmann et al. 1991; Sammarco and Fibularis digiti minimi has multiple presentations. It
Brainard 1991; Chaney et al. 1996; Trono et al. 1999; Donley may present as a tendinous slip that originates from the ten-
and Leyes 2001; Martinelli and Bernobi 2002; Zammit and don of fibularis tertius or, more often, the tendon of fibu-
Singh 2003; Sookur et al. 2008; Lotito et al. 2011; Lambert laris brevis (Cunningham and St. John Brooks 1889–1891;
2016). Its presence may cause peroneal compartment syn- Le Double 1897; Macalister 1867b, 1875; Jadhav et al. 2013;
drome (Wenning et al. 2019). Fibularis quartus and its vari- Lambert 2016; Chaney et al. 2018; Olewnik et al. 2019b).
ants may mimic a soft tissue mass during medical imaging Alternatively, it may present as an independent muscle
(Bergman et al. 1988; Tubbs et al. 2008b). Sookur et al. belly that originates from fibularis brevis or the distal fib-
(2008) note that the fibularis quartus tendon may be mis- ula (Macalister 1875; Mori 1964; Zammit and Singh 2003;
taken for a longitudinal tear of the fibular tendons. However, Sookur et al. 2008; Lambert 2016). Fibularis digiti minimi
fibularis quartus has been found in tandem with longitudi- may also present as a small, independent muscle belly that
nal splitting of the fibularis brevis tendon (Sobel et al. 1990; arises in the foot from the tendon of fibularis brevis and
Rosenberg et al. 1997; Trono et al. 1999; Chepuri et al. 2001; sends its own tendon to digit five (Cunningham and St. John
Zammit and Singh 2003). It has also been associated with Brooks 1889–1891; Raheja et al. 2005).
impingement, subluxation, tendinitis, and tenosynovitis of Fibularis digiti minimi has variable insertions onto
the fibular tendons (Trono et al. 1999; Chepuri et al. 2001; the fifth digit, including the base and/or head of the fifth
Lambert 2016; Opdam et al. 2017; Agha et al. 2018). metatarsal, the proximal phalanx, or the extensor expan-
sion (Cunningham and St. John Brooks 1889–1891; Le
Double 1897; Macalister 1875; Sobel et al. 1990; Zammit
Fibularis digiti minimi (Figure 5.9)
and Singh 2003; Raheja et al. 2005; Sookur et al. 2008;
See also: Fibularis quartus Jadhav et al. 2013; Lambert 2016; Chaney et al. 2018;
Olewnik et al. 2019b). It may insert via two tendons onto
Synonyms the base and head of the fifth metatarsal (Sobel et al.
This muscle may also be referred to as peroneus quintus 1990; Sookur et al. 2008). It may also have an insertion
(Knott 1883b), peroneus digiti quinti or peroneus quinti into both the fourth and fifth metatarsals (Jadhav et al.
digiti (Macalister 1867b, 1875), or fibularis digiti quanti 2013; Demir et al. 2015). It may join with or insert onto
(Olewnik et al. 2019b). the extensor digitorum longus tendon going to digit five
(Knott 1883b; Jadhav et al. 2013; Demir et al. 2015;
Typical Presentation Olewnik et al. 2019b).
This muscle is present only as a variation or anomaly.
Innervation
Comparative Anatomy When it presents as an independent muscle belly, fibularis
Muscles and tendinous slips that may correspond to fibu- digiti minimi is likely innervated by the superficial fibular
laris digiti minimi have been found in gorillas, common nerve, as are the other variants of fibularis quartus (e.g.,
chimpanzees, and bonobos (Champneys 1872; Macalister Tubbs et al. 2008b; Athavale et al. 2012b).
1873; MacDowell 1910; Raven 1950; Gibbs 1999; Diogo et
al. 2010, 2013a, 2017). In gorillas, slips have been found (1) Prevalence
extending between fibularis brevis and the extensor expan- In a sample of 45 lower limbs, Cunningham and St. John
sion of the fifth digit and (2) inserting onto the proximal Brooks (1889–1891) found fibularis digiti minimi in well-
and intermediate phalanges of digit five (Macalister 1873; developed form (a slip from the tendon of fibularis brevis
Raven 1950; Gibbs 1999; Diogo et al. 2010). In common that inserted onto the extensor expansion of digit five) in 21
chimpanzees, the tendon of fibularis brevis may send a slip cases (46.7%) and in rudimentary form (a slip that ended
to the proximal and/or intermediate phalanges of digit five on the dorsal surface of the fifth metatarsal) in five cases
(Champneys 1872; MacDowell 1910; Diogo et al. 2013a). (11.1%).
In one bonobo dissected by Diogo et al. (2017), extensor Mori (1964) found “peroneus digiti quinti” extending
digitorum longus sent an extra tendon to the fifth digit with from the distal fibula to the fifth digit in 2 out of 73 legs
attachments onto the metatarsal, proximal phalanx, and (2.7%). Mori (1964) also notes that the fibularis brevis ten-
extensor expansion of this digit. don sent a slip to the distal phalanx of digit five in 20% of
cases, with this slip having a separate muscular belly in 8%
Variations of cases. These could also be presentations of fibularis digiti
Description minimi. Bareither et al. (1984) found fibularis digiti minimi
Fibularis digiti minimi is regarded as a variant of fibularis in 178 out of 298 cadaveric feet (59.7%). Sobel et al. (1990)
quartus that inserts onto the fifth digit (Bergman et al. found fibularis digiti minimi extending between fibularis
1988; Sobel et al. 1990; Sookur et al. 2008; Lambert 2016). brevis and the fifth metatarsal in 2 out of 124 legs (1.6%). In
a sample of 102 legs from 80 cadavers, Zammit and Singh Hootnick et al. (1987) describe an individual that had
(2003) found fibularis digiti minimi in two legs (2%). a right limb with congenital tibial aplasia, talocalcaneal
Jadhav et al. (2013) found slips corresponding to fibu- synchondrosis, and an adducted foot with five toes. In this
laris digiti minimi in 51 out of 100 cadaveric lower limbs limb, there was an accessory muscle that originated from
(51%). They inserted onto the dorsal digital expansion of the fibula deep to fibularis tertius and inserted via a long
digit five in 41% of these 51 cases, onto the common exten- tendon onto the fifth metatarsal.
sor tendon in 19% of cases, onto the base of the proximal
phalanx of digit five in 16% of cases, onto the head of the Prevalence
fifth metatarsal in 21% of cases, and onto both the shaft of Ramirez-Castro and Bersu (1978) found fibularis digiti
the fourth metatarsal and the head of the fifth metatarsal in minimi in five out of eight infants (62.5%) with trisomy 18.
3% of cases. In their literature review, Smith et al. (2015) found that fibu-
In a sample of 25 cadaveric lower limbs, Demir et al. laris digiti minimi was present in 2 out of 20 individuals
(2015) found fibularis digiti minimi in eight limbs (32%). with trisomy 13 (10%) and in 3 out of 17 individuals with
Five of these eight cases (62.5%) had a single tendon and trisomy 18 (17.6%). It seems like this assessment neglected
three (37.5%) had a doubled tendon, thus yielding 11 ten- to include the specimens described by Ramirez-Castro and
dons. Out of these 11 tendons, three (27.3%) inserted onto Bersu (1978).
the base of the fifth metatarsal, two (18.1%) inserted onto
the “dorsolateral ridge” of the fifth metatarsal, four (36.4%) Clinical Implications
inserted onto the fifth digit near the proximal phalanx, one N/A
(9.1%) inserted onto the extensor digitorum longus tendon
going to digit five, and one (9.1%) inserted onto both the MUSCLES OF THE FOOT
fourth metatarsal and the fifth metatarsal.
Yammine (2015c) conducted a comprehensive review of Abductor hallucis (Figure 5.10)
fibularis digiti minimi. This author found that out of a total
Synonyms
of 1,112 feet from eight cadaveric studies, fibularis digiti
minimi had a “true” prevalence of 34.3%. Based on a total N/A
of 194 cadavers from three cadaveric studies, fibularis digiti
minimi had a “crude” prevalence of 21.5%.
Chaney et al. (2018) dissected 52 limbs from 26 cadav- Description
ers to determine the prevalence of fibularis digiti minimi. Abductor hallucis originates from the flexor retinaculum,
A fully present fibularis digiti minimi—defined as a tendon the plantar aponeurosis, and the medial process of the cal-
that originated from the fibularis brevis tendon and inserted caneal tuber (Standring 2016). Its tendon inserts onto the
onto the extensor expansion of digit five—was present in base of the proximal phalanx of digit one (Standring 2016).
17 limbs (32.7%). A rudimentary fibularis digiti minimi—
defined as either a thin tendon or a tendon that did not reach Innervation
the extensor expansion—was present in 20 limbs (38.5%). Abductor hallucis is innervated by the medial plantar nerve
Olewnik et al. (2019b) found “fibularis digiti quanti” in 18 (Standring 2016).
out of 102 lower limbs (17.6%). It inserted onto the base
of the proximal phalanx of digit five in eight cases (7.8%), Comparative Anatomy
onto the extensor expansion of digit five in five cases (4.9%), Abductor hallucis has a similar typical presentation in the
and fused with the extensor digitorum longus tendon in five apes, originating from the medial and plantar surfaces of
cases (4.9%). the calcaneus and the plantar aponeurosis and inserting
onto the base of the proximal hallucal phalanx (Hepburn
1892; Beddard 1893; Sonntag 1924; Raven 1950; Miller
Anomalies 1952; Gibbs 1999; Diogo et al. 2010, 2012, 2013a,b, 2017).
Description It may be blended with flexor hallucis brevis in orangutans
Ramirez-Castro and Bersu (1978) found fibularis digiti and gibbons, and with flexor digitorum brevis in orang-
minimi in five infants with trisomy 18. These muscles origi- utans (Beddard 1893; Sonntag 1924; Gibbs 1999; Diogo et
nated from the distal fibula and inserted onto the base of the al. 2012, 2013b).
fifth metatarsal and sent tendinous slips to the base of digit
five. Aziz (1979) found fibularis digiti minimi bilaterally Variations
in one neonate with trisomy 18. It extended from the distal Description
fibula to join the insertion of fibularis brevis onto the fifth Abductor hallucis often has an insertion onto the medial
metatarsal. Aziz (1979, 1980, 1981) found fibularis digiti sesamoid bone and/or the “medial sesamoidal ligament”
minimi in two neonates with trisomy 13 and two neonates (fibers that extend from the medial sesamoid bone to the
with trisomy 18. These muscles originated from the distal joint capsule at the head of the first metatarsal) (Brenner
fibula and inserted into the fifth metatarsal. 1999; Agawany and Meguid 2010; Kafka et al. 2016;
FIGURE 5.10 First layer of plantar foot muscles in plantar (inferior) view.
Standring 2016). Abductor hallucis may send a tendinous (1883b) notes that Krause found a slip from abductor hal-
slip to the base of the proximal phalanx of digit two (Wood lucis going to the base of the proximal phalanx of digit two
1868; Macalister 1875; Knott 1883b) or to the proximal pha- in 9% of cases. Brenner (1999) studied the insertions of
langes of digits two and three (Bergman et al. 1988; Kafka abductor hallucis in 109 feet. The muscle inserted onto the
et al. 2016). It may send a slip to the flexor hallucis longus proximal phalanx of the hallux in 42 feet (38.5%), into the
tendon (Macalister 1875; Knott 1883b). It may also be con- medial sesamoid ligament and the medial sesamoid bone in
nected via a slip to the skin on the medial margin of the foot 65 feet (59.6%), and into the medial sesamoid bone only in 2
(Macalister 1875; Bergman et al. 1988; Standring 2016). feet (1.8%). Agawany and Meguid (2010) studied the inser-
Chittoria et al. (2015) report an unusual case in which tions of abductor hallucis in 15 feet. The muscle inserted
abductor hallucis had numerous tendinous slips along three onto the proximal phalanx of digit one in 7 feet (46.7%), into
quarters of its internal surface that attached to the medial the base of the proximal phalanx and the sesamoid bone via
intermuscular septum arising from the connection of the two slips in 5 feet (33.3%), into the sesamoid bone only in 1
tibial and central components of the plantar aponeurosis, foot (6.7%), and into both the base of the proximal phalanx
the medial surface of the hallucal metatarsal, and the inter- and the metatarsophalangeal joint capsule of digit one in 2
muscular septum abductor hallucis shares with flexor hal- feet (13.3%).
lucis brevis.
Edwards et al. (1969) found an accessory belly of abduc- Anomalies
tor hallucis that originated from the interior surface of the Description
navicular. Bhansali and Bhansali (1997) found an accessory Mieden (1982) describes two male fetuses with cyclo-
muscle that originated from the deep surface of the fascia pia and alobar holoprosencephaly. On the left side of one
covering the posterior tibial nerve just above the medial specimen, abductor hallucis was doubled. Hootnick et al.
malleolus, passed deep to the nerve, and then coursed (1987) describe an individual that had a right limb with con-
through the tarsal tunnel to unite with abductor hallucis. genital tibial aplasia, talocalcaneal synchondrosis, and an
adducted foot with five toes. In this limb, abductor hallucis
Prevalence was absent. In the female fetus with trisomy 18 dissected
Wood (1868) found a slip from abductor hallucis going to by Alghamdi et al. (2018), the distal end of abductor hallu-
the base of the proximal phalanx of digit two in 5 out of cis was fused with the medial head of flexor hallucis brevis
40 males (12.5%) and in 1 out of 30 females (3.3%). Knott bilaterally.
N/A Description
The tendons of flexor digitorum brevis may receive acces-
sory slips from other muscles, often flexor digitorum lon-
An accessory abductor hallucis can entrap the posterior gus or quadratus plantae (Wood 1868; Macalister 1875;
tibial nerve (Edwards et al. 1969; Bhansali and Bhansali Nathan and Gloobe 1974; Bergman et al. 1988; Kafka et al.
1997). It is unclear whether variations in the insertion of 2016; Standring 2016). Any of the digital tendons may be
abductor hallucis contribute to the etiology of hallux valgus absent, but the tendon to digit five is most commonly absent
(Brenner 1999). (Wood 1867b, 1868; Macalister 1875; Knott 1883b; Aasar
1947; Mori 1964; Sato 1970; Bergman et al. 1988; Nathan
Flexor digitorum brevis (Figure 5.10) and Gloobe 1974; Chaney et al. 1996; Claassen and Wree
2003; Lobo et al. 2008; Gugapriya 2012; Kafka et al. 2016;
Synonyms Standring 2016).
It may also be referred to as flexor brevis digitorum When a tendon is absent, particularly the tendon going to
(Macalister 1875). digit five, it may be replaced by a muscular slip originating
from the tendon of flexor digitorum longus, quadratus plan-
Typical Presentation tae, tendon of tibialis posterior, the medial process of the
Description calcaneus, the lateral band of the plantar aponeurosis, and/
or the lateral intermuscular septum (Wood 1864, 1867b,
Flexor digitorum brevis originates via a tendon from the
1868; Macalister 1875; Knott 1883b; Aasar 1947; Nathan
medial process of the calcaneal tuber and from the planar apo-
and Gloobe 1974; Bergman et al. 1988; Claassen and Wree
neurosis (Standring 2016). It ends in four tendons that travel to
2003; Asomugha et al. 2005; Gugapriya 2012; Kafka et al.
digits two through five (Standring 2016). These tendons split
2016; Standring 2016; Ramesh Rao and Rao 2017).
to accommodate the tendons of flexor digitorum longus and
The tendon to digit five may also fail to divide around
then insert onto both sides of the shaft of the intermediate
the tendon of flexor digitorum longus (Macalister 1875).
phalanx of their respective digits (Standring 2016).
This tendon may also end on the anterior fascia of digit five
Innervation (Macalister 1875; Bergman et al. 1988). This tendon may
be thin or otherwise reduced (Claassen and Wree 2003;
Flexor digitorum brevis is innervated by the medial plantar
Yalçin and Ozan 2005b; Bernhard et al. 2013). The inser-
tion onto digit five may be split into two slips, one going
Comparative Anatomy to the proximal phalanx and the other inserting with flexor
digitorum longus onto the distal phalanx (Yalçin and Ozan
Flexor digitorum brevis has a similar typical presentation in
2005b).
the apes, extending from the calcaneus to various combina-
Yalçin and Ozan (2005b) describe a case in which flexor
tions of the intermediate phalanges of the toes (Champneys
digitorum brevis had a superficial head and a deep head.
1872; Hepburn 1892; Beddard 1893; Dwight 1895;
Both heads contributed to the tendons going to digits three
MacDowell 1910; Sonntag 1924; Raven 1950; Miller 1952;
and four, and the deep head formed separate muscle bellies
Gibbs 1999; Diogo et al. 2010, 2012, 2013a,b, 2017). As in
to send tendons to digits two and five. This morphology is
humans, the insertion onto the fifth digit is not constant.
similar to that seen in some ape specimens, though differ-
In gibbons, there may be an origin from flexor digi-
ent digits are targeted by the superficial and deep heads (see
torum longus, and the tendons usually insert onto digits
comparative anatomy, above).
two, three, and four with an occasional insertion onto digit
five (Hepburn 1892; Beddard 1893; Diogo et al. 2012). In
orangutans, the tendon always inserts onto digits two and Prevalence
three, occasionally onto digit four, and rarely onto digit five Wood (1868) found that the flexor digitorum brevis tendon
(Chapman 1880; Hepburn 1892; Beddard 1893; Primrose to digit five was absent in 10 out of 68 males (14.7%) and
1900; Sonntag 1924; Diogo et al. 2013b). In common chim- in 5 out of 34 females (14.7%). Knott (1883b) found that the
panzees and bonobos, the tendons always go to digits two tendon to digit five was absent in 3 out of 30 cases (10%).
and three and sometimes to digits four and five (Hepburn Knott (1883b) also notes that Krause found it was absent in
1892; Miller 1952; Diogo et al. 2013a, 2017). In gorillas, a 15% of cases. Mori (1964) found that the tendon to digit five
superficial head of the muscle inserts onto digits two and was absent in 16% of feet. Sato (1970) found that the flexor
three, and a deep head inserts onto digit four and sometimes digitorum brevis tendon to digit five was absent in 106 out
five (Raven 1950; Diogo et al. 2010). When the tendons to of 380 limbs (27.9%) from Kyushu Japanese males and in
digits four and five are present in the apes besides goril- 72 out of 252 limbs (28.6%) from females.
las, they are often described as deriving from a deep head Out of a sample of 100 feet, Nathan and Gloobe (1974)
of flexor digitorum brevis (Hepburn 1892; Primrose 1900; found that the flexor digitorum brevis tendon to digit five
Gibbs 1999; Diogo et al. 2012, 2013a,b). was absent in 23 cases (23%). The tendons to both the fourth
and fifth digits were absent in 3 feet (3%). Flexor digito- Absence of the digit five tendon is also a common feature
rum longus sent slips to the flexor digitorum brevis tendon in individuals with trisomy 18 (Ramirez-Castro and Bersu
to digit five in 20 cases (20%) and to the brevis tendons 1978; Alghamdi et al. 2018) and trisomy 21 (Bersu 1980).
to digits four and five in 3 cases (3%). The fifth tendon of On the right foot of a male neonate with Meckel syn-
flexor digitorum brevis had an origin from flexor digitorum drome, Pettersen (1984) found that flexor digiti minimi
longus in five cases (5%), from the intermuscular septum in brevis attached to digit five and received a slip from flexor
one case (1%), and from tibialis posterior in one case (1%). digitorum brevis that did not split to accommodate the
Chaney et al. (1996) found that the tendon to digit five tendon of flexor digitorum longus. The transverse head
was absent in 181 out of 284 limbs (63.7%). In a sample of of adductor hallucis also a partial origin from the flexor
11 feet from 8 cadavers, Kura et al. (1997) found that the digitorum brevis to the fifth digit. In a fetus with cranio-
tendon to digit five was absent in 4 feet (36.4%). Out of a rachischisis dissected by Alghamdi et al. (2017), the flexor
sample of 97 feet from 60 cadavers, Bernhard et al. (2013) digitorum brevis tendon to digit five was absent on both
found that 47 did not have a tendon to the fifth digit (48.4%), sides. Hootnick et al. (1987) describe an individual that had
25 feet had small tendons to the fifth digit (25.8%), and the a right limb with congenital tibial aplasia, talocalcaneal
remaining 25 feet (25.8%) demonstrated the typical presen- synchondrosis, and an adducted foot with five toes. In this
tation of flexor digitorum brevis. Yalçin and Ozan (2005b) limb, flexor digitorum brevis did not send a tendon to the
studied flexor digitorum brevis in 33 feet from 15 cadavers. fifth digit.
The muscle had a typical presentation in 15 feet (45.5%).
The tendon to digit five was small in 12 feet (36.4%) and Prevalence
absent in 6 feet (18.2%). In their literature review, Smith et al. (2015) found that the
In a sample of 60 feet from 30 Nepalese cadavers, Lobo flexor digitorum brevis tendon to digit five was absent in 5
et al. (2008) found that the flexor digitorum brevis tendon out of 20 individuals with trisomy 13 (25%). Based on 16
for the fifth digit was absent in all 60 feet (100%). Gugapriya sides from eight infants with trisomy 18, Ramirez-Castro
(2012) examined flexor digitorum brevis in 30 feet from 15 and Bersu (1978) found that the flexor digitorum brevis ten-
cadavers from the northern Tamil Nadu region of India. The don to digit five was absent in 11 out of 16 sides (68.75%) and
tendon to digit five was absent in 25 feet (83.3%). In a sam- from seven out of eight infants (87.5%). According to Bersu
ple of 270 feet from 135 Sri Lankan cadavers, Ilayperuma (1980), the tendon to digit five was absent on at least one side
(2012) found that the tendon to digit five was absent bilater- in two out of five individuals with trisomy 21 (40%).
ally in 194 limbs (71.9%) from 67 cadavers. Ferreira Arquez
(2017) studied flexor digitorum brevis in 34 feet from 17 Clinical Implications
cadavers. In 2 feet (5.9%) the tendon to digit five was absent. Understanding variations in the presentation of the flexor
In the remaining 32 feet (94.1%), the muscle sent tendons to digitorum brevis tendons is important for planning tendon
digits two through five. transfer surgeries (Ilayperuma 2012).
Yammine (2015d) conducted a comprehensive review of
flexor digitorum brevis. This author found that out of a total
Abductor digiti minimi (Figure 5.10)
of 2,789 feet from 22 studies, the “true” prevalence of the
absence of the tendon to digit five was 31.3%. Based on a Synonyms
total of 416 cadavers from six studies, the “crude” preva- N/A
lence of the absence of the tendon to digit five was 47%.
Based on a total of 476 feet from four studies, the “true” Typical Presentation
prevalence of the tendon to digit five presenting as a thin Description
slip was 47.7%. Based on a total of 223 feet from four stud- Abductor digiti minimi originates from the calcaneal
ies, the “true” prevalence of variations in origin of the fifth tuberosity and the plantar aponeurosis (Standring 2016).
tendon was 12.7%. Its tendon blends with that of flexor digiti minimi brevis
to insert onto the base of the proximal phalanx of digit five
Anomalies (Standring 2016).
Description
On the left foot of a boy with trisomy 13q, Pettersen (1979) Innervation
found that flexor digitorum brevis only had two tendons of Abductor digiti minimi is innervated by the lateral plantar
insertion. These tendons inserted onto the middle phalanges nerve (Standring 2016).
of digits three and four. On the right foot of this individ-
ual, only the tendon to digit five was absent. Pettersen et al. Comparative Anatomy
(1979) found that flexor digitorum brevis lacked a tendon to Abductor digiti minimi has a similar typical presentation in
digit five in two fetuses and two neonates with trisomy 13. the apes, extending from the calcaneus and the plantar apo-
In one of these cases, flexor digitorum brevis was a distinctly neurosis to the proximal phalanx of digit five (Champneys
separate muscle. The tendon to digit five was also absent 1872; Hepburn 1892; Beddard 1893; Sonntag 1924; Raven
bilaterally in another neonate with trisomy 13 (Aziz 1980). 1950; Miller 1952; Gibbs 1999; Vereecke et al. 2005; Diogo
et al. 2010, 2012, 2013a,b, 2017). The insertion may be split Prevalence
into two tendons in gibbons and bonobos (Vereecke et al. Knott (1883b) found that abductor digiti minimi sent a ten-
2005; Diogo et al. 2012). dinous slip to the base of the fifth metatarsal in 60% of
subjects.
Variations
Description Anomalies
Abductor digiti minimi may be fused with flexor digiti min- Description
imi brevis (Macalister 1875; Kopuz et al. 1999). Its insertion On the left side of a female fetus with trisomy 13, Pettersen
onto the proximal phalanx may be separate from the inser- et al. (1979) found that abductor digiti minimi had an acces-
tion of flexor digiti minimi brevis (Macalister 1875; Knott sory slip. On the left side of male neonate with trisomy 13,
1883b). Abductor digiti minimi is associated with several Pettersen et al. (1979) found that abductor digiti minimi
accessory muscles (Bergman et al. 1988; Kafka et al. 2016; attached to the extra digit in that foot instead of onto digit
Standring 2016). A short and deep accessory head may five. On the right foot of a male neonate with Meckel syn-
originate from the tuberosity (styloid process) of the fifth drome, Pettersen (1984) observed that abductor digiti min-
metatarsal and insert onto the base of the proximal pha- imi went to the sixth digit and sent a small slip to the fifth
lanx of digit five with abductor digiti minimi (Macalister digit. On the left foot, this muscle (along with flexor digiti
1875; Knott 1883b; Bergman et al. 1988; Kopuz et al. 1999; minimi brevis) was represented by a mass on the plantar
Kafka et al. 2016; Standring 2016). Abductor accessorius and lateral sides of digit six with vague attachments to this
digiti minimi arises from the lateral plantar process of the digit and partially to digit five.
calcaneus and inserts into the lateral aspect of the base of
the proximal phalanx of digit five (Bergman et al. 1988; Prevalence
Nayyar et al. 2010; Kafka et al. 2016). In their literature review, Smith et al. (2015) found that
Abductor digiti minimi may send a tendinous slip to the abductor digiti minimi had an extra slip in only 1 out of 24
base of the fifth metatarsal (Knott 1883b). In some cases, individuals with trisomy 13 (4.2%).
this slip may present as abductor ossis metatarsi digiti
quinti, a supernumerary muscle that extends between the Clinical Implications
lateral plantar process of the calcaneus and the tuberosity An accessory abductor of digit five may present as a soft
of the fifth metatarsal (see the entry for this muscle) (Wood tissue lump on the lateral border of the foot (Carmont et al.
1864, 1867b, 1868; Macalister 1875; Knott 1883b; Bergman 2002).
et al. 1988; Chaney et al. 1996; Kopuz et al. 1999; Kafka
et al. 2016; Standring 2016). Abductor ossis metatarsi digiti quinti (Figure 5.10)
Kopuz et al. (1999) report a left foot in which there were
multiple accessory muscles present in association with the Entry adapted by permission from Springer Nature Customer
normal abductor digiti minimi. Abductor ossis metatarsi Service Centre GmbH: Springer Current Molecular Biology
digiti quinti arose from the lateral process of the calca- Reports, Muscles Lost in Our Adult Primate Ancestors
neus and inserted onto the tuberosity of the fifth metatar- Still Imprint in US: on Muscle Evolution, Development,
sal. A “distal belly” was present extending between the Variations, and Pathologies. E. Boyle, V. Mahon, R. Diogo,
tuberosity of the fifth metatarsal and the lateral aspect of 2020.
the base of the proximal phalanx of digit five. Lastly, a See also: Abductor digiti minimi
medial accessory muscle was present extending between
the plantar surface of the calcaneus and the base of the Synonyms
fifth metatarsal. This muscle is also referred to as abductor ossis metatarsi
Boupha and Wree (1995) report a bicipital accessory quinti (Wood 1864), abductor os metatarsi minimi digiti
muscle situated in between the superficial and deep poste- (digiti minimi) (Macalister 1875), or abductor metatarsi
rior crural muscles that they name “abductor digiti minimi quinti (Gruber 1886).
longus.” The tibial head originated from the tibia distal to
the soleal line. The fibular head originated from the poste- Typical Presentation
rior intermuscular septum just proximal to the lateral mal- This muscle is only present as a variation.
leolus. The heads united to end in a tendon that coursed
behind the medial malleolus posterior to the tendon of Comparative Anatomy
flexor hallucis longus. After entering the plantar aspect Ferrero et al. (2012) are the first to report this muscle in
of the foot, the tendon passed laterally and inserted with gibbons and found an insertion onto the proximal phalanx
abductor digiti minimi onto the proximal phalanx of the of the fifth digit. In the other apes, abductor ossis metatarsi
fifth digit. These authors designate this muscle as a novel digiti quinti is frequently present as a distinct muscle that
variation but note that it shares some similarities with other extends between the calcaneus and the tuberosity of the
accessory muscles of the calf (e.g., tensor fasciae plantaris fifth metatarsal (Champneys 1872; Beddard 1893; Primrose
(tibiotarsalis), flexor digitorum accessorius longus). 1899; Raven 1950; Miller 1952; Gibbs 1999; Diogo et al.
2010, 2012, 2013a,b, 2017). Wood (1868) notes that similar the plantar surface of the calcaneus and the base of the
muscles have been found monkeys, cats, bears, and other fifth metatarsal.
mammals.
Innervation
Variations Abductor ossis metatarsi digiti quinti is innervated by the
Description lateral plantar nerve (Gibbs 1999; Kopuz et al. 1999).
Abductor ossis metatarsi digiti quinti is a variation of abduc-
tor digiti minimi that is typically not present in humans Prevalence
(Swindler and Wood 1973). It is a supernumerary muscle Wood (1867b) found abductor ossis metatarsi digiti quinti in
that originates from the lateral plantar process of the calca- 17 out of 36 subjects (47.2%) and later (Wood 1868) found
neus and inserts onto the tuberosity (styloid process) of the this muscle in 19 out of 36 subjects (52.8%). Knott (1883b)
fifth metatarsal (Wood 1864, 1867b, 1868; Macalister 1875; found this muscle in 3 out of 40 cases (7.5%). Le Double
Knott 1883b; Bergman et al. 1988; Chaney et al. 1996; (1897) reports a prevalence of 43% (from examinations of
Kopuz et al. 1999; Kafka et al. 2016; Standring 2016). It 65 feet) and of 45% (from examinations of 40 feet). Chaney
may have an origin from the medial process of the cal- et al. (1996) found this muscle in 125 out of 275 feet (45.5%).
caneus (Wood 1867b; Macalister 1875). Its insertion may
Anomalies
shift distally to the midsection or the anterior portion of the
fifth metatarsal (Wood 1867b; Macalister 1875; Bergman et N/A
al. 1988; Kafka et al. 2016). It may be fused with abductor
digiti minimi (Macalister 1875; Bergman et al. 1988).
Kopuz et al. (1999) report a left foot in which there were N/A
multiple accessory muscles present in association with the
normal abductor digiti minimi. Abductor ossis metatarsi Quadratus plantae (Figure 5.11)
digiti quinti arose from the lateral process of the calca-
neus and inserted onto the tuberosity of the fifth metatar- Synonyms
sal. A “distal belly” was present extending between the This muscle may also be referred to as flexor accessorius
tuberosity of the fifth metatarsal and the lateral aspect of pedis (Macalister 1875), flexor accessorius (Standring 2016),
the base of the proximal phalanx of digit five. Lastly, a flexor digitorum accessorius (Knott 1883b), flexor digito-
medial accessory muscle was present extending between rum accessorius brevis (Verma et al. 2018), moles carnea or
FIGURE 5.11 Second layer of plantar muscles and the plantar interossei in plantar (inferior) view.
massa carnea (Sylvius), caro plantae pedis quadrata, or caro Quadratus plantae may insert entirely onto, or connect
quadrata (Pretterklieber 2018). via an accessory slip to, the tendon of flexor hallucis lon-
gus (Macalister 1875; Knott 1883b; Bergman et al. 1988;
Typical Presentation Tubbs et al. 2004b; Hur et al. 2011b; Athavale et al. 2012a;
Description Gupta and Nasir 2013; Pretterklieber 2017, 2018; Beger et
The medial head of quadratus plantae originates from the al. 2018b). Hur et al. (2011b) and Beger et al. (2018b) found
medial plantar surface of the calcaneus (Standring 2016). that the insertion of quadratus plantae into the tendons of
The lateral head originates from the lateral plantar sur- both long flexor muscles is common. Quadratus plantae is
face of the calcaneus (Standring 2016). Quadratus plantae often involved in the variable blending of the flexor digi-
inserts into the tendon of flexor digitorum longus at the torum longus and flexor hallucis longus tendons, which is
juncture where it splits into four tendons that go to digits referred to as the chiasma plantare or the knot of Henry
two through five (Standring 2016) (O’Sullivan et al. 2005; LaRue and Anctil 2006; Plaass et al.
2013; Pretterklieber 2017, 2018; Beger et al. 2018b; Elvan
Innervation et al. 2019). Quadratus plantae may send an accessory slip
to the flexor tendon of digit five or to the flexor digitorum
Quadratus plantae is innervated by the lateral plantar nerve
brevis tendon of digit three (Macalister 1875). It may receive
(Standring 2016).
a slip from fibularis brevis (Macalister 1875). It may also
Comparative Anatomy be associated with a slip that inserts onto the distal phalanx
of digit five (Macalister 1875; Claassen and Wree 2003). Its
Quadratus plantae is absent in gibbons but sometimes found
insertion may expand to the lumbricals (Verma et al. 2018).
in the other apes with an insertion onto the flexor digito-
The accessory muscle flexor digitorum accessorius lon-
rum longus tendon (Gibbs 1999; Diogo et al. 2010, 2012,
gus originates from the posterior compartment of the lower
2013a,b, 2017). When the muscle is present in orangutans,
leg and usually inserts the main tendon of flexor digitorum
the lateral head is more common than the medial head
longus and/or into quadratus plantae (see the entry for this
(Chapman 1880; Hepburn 1892; Gibbs 1999; Lewis 1962;
muscle) (Hallett 1848; Wood 1864, 1868; Bergman et al.
Diogo et al. 2013b). When present in the gorillas, com-
1988; Sammarco and Conti 1994; Peterson et al. 1995;
mon chimpanzees, and bonobos, the medial head is absent
Cheung et al. 1999; Gümüşalan and Kalaycioğlu 2000;
(Champneys 1872; Hepburn 1892; Gibbs 1999; Diogo et al.
Sookur et al. 2008; Bowers et al. 2009; Georgiev et al. 2009;
2010, 2013a, 2017).
Hur et al. 2011b; Upasna et al. 2011; Standring 2016). This
muscle may be described as high origin of quadratus plan-
Variations
tae (e.g., Macalister 1875). Hallett (1848) notes that quadra-
Description tus plantae may be unusually large when flexor digitorum
Quadratus plantae may be diminutive, missing either accessorius longus is present. The presence of flexor digi-
the lateral or medial head, or entirely absent (Bergman torum accessorius longus may also be associated with the
et al. 1988; Kura et al. 1997; Hur et al. 2011b; Athavale et al. absence of quadratus plantae (Pác and Malinovský 1985).
2012a; Standring 2016; Pretterklieber 2018; Edama et al.
2019). Three heads may be present (Yalçin and Ozan Prevalence
2005b; Pretterklieber 2018). Its fibers or its insertion may In a sample of 18 feet, Lewis (1982) found that quadratus
be split into a superficial and deep layer (Bergman et al. plantae contributed to the flexor tendons going to digits two
1988; Verma et al. 2018). It may originate more proximally and three in 4 cases (22.2%) and into the flexor tendons
on the calcaneus, from the upper surface of the calcaneal going to digits two, three, and four in 14 cases (77.8%). In
tuber near the insertion of the plantaris tendon or from the a sample of 11 feet from eight cadavers, Kura et al. (1997)
Achilles tendon (Macalister 1875; Knott 1883b). It may also found that the lateral head of quadratus plantae was absent
originate from the calcaneocuboid ligament (Macalister in 1 foot (9.1%). Hur et al. (2011b) examined quadratus plan-
1875; Pretterklieber 2018). Pretterklieber (2018) lists sev- tae in 50 cadaveric specimens. The muscle was absent on
eral other origins for both the medial and lateral heads (see one side of one cadaver (2%). Forty muscles (80%) had two
prevalence information, below). heads of origin, five muscles had a medial head only (10%),
Its insertion onto the tendon(s) of flexor digitorum lon- and four muscles had a lateral head only (8%). Quadratus
gus is variable (Macalister 1875; Lewis 1962; Bergman et al. plantae inserted onto both the tendon of flexor digitorum
1988; Standring 2016; Pretterklieber 2017; Elvan et al. 2019). longus and tendinous slips of flexor hallucis longus in 48
The tendons of flexor digitorum longus that go digit three cases (96%) and into the flexor digitorum longus tendon
(in rare cases), digit four, and digit five (frequently) may be only in two cases (4%).
missing contributions from quadratus plantae (Macalister Athavale et al. (2012a) studied quadratus plantae in 47
1875; Lewis 1962; Standring 2016; Pretterklieber 2017, 2018; cadaveric limbs. The medial head was absent in five limbs
Edama et al. 2019; Elvan et al. 2019). The quadratus plantae (10.6%). In four limbs (8.5%), the medial head did not unite
contribution to the flexor tendon going to digit two may also with the lateral head. Out of the 42 limbs in which it was
be lacking (Pretterklieber 2017, 2018; Elvan et al. 2019). present, the medial head inserted into the flexor digitorum
longus tendon in 35 limbs (83.3%), into the flexor digitorum 6 feet (30%) and to both this slip and the tendon of flexor
longus tendon and a slip from flexor hallucis longus in four digitorum longus in the 14 remaining cases (70%). Edama
limbs (9.5%), into a slip from flexor hallucis longus in one et al. (2019) studied quadratus plantae in 116 legs from 62
limb (2.4%), and into the digital tendons of flexor digitorum Japanese cadavers. Quadratus plantae was comprised of
longus in two limbs (4.8%). Out of the 47 limbs in which it both the medial and lateral heads in 101 legs (87.1%). In 11
was present, the lateral head inserted into the flexor digito- cases (9.5%) the lateral head was absent, and in four cases
rum longus tendon in 44 limbs (93.6%) and into the flexor (3.4%) the medial head was absent.
digitorum longus tendon and a slip from flexor hallucis lon- Out of ten sides from five fetuses, Elvan et al. (2019)
gus in three limbs (6.4%). found that quadratus plantae contributed to the long flexor
Pretterklieber (2017) examined the contributions of flexor tendon going to digit two on one side (10%), to the long
digitorum longus, flexor hallucis longus, and quadratus plan- flexor tendon going to the third and fourth digits on all ten
tae to the long flexor tendons in 100 cases. Quadratus plantae sides (100%), and to the long flexor tendon going to the fifth
contributed to the first tendon in 14 cases (14%), to the sec- digit on four sides (40%).
ond tendon in 97 cases (97%), to the third tendon in 98 cases
(98%), to the fourth tendon in 99 cases (99%), and to the fifth Anomalies
tendon in 51 cases (51%). Pretterklieber (2018) studied qua- Description
dratus plantae in 100 feet from 50 cadavers. The muscle had Pettersen (1979) found that quadratus plantae was rudimen-
one head in 34 cases (34%), two heads in 57 cases (57%), tary in both feet of a boy with trisomy 13q. In both feet, the
and three heads in 9 cases (9%). In the nine cases where the lateral head was absent. In the left foot, the medial head was
muscle had three heads, these middle heads originated from diminutive and originated from the posteromedial aspect of
the plantar or medial calcaneal surface in five cases (56%), the calcaneus near the insertion of the calcaneal tendon. The
the long plantar ligament in five cases (56%), the medial plan- muscle did not have fleshy fibers as it crossed the longitudinal
tar intermuscular septum in two cases (22%), and the plantar arch of the foot. It had a “weblike” attachment onto the deep
calcaneocuboid ligament in one case (11%). surface of flexor digitorum brevis, into the part of the flexor
The medial head was absent in one foot (1%). Out of the hallucis longus tendon that unites with flexor digitorum lon-
99 cases in which it was present, the medial head had origins gus, and into the deep plantar fascia and ligaments of the
from the medial calcaneal surface in all cases (100%), the foot. In the right foot, the medial head had a similar origin
long plantar ligament in 92 cases (93%) the plantar calca- and was also diminutive but sent fleshy fibers across the foot
neocuboid ligament in 79 cases (80%), the plantar calcaneal that ended with a tendinous attachment into flexor digitorum
surface in 62 cases (63%), the medial calcaneal process in 12 longus. Hootnick et al. (1987) describe an individual that had
cases (12%), the tendon of tibialis posterior in 8 cases (8%), a right limb with congenital tibial aplasia, talocalcaneal syn-
the sheath of the flexor hallucis longus tendon in 8 cases (8%), chondrosis, and an adducted foot with five toes. In this limb,
the medial plantar intermuscular septum in 5 cases (5%), the quadratus plantae originated from the calcaneus and inserted
flexor retinaculum in 2 cases (2%), the lateral plantar inter- into the common flexor tendon sheet in the foot.
muscular septum in 2 cases (2%), the connection between
quadratus plantae and abductor hallucis in 1 case (1%), and Prevalence
from abductor hallucis in 1 case (1%) (Pretterklieber 2018).
The lateral head was absent in 31 feet (31%). Out of the
absence of the lateral head of quadratus plantae was only
69 cases in which it was present, the lateral head had ori-
observed in 1 out of 20 individuals with trisomy 13 (5%).
gins from the long plantar ligament in 62 cases (90%), the
lateral plantar process in 44 cases (64%), the plantar surface Clinical Implications
of the calcaneus in 11 cases (16%), the lateral surface of the
Athavale et al. (2012a) suggest that a bulky medial head of
calcaneus in eight cases (12%), the plantar calcaneocuboid
quadratus plantae may cause tarsal tunnel syndrome.
ligament in two cases (3%), the lateral plantar intermuscu-
lar septum in two cases (3%), and the medial plantar inter-
muscular septum in one case (1%) (Pretterklieber 2018). Lumbricales (Lumbricals) (Figure 5.11)
Pretterklieber (2018) was able to identify the insertions
of quadratus plantae onto the long flexor tendons in 44 Synonyms
cases. Quadratus plantae contributed to the first tendon in A singular lumbrical muscle may be referred to as lumbri-
two cases (5%), the second tendon in 43 cases (98%), the calis, and multiple lumbrical muscles may be referred to as
third tendon in 44 cases (100%), the fourth tendon in 42 lumbricales or lumbricales pedis (Wood 1868; Macalister
cases (96%), and the fifth tendon in 19 cases (43%). 1875).
Beger et al. (2018b) studied the connections of the master
knot of Henry in 20 feet from ten cadavers. In 4 feet (20%), Typical Presentation
the medial head of quadratus plantae was either mostly ten- Description
dinous (one case) or half tendinous (three cases). Quadratus The four lumbrical muscles originate from the digital ten-
plantae attached to the slip of flexor hallucis longus only in dons of flexor digitorum longus at the points where they
separate from the common flexor digitorum longus tendon The lumbrical muscles may send accessory tendons to the
(Standring 2016). The first lumbrical muscle originates proximal phalanges of their target digit (Oukouchi et al.
from the tendon going to digit two and goes to digit two 1992). Oukouchi et al. (1992) suggest that the insertion onto
(Standring 2016). The second lumbrical originates from proximal phalanges and bifurcated insertions onto two adja-
the tendons going to digits two and three and goes to digit cent digits may be atavistic presentations of the lumbricals.
three (Standring 2016). The third lumbrical originates from
the tendons going to digits three and four and goes to digit Prevalence
four (Standring 2016). The fourth lumbrical originates from In a sample of 102 subjects, Wood (1868) found that the
the tendons going to digits four and five and goes to digit lumbricals only showed variations in seven subjects (6.9%).
five (Standring 2016). Each lumbrical inserts into the dorsal The second lumbrical was absent bilaterally in one subject.
digital expansion on the proximal phalanx of its target digit The third lumbrical originated from flexor digitorum brevis
(Standring 2016). in one subject and was doubled bilaterally in another sub-
ject. The fourth lumbrical muscle was absent in three sub-
Innervation jects (one bilateral case, two unilateral cases) and doubled
The first lumbrical is innervated by the medial plantar bilaterally in another subject and had bifurcated insertions
nerve and the others are innervated by the lateral plantar that attached onto both digits four and five.
nerve (Standring 2016). Oukouchi et al. (1992) studied the lumbrical muscles
in 25 feet from Japanese cadavers. Accessory slips to the
proximal phalanges of the corresponding digit were sent by
Comparative Anatomy the first lumbrical in three cases (12%), the second lum-
The lumbrical muscles have similar typical presentations brical in two cases (8%), the third lumbrical in three cases
in the apes, with origins from the tendons of flexor digito- (12%), and the fourth lumbrical in four cases (16%). In one
rum longus and flexor hallucis longus and insertions onto foot (4%), the third and fourth lumbricals were split at their
the medial aspects of the dorsal digital expansions of digits insertions, with the former sending slips to digits three and
two through five (Chapman 1880; Hepburn 1892; Beddard four and the latter sending slips to digits four and five.
1893; Dwight 1895; Sonntag 1924; Raven 1950; Miller In a sample of 257 cadaveric feet, Chaney et al. (1996)
1952; Gibbs 1999; Diogo et al. 2010, 2012, 2013a,b, 2017). found that all the lumbricals were absent in 8 cases (3.1%).
As in humans, the lumbricals may be missing in some One lumbrical was present in 9 cases (3.5%), two lumbri-
cases (Beddard 1893) or may have double origins from the cals were present in 31 cases (12.1%), three lumbricals were
flexor digitorum longus and flexor hallucis longus tendons present in 70 cases (27.2%), and all four lumbricals were
(Champneys 1872; Chapman 1880; Hepburn 1892; Beddard present in 139 cases (54.1%). In a sample of 11 feet from
1893; Dwight 1895; Sonntag 1924; Gibbs 1999; Diogo et al. eight cadavers, Kura et al. (1997) found that the third and
2012, 2013a,b). fourth lumbrical muscles were absent in 1 foot (9.1%).
In a sample of 47 lower limbs, Athavale et al. (2012a)
Variations found that slips of flexor hallucis longus gave rise to the first
Description lumbrical muscle in 18 cases (38.3%) and to other lumbrical
One or more of the lumbrical muscles may be absent muscles in 9 cases (19.15%). In a sample of 66 cadaveric
(Wood 1868; Macalister 1875; Knott 1883b; Ochiltree specimens, Hur et al. (2015b) found that the first lumbrical
1912; Bergman et al. 1988; Chaney et al. 1996; Kura et al. had two bellies with origins from the digit two tendon of
1997; Kafka et al. 2016). Any one of the lumbricals may flexor digitorum longus and a slip of flexor hallucis longus
originate with two bellies (i.e., are doubled) instead of in 55 specimens (83.3%). It originated as a single belly from
having a unified origin from one or more of the flexor ten- the digit two tendon of flexor digitorum longus only in one
dons (Wood 1868; Macalister 1875; Bergman et al. 1988; case (1.5%) and as a single belly from the tendinous slip
Hur et al. 2015b). The lumbricals may receive contribu- of flexor hallucis longus only in the remaining ten cases
tions from flexor digitorum longus, flexor hallucis lon- (15.2%). The second lumbrical originated from the digit
gus, and/or quadratus plantae (Macalister 1875; Bergman three tendon of flexor digitorum longus and from tendinous
et al. 1988; Hur et al. 2015b). The lumbricals may also slips of flexor hallucis longus going to digits two and three
lack origins from the flexor digitorum longus tendons and in all specimens (100%). The third lumbrical originated
instead originate from the tendon(s) of flexor digitorum from tendinous slips of flexor hallucis longus going to dig-
brevis, tibialis posterior (in the case of the first lumbri- its three and four in 46 specimens (69.7%). The fourth lum-
cal), and/or flexor hallucis longus (Wood 1868; Macalister brical originated from the tendinous slip of flexor hallucis
1875; Bergman et al. 1988; Athavale et al. 2012a; Hur et al. longus going to digit four in 12 specimens (18.2%). Deep
2015b; Kafka et al. 2016). muscle fibers of the fourth lumbrical originated from the
The third or fourth lumbricals may be bifurcated at their tendinous slip of flexor hallucis longus going to digit two in
insertions, particularly when these muscles are doubled, three specimens (4.5%), from the slip going to digit three in
and attach onto the contiguous sides of the corresponding 19 specimens (28.8%), and from the slip going to digit four
digits (Wood 1868; Macalister 1875; Oukouchi et al. 1992). in 10 specimens (15.2%).

Mieden (1982) describes an infant with median cleft lip, Flexor hallucis brevis arises from a bifurcated tendon,
hypotelorism, and alobar holoprosencephaly. The fourth with the lateral part originating from the cuboid and the
lumbrical muscle was absent on the left side (Mieden 1982). lateral cuneiform and the medial part originating from the
The author does not specify if it was a hand or foot lum- tibialis posterior tendon and the medial intermuscular sep-
brical. On the left side of a child with trisomy 21, there tum (Standring 2016). The muscle splits into medial and
were double lumbrical muscles between digits four and five lateral portions that each have a tendinous attachment onto
(Bersu 1980). In the right foot of the fetus with craniora- their respective side of the base of the proximal hallucal
chischisis dissected by Alghamdi et al. (2018), the first lum- phalanx (Standring 2016). Upon insertion, the medial por-
brical muscle was absent. Hootnick et al. (1987) describe an tion joins with the abductor hallucis tendon and the lateral
individual that had a right limb with congenital tibial apla- portion joins with the adductor hallucis tendon (Standring
sia, talocalcaneal synchondrosis, and an adducted foot with 2016).
five toes. In this limb, the four lumbricals partially arose
from the common flexor tendon sheet found in the foot. Innervation
Flexor hallucis brevis is innervated by the medial plantar
Prevalence nerve (Standring 2016).
Bersu (1980) found double lumbrical muscles in one out of
five individuals with trisomy 21 (20%). Comparative Anatomy
Flexor hallucis brevis has a similar typical presentation in
Clinical Implications the apes. It has two heads that extend from the medial and/
Chaney et al. (1996) note that the absence of a lumbrical or lateral cuneiform, the first metatarsal, and sometimes
can result in claw toe or a sagittal plane deformity of a digit the tibialis posterior tendon to the proximal hallucal pha-
that can lead to calluses. lanx, often fusing with abductor hallucis, adductor hallucis,
or opponens hallucis and exhibiting additional insertions
into the first metatarsal (Champneys 1872; Brooks 1887;
Flexor hallucis brevis (Figure 5.12)
Hepburn 1892; Beddard 1893; Dwight 1895; Sonntag 1924;
Synonyms Raven 1950; Miller 1952; Lewis 1964; Gibbs 1999; Vereecke
This muscle may also be referred to as flexor brevis hallucis et al. 2005; Diogo et al. 2010, 2012, 2013a,b, 2017). It may
(Macalister 1875). have an origin from the calcaneus, first tarsometatarsal
FIGURE 5.12 Third layer of plantar foot muscles in plantar (inferior) view. Contrahentes pedis and flexor hallucis brevis are illus-
trated on the right side, and the other muscles are illustrated on the left side.
joint, or the navicular in gorillas (Gibbs 1999; Diogo et al. pedis, transversus pedis, transversus plantae, abducteur
2010). It may have an origin from the cuboid in common transverse, or adductor transversus hallucis (Knott 1883b).
chimpanzees (Beddard 1893).
Description The transverse head of adductor hallucis originates from
The lateral head of flexor hallucis brevis may be fused the plantar metatarsophalangeal ligaments of digits three,
with adductor hallucis, and the medial head may be fused four, and five and from the deep transverse metatarsal liga-
with abductor hallucis (Macalister 1875; Le Double 1897; ments between these digits (Standring 2016). The oblique
Bergman et al. 1988; Kafka et al. 2016). It may receive head originates from the bases of metatarsals two, three,
accessory slips from the calcaneus, medial cuneiform, and four and from the sheath of the fibularis longus tendon
long plantar ligament, the tendon of flexor digitorum lon- (Standring 2016). The medial portion of the oblique head
gus, or the tendon of tibialis posterior (Wood 1867b, 1868; joins with flexor hallucis brevis and inserts onto the lateral
Macalister 1875; Le Double 1897; Bergman et al. 1988; sesamoid bone of the hallux (Standring 2016). The lateral
Kafka et al. 2016; Standring 2016). It may send a slip to portion of the oblique head merges with the transverse head
the proximal phalanx of digit two (Wood 1867b; Macalister and inserts onto the lateral sesamoid bone and onto the base
1875; Le Double 1897; Bergman et al. 1988; Kafka et al. of the proximal hallucal phalanx (Standring 2016).
2016; Standring 2016).
Innervation
Prevalence Adductor hallucis is innervated by the lateral plantar nerve

Description Adductor hallucis has a similar typical presentation in the
Pettersen et al. (1979) found that flexor hallucis brevis had apes. In most cases, this muscle has (1) an oblique head
anomalous presentations in one neonate and one fetus with that originates from the bases of the second, third, and
trisomy 13. In the neonate, the left flexor hallucis brevis sometimes fourth metatarsals; (2) a transverse head that
had an accessory head of origin from the medial cuneiform. originates from the distal plantar surfaces of metatarsals
The other “normal” heads of this muscle originated from two through four and their corresponding metatarsophalan-
the base of the first metatarsal. In the fetus, the muscle on geal joints and ligaments; and (3) insertions onto the first
both sides of the body originated from the first metatarsal metatarsal, the base of the proximal hallucal phalanx, and
distal to its base. sometimes the lateral hallucal sesamoid bone (Champneys
On the right side of the fetus with craniorachischisis 1872; Brooks 1887; Hepburn 1892; Beddard 1893; Dwight
dissected by Alghamdi et al. (2017), flexor hallucis brevis 1895; Sonntag 1924; Raven 1950; Miller 1952; Lewis 1964;
was absent. On the left side of the specimen, flexor hallucis Gibbs 1999; Vereecke et al. 2005; Diogo et al. 2010, 2012,
brevis extended from the lateral cuneiform and base of the 2013a,b, 2017).
first metatarsal to the base of the proximal hallucal phalanx The two heads are always fused in gibbons (Brooks
and sent fibers to the tendon of abductor hallucis. In the 1887) and variably united in common chimpanzees (Gibbs
female fetus with trisomy 18 dissected by Alghamdi et al. 1999). The oblique head in gorillas and bonobos does not
(2018), the distal end of abductor hallucis was fused with have an origin from the base of the fourth metatarsal
the medial head of flexor hallucis brevis bilaterally. (Miller 19552; Vereecke et al. 2005; Diogo et al. 2010,
2017). Attachments to the fourth metatarsal are absent
Prevalence entirely in orangutans (Beddard 1893; Brooks 1887;
In their literature review, Smith et al. (2015) found that Gibbs 1999) and in some bonobos (Diogo et al. 2017).
accessory slips of flexor hallucis brevis were only present in The oblique head in orangutans may have an origin from
1 of 20 individuals with trisomy 13 (5%). the sheath of the fibularis longus tendon (Beddard 1893;
Brooks 1887; Sonntag 1924).
N/A Variations
Description
Adductor hallucis (Figure 5.12) The central component of the transverse head may be insep-
arable from the medial and lateral components (Kura et al.
Synonyms 1997). The origin of the transverse head from the fifth digit
The transverse head of adductor hallucis was considered a may be absent (Macalister 1875; Le Double 1897; Bergman
separate muscle in earlier texts and may be referred to as et al. 1988; Arakawa et al. 2003; Standring 2016). Macalister
transversalis pedis Casserii (Macalister 1875), transversalis (1875) and Knott (1883b) note that the attachments to the
third and fourth digits may also be absent. More often, the Arakawa et al. (2003) also identified the common ori-
entire transverse head of adductor hallucis may be absent gin sites of the transverse head as the capsules of the third,
(Macalister 1875; Knott 1883b; Cralley and Schuberth fourth, and fifth metatarsophalangeal joints and the deep
1979; Kura et al. 1997; Kafka et al. 2016; Standring 2016). transverse metatarsal ligaments between them. These
The oblique head of adductor hallucis may originate authors classified the transverse head into three types. Type
entirely from the sheath of the fibularis longus tendon A was described as the narrow type and had a confined
(Macalister 1875). The oblique head of adductor hallucis range of origin, only originating from the third, fourth, and
may send a slip to the base of the proximal phalanx of digit fifth metatarsophalangeal joints and the deep transverse
two (Wood 1867b; Macalister 1875; Knott 1883b; Arakawa metatarsal ligaments between them. Type B was described
et al. 2003; Kafka et al. 2016; Standring 2016) or more as the lateral type and originated from the common ori-
rarely digit three (Ochiltree 1912; Bergman et al. 1988). The gin sites only. Type C was described as the wide type and
oblique head may receive a slip from the tendon of tibialis originated from the common origin sites, the aponeurosis
posterior (Gunal et al. 1994). between the third plantar and the fourth dorsal interosse-
Chouke (1927) uses “adductor hallucis accessorius” to ous muscles, and the deep band of the fibular portion of the
refer to a trigastric supernumerary muscle that has some plantar aponeurosis.
similarities with the accessory long flexors of the calf.
Adductor hallucis accessorius originated as a fleshy belly Prevalence
(first belly) from the medial margin of the tibia below the Knott (1883b) found that the attachment of “transversus
origin of flexor digitorum longus, about three inches above pedis” to the heads of the third and fourth metatarsals
the medial malleolus. It also had an origin from the deep was absent in 5 out of 34 cases (14.7%). In a sample of
fascia of the leg. The muscle coursed superficial to the tibial 91 cadaveric feet, Cralley and Schuberth (1979) found that
nerve and became tendinous at the level of the medial malle- the transverse head of adductor hallucis was absent in 6%
olus. The tendon passed under the flexor retinaculum medial of cases. In a sample of 11 feet from eight cadavers, Kura
to the synovial sheath of the flexor hallucis longus tendon et al. (1997) found that the transverse head of adductor
and posterior to the tibial nerve. The tendon then received a hallucis was absent in 1 foot (9.1%). In 10 feet (90.9%),
fleshy bundle (second belly) from the posterior portion of the the central component of the transverse head of adductor
medial surface of the calcaneus that originated from under hallucis could not be separated from the medial or lateral
the origin of abductor hallucis. In the sole of the foot, adduc- components.
tor hallucis accessorius became tendinous again and passed Arakawa et al. (2003) studied adductor hallucis in 45
to the plantar surface of the medial cuneiform. At this point, feet from 34 cadavers. These authors classified the oblique
it became fleshy again (third belly) and joined with the head into four types (see above for descriptions). Type A
oblique head of adductor hallucis just before its insertion. was found in 21 cases (46.7%), type B was found in 15 cases
Arakawa et al. (2003) identified the common origin sites (33.3%), type C was found in four cases (8.8%), and type D
of the oblique head as the sheath of the fibularis longus ten- was found in five cases (11.1%). These authors could only
don, the long plantar ligament, the bases of metatarsals two identify the origins and insertions of the transverse head
through four and the plantar metatarsal ligaments between in 34 specimens and classified this head into three types
them, and the lateral cuneiform. These authors then classi- (see above for descriptions). Type A was found in 14 cases
fied the oblique head into four types. Type A was described (41.2%), type B was found in 10 cases (29.4%), and type C
as the narrow type and had a confined range of origin. This was found in 10 cases (29.4%).
type was divided into three subtypes. In subtype 1, all the
common origins were present. In subtype 2, the origin from Anomalies
the long plantar ligament was missing. In subtype 3, the Description
origin from the lateral cuneiform was missing. Type B was In a study of individuals with trisomy 13, Pettersen et al.
described as the lateral type and exhibited all the common (1979) noted that the transverse head of adductor hallucis
origin sites as well as an origin via an aponeurotic slip from was absent on the left side of one neonate and bilaterally
the base of the fifth metatarsal. Type C was described as the in one fetus. The muscle was hypoplastic bilaterally in a
wide type and had broad origins. In addition to the com- second neonate and on the right side of a second fetus. On
mon origin sites, type C oblique heads originated laterally the left side of this second fetus, it originated from the deep
from a slip from the base of the fifth metatarsal and had surface of the flexor digitorum longus tendon to digit five
possible medial origins from the tibialis posterior tendon, but had a typical insertion.
medial intermuscular septum, plantar tarsometatarsal liga- On the right foot of a male neonate with Meckel syn-
ment between the medial cuneiform and the base of the sec- drome, Pettersen (1984) observed that the transverse head of
ond metatarsal, or the fibularis longus tendon inserting into adductor hallucis had a partial origin from the flexor digito-
the first dorsal interosseous muscle. Type D was described rum brevis to the fifth digit. On the left foot, the transverse
as the medial type and exhibited all the common origin head was joined at its origin to flexor digiti minimi brevis
sites and had potential medial origins from the medial sites and to the interosseous muscles of digit four. Hootnick et al.
found in Type C. (1987) describe an individual that had a right limb with
congenital tibial aplasia, talocalcaneal synchondrosis, and Variations

an adducted foot with five toes. In this limb, the transverse Description
head of adductor hallucis was absent.
Contrahentes of the foot (contrahentes 3, 4, 5) are typi-
The transverse head of adductor hallucis was absent
cally only present in the early stages of human devel-
in the right foot of the fetus with trisomy 18 and cyclopia
opment and then disappear completely later in ontogeny
described by Smith et al. (2015). On the right side of the
(Cihak 1972; Diogo et al 2019). Adductor hallucis derives
fetus with craniorachischisis dissected by Alghamdi et al.
from contrahentes 1 and 2 (Cihak 1972; Arakawa et al.
(2017), the transverse head of adductor hallucis was absent.
2003).
The oblique head originated from the medial cuneiform and
The oblique head develops from the tibial part of the con-
had a typical insertion onto the proximal phalanx of digit
trahentes layer at CR18 mm (crown-rump length of 18 mm),
one. On the left foot, the oblique head originated from the
and the transverse head develops from the fibular part of
bases of the third and fourth metatarsals and the transverse
the contrahentes layer at CR30 mm (crown-rump length
head originated from the distal ends of these metatarsals.
of 30 mm) (Cihak 1972; Diogo et al 2019). Contrahentes 3,
The heads inserted together onto the base of the proximal
4, and 5 persist in the embryo until CR25–35 mm (crown-
phalanx of digit one.
rump length of 25 to 35 mm) and then become completely
Prevalence lost (Cihak 1972, Diogo et al. 2019).
Contrahentes are very rare in karyotypically normal
humans. Hirsch and Vekkos (1984) found contrahentes
absence of the transverse head of adductor hallucis was
muscles in 2 feet. In both cases, the contrahentes muscles
noted in 2 out of 20 individuals (10%) with trisomy 13.
occupied the approximate position of the oblique head of
adductor hallucis but were larger and had more extensive
attachments. The transverse head of adductor hallucis
Expansion of the tibialis posterior tendon into the oblique was absent in both feet. In one case, the muscle inserted
head of adductor hallucis is associated with hallux valgus onto the metatarsophalangeal joint capsules of digits one
(Gunal et al. 1994). and two. In the second case, the muscle inserted onto the
metatarsophalangeal joint capsules of digits two through
Contrahentes pedis (Figure 5.12) five.
Entry adapted by permission from Springer Nature Customer Innervation
Service Centre GmbH: Springer Current Molecular Biology
Contrahentes pedis are innervated by the lateral plantar
Reports, Muscles Lost in Our Adult Primate Ancestors
nerve (Miller 1952; Hirch and Vekkos 1984).
Still Imprint in US: on Muscle Evolution, Development,
Variations, and Pathologies. E. Boyle, V. Mahon, R. Diogo, Prevalence
2020.
N/A
See also: Adductor hallucis
Anomalies
Synonyms
N/A
Contrahens refers to a singular muscle, and contrahentes
refer to multiple muscles (Cihak 1972).
Opponens hallucis (Figure 5.12)
Comparative Anatomy
Descriptions of contrahentes of the feet are often neglected Entry adapted by permission from Springer Nature Customer
in descriptions of ape anatomy. In gibbons, Hirasaki and Service Centre GmbH: Springer Current Molecular Biology
Kumakura (2010) note the presence of a contrahens muscle Reports, Muscles Lost in Our Adult Primate Ancestors
originating from the sheath of the fibularis longus tendon Still Imprint in US: on Muscle Evolution, Development,
and the third metatarsal and inserting onto the tibial side Variations, and Pathologies. E. Boyle, V. Mahon, R. Diogo,
of the proximal phalanx of digit five. In bonobos, the con- 2020.
trahens muscle extending from the contrahens raphe to See also: Adductor hallucis, Flexor hallucis brevis
digit five is often present and well-developed while that to
digit four is often thin, reduced, or absent (Hepburn 1892; Synonyms
Miller 1952; Lewis 1989; Diogo et al. 2017). Contrahentes N/A
with variable attachments to digits two, four, and five are
common in lemuriform primates and some monkeys (Gebo Typical Presentation
1985; Hirasaki and Kumakura 2010). This muscle is only present as a variation.
Comparative Anatomy (Standring 2016). Its tendon blends with that of abductor
Among the apes, opponens hallucis has been described in digiti minimi to insert onto the base of the proximal pha-
detail only in orangutans (Chapman 1880; Brooks 1887; lanx of digit five (Standring 2016).
Primrose 1899; Boyer 1935; Diogo et al. 2013b) and seems
to be present in some common chimpanzees and bonobos Innervation
(Chapman 1879; Champneys 1872; Diogo et al. 2013a, 2017). Flexor digiti minimi brevis is innervated by the lateral plan-
Though it has been described in one gibbon, the presence tar nerve (Standring 2016).
of this muscle in gibbons has not been confirmed by other
Comparative Anatomy
dissections (Diogo et al. 2012). When present in orangutans,
common chimpanzees, and bonobos, the muscle originates Flexor digiti minimi brevis has a similar typical presentation
from the medial cuneiform and inserts onto the first metatar- in the apes, originating from the base of the fifth metatarsal
sal (Brooks 1887; Primrose 1899; Diogo et al. 2013a,b, 2017). and/or the plantar tarsometatarsal ligament extending to the
In orangutans, Brooks (1887) notes an origin from a carti- tuberosity of this metatarsal and inserting onto the proxi-
laginous nodule in the tibialis posterior tendon, and Boyer mal phalanx of digit five (Champneys 1872; Sonntag 1924;
(1935) notes an insertion into the base of the proximal hallu- Boyer 1935; Raven 1950; Miller 1952; Gibbs 1999; Vereecke
cal phalanx. It is also often fused with flexor hallucis brevis et al. 2005; Ferrero 2011; Ferrero et al. 2012; Diogo et al.
in orangutans (Brooks 1887; Primrose 1889; Boyer 1935). 2010, 2012, 2013a,b, 2017). There may be an origin from the
fourth metatarsal in gorillas (Raven 1950; Gibbs 1999) and
Variations from the sheath of fibularis longus in orangutans (Sonntag
Description 1924; Gibbs 1999). It may be absent in some common chim-
panzees (Champneys 1872; Diogo et al. 2013a).
Opponens hallucis has typically been described as an acces-
sory slip of adductor hallucis that inserts onto the first meta- Variations
tarsal (Macalister 1875; Knott 1883b; Playfair McMurrich
Description
1906; Standring 2016). Kafka et al. (2016) state that oppo-
nens hallucis describes a slip originating from either the Flexor digiti minimi brevis may be fused with abductor
oblique or transverse head of adductor hallucis that inserts digiti minimi (Macalister 1875; Kopuz et al. 1999). It may
onto the proximal hallucal phalanx. Bergman et al. (1988) also be fused with the tendon of abductor ossis metatarsi
state that opponens hallucis is an accessory slip of flexor digiti quinti (Macalister 1875). It may originate entirely
hallucis brevis that inserts onto the first metatarsal. from the sheath of fibularis longus (Macalister 1875). Jana
and Roy (2011) note an unusual case in which flexor digiti
Innervation minimi brevis arose from the lateral plantar process of the
In orangutans, opponens hallucis is innervated by the calcaneus with abductor digiti minimi and then passed onto
medial plantar nerve (Brooks 1887). the lateral side of the fifth metatarsal. The tendon of flexor
digiti minimi brevis received the lateral slip of the fibularis
Prevalence tertius insertion at the base of the fifth metatarsal (Jana and
No prevalence information is available, but Macalister Roy 2011). The insertion of flexor digiti minimi brevis onto
(1875, p. 129) describes it as a “frequent constituent element the proximal phalanx may be separate from the insertion
in the human foot.” of abductor digiti minimi (Macalister 1875; Knott 1883b).
A deep fascicle of flexor digiti minimi brevis that inserts
Anomalies onto the fifth metatarsal may be referred to as opponens
N/A digiti minimi (see the entry for this muscle) (Wood 1864,
Macalister 1875; Playfair McMurrich ‎ 1906; Bergman et al.
Clinical Implications 1988; Kafka et al. 2016; Standring 2016).
N/A
Prevalence
Flexor digiti minimi brevis (Figure 5.12) N/A
Synonyms Anomalies
It may also be referred to as flexor brevis minimi digiti Description
(Macalister 1875) or flexor brevis quinti digiti (Playfair On the right foot of a male neonate with Meckel syn-
McMurrich 1906). drome, Pettersen (1984) found that flexor digiti minimi
brevis attached to the fifth digit and received a slip from
flexor digitorum brevis that did not split to accommodate
Description the tendon of flexor digitorum longus. On the left foot,
Flexor digiti minimi brevis originates from the base of the this muscle was fused with the origin of adductor hallu-
fifth metatarsal and from the sheath of fibularis longus cis and was represented by a mass (along with abductor
digiti minimi) on the plantar and lateral sides of digit six Diogo et al. (2019) found that opponens digiti minimi is
with vague attachments to this digit and partially to digit well-differentiated from flexor digiti minimi brevis at CR
five. Hootnick et al. (1987) describe an individual that had 36mm and persists until at least CR51 mm, or gestational
a right limb with congenital tibial aplasia, talocalcaneal week 11.
synchondrosis, and an adducted foot with five toes. In this When present in adults, this muscle is a distinct, deep
limb, flexor digiti minimi brevis originated from the cal- fascicle of flexor digiti minimi brevis that originates from
caneus abnormally deep in the foot and passed deep to the base of the fifth metatarsal and inserts along the lat-
quadratus plantae. eral aspect of the shaft of the fifth metatarsal (Wood 1864,
Macalister 1875; Playfair McMurrich 1906; Bregman et al.
Prevalence 1988; Kafka et al. 2016; Standring 2016). Another presenta-
N/A tion in an adult male cadaver was noted by Rana and Das
(2006), where this muscle had the same origin but inserted
Clinical Implications onto the distal phalanx of the fifth digit. It may be present
N/A bilaterally (Wood 1868; Rana and Das 2006). Opponens
digiti minimi may receive a slip from the fibularis longus
tendon (Edama et al. 2020).
Opponens digiti minimi (Figure 5.12)
Entry adapted by permission from Springer Nature Customer Innervation
Service Centre GmbH: Springer Current Molecular Biology Opponens digiti minimi is innervated by the lateral plantar
Reports, Muscles Lost in Our Adult Primate Ancestors nerve (Gibbs 1999; Rana and Das 2006).
Still Imprint in US: on Muscle Evolution, Development,
Variations, and Pathologies. E. Boyle, V. Mahon, R. Diogo, Prevalence
2020. Wood (1868) found opponens digiti minimi in 6 out of 36
subjects (16.7%).
Synonyms
This muscle is also referred to as opponens quinti digiti Anomalies
(Playfair McMurrich ‎ 1906) or opponens minimi digiti N/A
(Macalister 1875).
Interossei plantares (Plantar Interossei) (Figure 5.11)
Comparative Anatomy
While typically absent in adult humans, opponens digiti Synonyms
minimi is present in adult apes (Gibbs 1999; Diogo et al. N/A
2010, 2012, 2013a,b, 2017). In most cases, opponens digiti
minimi has variable connections to flexor digiti minimi Typical Presentation
brevis, originates from the base of the fifth metatarsal, and Description
inserts onto to the lateral border of the fifth metatarsal and/ The plantar interossei are comprised of three muscles that
or the proximal phalanx of digit five (Primrose 1899; Raven are situated beneath the three lateral metatarsals (Standring
1950; Miller 1952; Gibbs 1999; Diogo et al. 2010, 2012, 2016). Each muscle originates from the base and medial
2013a,b, 2017). In orangutans and bonobos, this muscle side of one metatarsal and inserts onto the dorsal digital
has an origin from the sheath of the fibularis longus tendon expansion and base of the proximal phalanx of its corre-
(Miller 1952; Gibbs 1999; Ferrero 2011; Ferrero et al. 2012; sponding digit (Standring 2016). The first plantar interosse-
Diogo et al. 2013b, 2017). ous muscle thus originates and inserts onto digit three, the
second plantar interosseous muscle originates and inserts
Variations
onto digit four, and the third plantar interosseous muscle
Description originates and inserts onto digit five (Standring 2016).
Opponens digiti minimi is typically only present in the
early stages of human development, and then disappears Innervation
later during gestation. Bardeen (1906) states that the blas- The plantar interossei are innervated by the lateral plantar
tema of opponens digiti minimi and flexor digiti minimi nerve (Standring 2016).
brevis appears at CR12 mm, and then the muscles differ-
entiate at CR18 mm. Cihak (1972) suggests that this muscle Comparative Anatomy
appears at CR20 mm (crown-rump length of 20 mm) and While the functional axis of the foot is digit two in humans,
differentiates from flexor digiti minimi brevis at CR40 mm. in the apes, the digit around which the interosseous muscles
are arranged is typically digit three (Dwight 1895; Boyer onto digits three and four in one case (0.7%). The tendon of
1935; Sonntag 1924; Raven 1950; Miller 1952; Gibbs 1999; the third plantar interosseous split to insert onto digits four
Diogo et al. 2010, 2012, 2013a,b, 2017; Hirasaki and Oishi and five in two cases (1.4%).
2018). In most apes, the plantar interossei are attached along Edama et al. (2020) studied the insertion of fibularis
the lateral side of digit two and the medial sides of digits four longus in 104 feet from 52 Japanese cadavers. The ante-
and five (Dwight 1895; Boyer 1935; Sonntag 1924; Raven rior frenular ligament was present in 33 feet (31.7%), and
1950; Miller 1952; Gibbs 1999; Diogo et al. 2010, 2012, it attached to the second plantar interosseous muscle in 13
2013a,b, 2017; Hirasaki and Oishi 2018). Accessory attach- feet (12.5%) and the third plantar interosseous muscle in 29
ments to metatarsals one and three have been observed in feet (27.9%).
gorillas and orangutans (Raven 1950; Gibbs 1999; Ferrero
2011; Ferrero et al. 2012; Diogo et al. 2010, 2013b). Two Anomalies
accessory plantar interossei attached to the medial and lat- N/A
eral sides of the third metatarsal were observed by Dwight
(1895) in common chimpanzees. Hirasaki and Oishi (2018) Clinical Implications
found that the interossei exhibited the human presentation N/A
and were thus arranged around digit two in two out of ten
chimpanzees (20%), one out of three gorillas (33.3%), and
Interossei dorsales (Dorsal Interossei) (Figure 5.13)
in one bonobo.
Synonyms
Variations N/A
Description
The second and third plantar interosseous muscles may Typical Presentation
receive a slip from the tendon of fibularis longus (Edama Description
et al. 2020). The interossei may have accessory origins from The dorsal interossei are comprised of four bipennate
the plantar ligaments and/or the fascia of adjacent muscles muscles that occupy the spaces in between the metatarsals
(Kalin and Hirsch 1987). The first plantar interosseous (Standring 2016). Each muscle originates via two heads
muscle may extend into the dorsal part of the second inter- from adjacent metatarsals and inserts into the base of the
osseous space (Manter 1945). The second and third plantar proximal phalanx and dorsal digital expansion of its target
interossei may originate from the sheath of the fibularis lon- digit (Standring 2016). The first dorsal interosseous muscle
gus tendon (Ochiltree 1912). The second plantar interosseus originates from the sides of the first and second metatarsals
may take partial origin from adductor hallucis (Kalin and and inserts onto the medial aspect of digit two (Standring
Hirsch 1987). Accessory slips may be present connecting 2016). The second dorsal interosseous muscle originates
the plantar and dorsal interossei (Manter 1945). The plan- from the sides of the second and third metatarsals and
tar and dorsal interossei may also be fused (Manter 1945). inserts into the lateral aspect of digit two (Standring 2016).
The tendons of the interossei may be bifurcated upon inser- The third dorsal interosseous muscle originates from the
tion and attach to two separate digits (Hallisy 1930; Manter sides of the third and fourth metatarsals and inserts into the
1945). The insertions into the dorsal digital expansions may lateral aspect of digit three (Standring 2016). The fourth
be missing in some cases (Manter 1945; Oukouchi et al. dorsal interosseous muscle originates from the sides of
1992). the fourth and fifth metatarsals and inserts into the lateral
aspect of digit four (Standring 2016).
Prevalence
Manter (1945) studied the variations of the interosseous Innervation
muscles in 149 cadaveric feet. In 23 cases (15.4%), the ori- The dorsal interossei are innervated by the lateral plantar
gin of the first plantar interosseous muscle extended into nerve (Standring 2016).
the dorsal part of the second interosseous space. It occupied
the full length of this space in 8 feet, three-fourth the length Comparative Anatomy
of the space in 8 feet, and half the length of the space in While the functional axis of the foot is digit two in humans,
7 feet. An accessory slip originating from the third dorsal in the apes, the digit around which the interosseous mus-
interosseous muscle and joining the first plantar interosse- cles are arranged is typically digit three (Champneys 1872;
ous muscle was found in 17 cases (11.4%). The first plantar Dwight 1895; Boyer 1935; Sonntag 1924; Raven 1950;
and second dorsal interosseous muscles were fused in two Miller 1952; Gibbs 1999; Diogo et al. 2010, 2012, 2013a,b,
cases (1.4%). The second plantar and third dorsal interosse- 2017; Hirasaki and Oishi 2018). In the apes, the dorsal
ous muscles were fused in four cases (2.7%). Manter (1945) interossei are bipennate, with the exception of the first
also found that the tendon of the first plantar interosseous dorsal interosseous muscle in some gibbons (Brooks 1887;
split to insert onto digits two and three in one case (0.7%). Gibbs 1999; Diogo et al. 2012). Typically, the first dorsal
The tendon of the second plantar interosseous split to insert interosseous muscle originates from the first and second
FIGURE 5.13 Short extensors of the foot and the dorsal interossei in dorsal (superior) view of the foot.
metatarsal and inserts onto the medial side of digit two, the plantar portion of the second dorsal interosseous muscle
second originates from the second and third metatarsal and may course plantarly and extend laterally over the first
inserted onto the medial side of digit three, the third origi- plantar interosseous muscle without directly attaching to it
nates from the third and fourth metatarsals and inserts onto (Manter 1945). The second dorsal interosseous muscle may
the lateral side of digit three, and the fourth originates from be doubled (Manter 1945). The tendons of the interossei
digits four and five and inserts onto the lateral side of digit may be bifurcated upon insertion and attach to two sepa-
four (Champneys 1872; Brooks 1887; Dwight 1895; Boyer rate digits (Hallisy 1930; Manter 1945). The insertions into
1935; Sonntag 1924; Raven 1950; Miller 1952; Gibbs 1999; the dorsal digital expansions may be missing in some cases
Diogo et al. 2010, 2012, 2013a,b, 2017; Hirasaki and Oishi (Manter 1945; Oukouchi et al. 1992).
2018). In gorillas and orangutans, the first dorsal interos- The fourth interosseous muscle may originate from the
seus muscle may originate from the medial cuneiform and fusion of the fibularis tertius and fibularis brevis tendons
second metatarsal (Brooks 1887; Boyer 1935; Raven 1950; (Olewnik 2019). Fibularis tertius may send an accessory
Diogo et al. 2010, 2013b). Two accessory dorsal interos- tendon to the fascia that covers the fourth interosseus
sei on the lateral side of a common chimpanzee foot were muscle (the fourth interosseous space) (Macalister 1875;
observed by Dwight (1895). Hirasaki and Oishi (2018) Manter 1945; Joshi et al. 2006; Lambert 2016; Olewnik
found that the interossei exhibited the human presentation 2019). Fibularis longus may have insertions into the first,
and were thus arranged around digit two in two out of ten second, or fourth dorsal interossei (Kalin and Hirsch
chimpanzees (20%), one out of three gorillas (33.3%), and 1987; Edama et al. 2020). The first dorsal interosse-
in one bonobo. ous muscle may also originate from the fibularis longus
tendon (Harbeson 1933, 1938; Manter 1945; Kalin and
Variations Hirsch 1987). The dorsal interosseous muscles may also
Description receive slips or originate from fibularis brevis, extensor
Any one of the dorsal interossei may be unipennate as a digitorum brevis, or flexor hallucis brevis (Wood 1867b;
variation (Macalister 1875; Manter 1945). The interossei Macalister 1875; Manter 1945; Kalin and Hirsch 1987;
may have accessory origins from the plantar ligaments Kafka et al. 2016).
and/or the fascia of adjacent muscles (Kalin and Hirsch An accessory slip deep to adductor hallucis between the
1987). Accessory slips may be present connecting the plan- first dorsal interosseous muscle and the first metatarsal may be
tar and dorsal interossei (Manter 1945). The plantar and present (Manter 1945). A well-developed fibrous band extend-
dorsal interossei may also be fused (Manter 1945). The ing from the plantar aspect of the first dorsal interosseous to
the proximal hallucal phalanx may also be present (Manter over the fourth interosseous space in 15 limbs (16.5%). In
1945). These variations may represent the “missing” plantar two limbs (2.2%), the tendon was fused with a slip from the
interosseous muscle in humans (Manter 1945). fibularis brevis tendon, and the fourth interosseous muscle
originated from this fusion. Edama et al. (2020) studied the
Prevalence insertion of fibularis longus in 104 feet from 52 Japanese
Harbeson (1933) studied the origin of the first dorsal inter- cadavers. Insertions onto the first dorsal interosseous mus-
osseous muscle in 27 feet from 15 subjects. In 20 cases, the cle were found in 38 feet (36.5%). The anterior frenular lig-
medial head of the muscle originated via a tendinous slip ament was present in 33 feet (31.7%), and it attached to the
from the fibularis longus tendon (74.1%). Harbeson (1938) fourth dorsal interosseous muscle in 20 feet (19.3%).
provides the prevalence of this variation for two subsequent
dissections in 1934 and 1937. In 1934, 24 feet from 13 sub- Anomalies
jects were dissected and the first dorsal interosseous muscle N/A
had an origin from the fibularis longus tendon in 11 feet
(45.8%). In 1937, 24 feet from 12 subjects were dissected Clinical Implications
and the first dorsal interosseous muscle had an origin from N/A
the fibularis longus tendon in 15 feet (62.5%). The over-
all prevalence of this variation from Harbeson’s research is
therefore 46 out of 75 feet (61.3%). Extensor digitorum brevis (Figure 5.13)
Manter (1945) studied the variations of the interosseous Synonyms
muscles in 149 cadaveric feet. The first dorsal interosseous
This muscle may also be referred to as the extensor brevis
muscle originated as a single head from the second meta-
digitorum pedis (Macalister 1875).
tarsal in 17 cases (11.4%). The second dorsal interosseous
muscle originated as single head from the second metatar- Typical Presentation
sal in 11 cases (7.4%). The third dorsal interosseous muscle
Description
originated as a single head from the fourth metatarsal in
five cases (3.4%). The fourth dorsal interosseous muscle Extensor digitorum brevis originates from the superolateral
originated as a single head from the fourth metatarsal in surface of the distal calcaneus, from the interosseous talo-
two cases (1.3%) and from the fifth metatarsal in 34 cases calcaneal ligament, and from the inferior extensor retinacu-
(22.8%). lum (Standring 2016). It ends in three tendons that insert
Manter (1945) found an accessory slip originating from onto the extensor digitorum longus tendons for digits two,
the third dorsal interosseous muscle that joined the first three, and four (Standring 2016).
plantar interosseous muscle in 17 cases (11.4%). In four
Innervation
cases (2.7%), the plantar portion of the second dorsal inter-
osseous muscle extended laterally over the first plantar Extensor digitorum brevis is innervated by the deep fibular
interosseous muscle but did not directly attach to it. The nerve (Standring 2016).
first plantar and second dorsal interosseous muscles were
Comparative Anatomy
fused in two cases (1.4%). The second plantar and third dor-
sal interosseous muscles were fused in four cases (2.7%). Extensor digitorum brevis has a similar typical presenta-
The second dorsal interosseous was doubled in two cases tion in the apes, originating from the calcaneus and send-
(1.4%). The first dorsal interosseous muscle received a ten- ing tendons to the dorsal aponeurosis of digits two, three,
dinous slip of origin from the fibularis longus tendon in and four (Champneys 1872; Hepburn 1892; Beddard 1893;
about 95 feet (63.5%). An accessory slip deep to adductor Sonntag 1924; Boyer 1935; Raven 1950; Miller 1952; Lewis
hallucis between the first dorsal interosseous muscle and 1966; Gibbs 1999; Vereecke et al. 2005; Diogo et al. 2010,
the first metatarsal was present in three cases (2%). A well- 2012, 2013a,b, 2017). A tendon for digit five may be present
developed fibrous band extending from the plantar aspect in about half of gibbons (Lewis 1966) and in some common
of the first dorsal interosseous to the proximal hallucal pha- chimpanzees and bonobos (Miller 1952; Diogo et al. 2013a,
lanx was present in 30 out of 54 feet (55.6%). Manter (1945) 2017). An accessory slip attaching to digit two has been
also found that the tendon of the second dorsal interosseous found in gibbons (Diogo et al. 2012). An origin from the
split to insert onto digits two and three in five cases (3.4%). base of the third metatarsal as well as a split insertion of the
The tendon of the third dorsal interosseous split to insert digit two tendon has been observed in orangutans (Gibbs
onto digits three and four in two cases (1.4%). The tendon of 1999; Diogo et al. 2013b).
the fourth dorsal interosseous split to insert onto digits four
and five in one case (0.7%). Variations
Olewnik (2019) studied fibularis tertius in a sample of Description
106 cadaveric lower limbs. Out of the 91 limbs in which the One or more tendons, or even the entire muscle, may be
muscle was present, the tendon had a single, broad insertion absent (Flower and Murie 1867; Macalister 1875; Bergman
onto the base and shaft of the fifth metatarsal and the fascia et al. 1988; Sirasanagandla et al. 2013b; Standring 2016). The
muscle belly or tendon going to digits two or three may be digitorum tendons to digits two through four were pres-
doubled (Wood 1867b, 1868; Macalister 1875; Sirasanagandla ent. In four limbs (9.1%), the tendon to the fourth digit was
et al. 2013b; Won and Oh 2018). Sirasanagandla et al. (2013b) absent. In one limb (2.3%), there was an accessory bun-
report a case where the tendon for digit two was tripled, hav- dle present that originated from extensor hallucis brevis
ing attachments to the proximal phalanx of digit two, the and the extensor digitorum brevis belly going to digit two
extensor digitorum longus tendon for digit two, and the fas- that merged with the first dorsal interosseous muscle. The
cia over the shaft of the third metatarsal. extensor digitorum brevis tendon for digit two was doubled
Accessory bundles of extensor digitorum brevis in two cases (4.5%) and tripled in one case (2.3%). In one
may be present (Bergman et al. 1988; Kura et al. 1997; case (2.3%), the extensor digitorum brevis tendon for digit
Sirasanagandla et al. 2013b; Won and Oh 2018). There may three was doubled.
be an extra tendon of extensor digitorum brevis that joins
the tendon of extensor digitorum longus going to digit five Anomalies
(Macalister 1875; Knott 1883b; Le Double 1897; Bergman Description
et al. 1988; Chaney et al. 1996; Standring 2016). Extensor Mieden (1982) describes an infant with median cleft lip,
digitorum brevis may send slips to the dorsal interos- hypotelorism, and alobar holoprosencephaly. The tendon of
sei (Wood 1867b, 1868; Macalister 1875; Knott 1883b; extensor digitorum brevis to the second digit was absent on
Bergman et al. 1988; Sirasanagandla et al. 2013b; Standring the left side (Mieden 1982). On the left side, the tendons of
2016). The muscle bellies of extensor digitorum brevis extensor digitorum brevis and longus were attached to the
may receive slips from tendons of extensor digitorum lon- metatarsophalangeal joints (Mieden 1982). In a male infant
gus (Macalister 1875). It may also receive accessory slips with triploidy studied by Moen et al. (1984), extensor digito-
from the talus, navicular, cuboid, lateral cuneiform, and/ rum brevis was absent bilaterally. On the left foot of a male
or third metatarsal (Macalister 1875; Bergman et al. 1988; neonate with Meckel syndrome, Pettersen (1984) found that
Standring 2016). the extensor digitorum brevis tendons to digits five and six
Won and Oh (2018) report a case in which the exten- were absent and the tendon to digit two was diminutive.
sor digitorum muscle belly to digit three was doubled and On both sides of the fetus with craniorachischisis dis-
associated with an accessory muscle in the right foot of sected by Alghamdi et al. (2017), extensor digitorum brevis
a cadaver. The extra muscular head originated from the did not send a tendon to the fourth digit. In the right foot of
cuboid and then joined with the extensor digitorum brevis a fetus with trisomy 18 and cyclopia dissected by Smith et
tendon going to digit three. The accessory muscle was cov- al. (2015), the extensor digitorum brevis tendon to the sec-
ered by extensor hallucis brevis, originated from the calca- ond digit bifurcated and attached onto digits two and three.
neus, and inserted onto the lateral third of the transverse In the female fetus with trisomy 18 dissected by Alghamdi
retinacular band. et al. (2018), there was a supernumerary muscle situated lat-
eral to the right extensor digitorum brevis. This muscle arose
Prevalence from the cuboid and sent a tendon to the dorsum of digit five.
Wood (1868) found that the extensor digitorum brevis ten- Furthermore, extensor hallucis brevis fused proximally with
don for digit two was doubled in 6 out of 40 males (15%) extensor digitorum brevis on both sides of the body.
and 3 out of 30 females (10%). A slip to the first interos-
seous muscle was found in 3 out of 40 males (7.5%) and 1 Prevalence
out of 30 females (3.3%). Knott (1883b) found an extensor N/A
digitorum brevis tendon going to digit five in 2 out of 40
cases (5%). Clinical Implications
Out of 291 cases, Chaney et al. (1996) found that both Hypertrophy of the extensor digitorum brevis bundles may
the extensor hallucis brevis tendon to digit one and the simulate ganglia (Montgomery and Miller 1988).
extensor digitorum tendons to digits two through four were
present in 186 cases (63.9%). Tendons to the second, third,
and fourth digits were present without the extensor hallu- Extensor hallucis brevis (Figure 5.13)
cis brevis tendon in 50 cases (17.2%). In 11 subjects (3.8%),
Synonyms
there were only two tendons present to various digits. There
were more than four tendons present in 44 cases (15.1%). N/A
In a subsample of 284 limbs, Chaney et al. (1996) found an
extensor digitorum brevis tendon to digit five in 51 limbs Typical Presentation
(18%). In a sample of 11 feet from 8 cadavers, Kura et al. Description
(1997) found accessory muscles medial to extensor digito- Extensor hallucis brevis is regarded as a distinct medial slip
rum brevis going to digit two in 4 feet (36.4%). of extensor digitorum brevis (Standring 2016). This muscle
Sirasanagandla et al. (2013b) studied extensor digito- thus originates from the superolateral surface of the distal
rum brevis in 44 limbs. In 36 limbs (81.8%) both the exten- calcaneus, from the interosseous talocalcaneal ligament,
sor hallucis brevis tendon to digit one and the extensor and from the deep aspect of the base of the inferior extensor
retinaculum (Standring 2016). Its tendon inserts onto the extensor digitorum brevis (extensor hallucis brevis). Out of
dorsal surface of the base of the proximal hallucal phalanx 291 cases, Chaney et al. (1996) found that both the extensor
(Standring 2016). hallucis brevis tendon to digit one and the extensor digito-
rum tendons to digits two through four were present in 186
Innervation cases (63.9%). Tendons to the second, third, and fourth dig-
Extensor hallucis brevis is innervated by the deep fbular its were present without the extensor hallucis brevis tendon
nerve (Standring 2016). in 50 cases (17.2%). In 11 subjects (3.8%), there were only
two tendons present to various digits. There were more than
Comparative Anatomy four tendons present in 44 cases (15.1%).
Extensor hallucis brevis has a similar typical presentation Al-Saggaf (2003) studied the insertions of extensor hal-
in the apes, originating from the calcaneus and/or exten- lucis longus in 60 cadaveric lower limbs. In three cases (5%),
sor digitorum brevis and inserting onto the base of the an accessory tendon of extensor hallucis longus inserted
proximal hallucal phalanx (Champneys 1872; Hepburn with extensor hallucis brevis onto the base of the proximal
1892; Beddard 1893; Sonntag 1924; Boyer 1935; Raven hallucal phalanx. In two cases (3.3%), the accessory ten-
1950; Miller 1952; Lewis 1966; Gibbs 1999; Vereecke don joined the tendon of extensor hallucis brevis, forming
et al. 2005; Diogo et al. 2010, 2012, 2013a,b, 2017). In a common tendon that inserted onto the base of the proxi-
orangutans, there may be an origin from the navicular and mal hallucal phalanx. Sirasanagandla et al. (2013b) studied
the muscle may be fused with the extensor hallucis lon- extensor digitorum brevis and extensor hallucis brevis in 44
gus tendon (Boyer 1935; Gibbs 1999; Diogo et al. 2013a). limbs. In 36 limbs (81.8%) both the extensor hallucis brevis
An accessory slip attaching to digit two has been found in tendon to digit one and the extensor digitorum tendons to
gorillas (Raven 1950). digits two through four were present. In one limb (2.3%),
there was an accessory bundle present that originated from
Variations extensor hallucis brevis and the extensor digitorum brevis
Description belly going to digit two that merged with the frst dorsal
Extensor hallucis brevis may be absent (Macalister 1875; interosseous muscle.
Knott 1883b; Bergman et al. 1988; Chaney et al. 1996). It
may receive a slip from or be joined with extensor hallu- Anomalies
cis longus (Macalister 1875; Knott 1883b; Hallisy 1930; Description
Bergman et al. 1988; Al-Saggaf 2003; Hill and Gerges In a male fetus with triploidy studied by Moen et al. (1984),
2008). Accessory bundles may be present (Bergman et al. extensor hallucis brevis was absent on the right side. On
1988; Sirasanagandla et al. 2013b; Won and Oh 2018). Won the left foot of a male neonate with Meckel syndrome,
and Oh (2018) report a case in which the extensor digito- Pettersen (1984) found that the extensor hallucis brevis
rum muscle belly to digit three was doubled and associated tendon was rudimentary. In the female fetus with trisomy
with an accessory muscle in the right foot of a cadaver. The 18 dissected by Alghamdi et al. (2018), extensor hallucis
accessory muscle was covered by extensor hallucis brevis, brevis fused proximally with extensor digitorum brevis on
originated from the calcaneus, and inserted onto the lateral both sides of the body.
third of the transverse retinacular band.
Prevalence
Prevalence N/A
Knott (1883b) found that the extensor hallucis brevis ten-
don was absent in 1 out of 40 cases (2.5%). Hallisy (1930) Clinical Implications
found that in 3 out of 290 feet (1%), the tendon of exten- Hypertrophy of extensor hallucis brevis may cause anterior
sor hallucis longus sent a slip to the most medial tendon of tarsal tunnel syndrome (Tennant et al. 2014).
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Index
Note: Italic page numbers refer to fgures. coccygeofemoralis 262–263
coccygeus 225–226
abductor accessorius digiti minimi 325 compressor venae dorsalis 231
abductor digiti minimi constrictor pharyngis inferior 61, 65
of the foot 322, 324–325 constrictor pharyngis medius 61, 64–65
of the hand 196, 204–205 constrictor pharyngis superior 61–62
abductor hallucis 321–323 contrahentes digitorum manus 192–193
abductor ossis metatarsi digiti quinti 322, 325–326 contrahentes pedis 330, 333
abductor pollicis brevis 196, 203–204 coracobrachialis 150, 151–153
abductor pollicis longus 175, 191–192 coracobrachialis longus 150, 153–154
accessory radiocarpus 164, 167 coracobrachialis profundus 150, 153
acromioclavicularis 144 coraco–cervicalis 76
adductor brevis 281, 283–284 costoclavicularis 143
adductor hallucis 330, 331–333 costocoracoideus 141–142
adductor longus 281, 282–283 costodeltoideus 145
adductor magnus 281, 284–285 corrugator supercilii 22, 23–24
adductor minimus see adductor magnus cremaster 217, 219–220
adductor pollicis 196–197 cricoarytenoideus lateralis 69–70, 71
adductor pollicis accessorius see interossei palmaris cricoarytenoideus posterior 68–69
amygdaloglossal muscle 58 cricocorniculate muscle 69
anconeus 173, 176 cricoepiglotticus 69–70
anomalous nasi 28 cricohyoid 67
anterior fbulocalcaneus 306, 314 cricomembranosus 69
anterior scalene see scalenus anterior cricothyrohyoid 70
anterior tibiotalus see tibioastragalus anticus cricothyroideus (cricothyroid) 65–67
articularis cubiti see subanconeus cricotrachealis 67
articularis genus 269, 275–276
aryepiglotticus 71 deep transverse perinei see transversus perinei profundus
arytenoideus obliquus 68, 70–71 deltoideus (deltoid) 143, 144–145
arytenoideus transversus 68, 70 depressor anguli oris 22, 31–32
arythyrocricoid 70–71 depressor labii inferioris 22, 32
atlantobasilaris internus 100 depressor septi nasi 22, 33–34
atlantomastoideus 248, 249 depressor supercilii 23
auricularis anterior 46–47 diaphragm 232–234
auricularis posterior 46, 47–48 digastricus anterior 73, 82–85
auricularis superior 46, 47 digastricus posterior 73, 85–87
auriculoglossus 52–55 dilator tubae 55–56
axillary arch see pectorodorsalis Dollo’s law 3–4
azygos–pharyngeus 62 dorsal interossei see interossei dorsalis
dorsoepitrochlearis 133–134, 159
basiodeltoideus 144–145
basioglossus 52 epitrochleoanconeus 158–159
biceps brachii 154, 155–157 Evo–Devo–Path (evolutionary developmental pathology) 2–5
biceps femoris 286, 288–290 extensor carpi radialis accessorius 173, 179–180
biventer cervicis see semispinalis extensor carpi radialis brevis 173, 178–179
brachialis 154–155 extensor carpi radialis intermedius 177–179
brachiofascialis 156 extensor carpi radialis longus 173, 176–178
brachioradialis 173–174 extensor carpi radialis tertius 177
buccinatorius (buccinator) 22, 30 extensor carpi ulnaris 173, 180
bulbospongiosus 228, 232 extensor digiti minimi 181, 183–184
extensor digiti minimi accessorius 184
capsularis sub–brachialis 155 extensor digiti minimi et quarti 184
cephalo–pharyngeus 59–60, 63 extensor digiti quarti 185
ceratoarytenoid 69 extensor digiti secundus 308
ceratocricoid 67–68 extensor digitorum 181–183
ceratoglossus 51–52 extensor digitorum brevis 337, 338–339
cervico–auriculo–occipitalis 21 extensor digitorum brevis manus 181, 187–188
cervico–costo–humeralis 76 extensor digitorum longus 306, 310–311
chondrocoracoideus see costocoracoideus extensor hallucis brevis 337, 339–340
chondroepitrochlearis 131, 137–138 extensor hallucis capsularis 308–310
chondrofascialis 137–138 extensor hallucis longus 306, 308–310
chondroglossus 51–52 extensor indicis 175, 181, 185–186
cleidofascialis 76 extensor indicis brevis see extensor digitorum brevis manus
cleidohyoideus (cleidohyoid) 73, 74–75 extensor indicis et medii communis 186–187
cleido–occipitalis cervicalis 46, 107–108 extensor medii digiti 181, 186–187
375
376 Index
extensor ossis primi metatarsi 308–309 inferior rectus see rectus inferior
extensor pollicis brevis 175, 189–190 inferior thyroid muscle 72
extensor pollicis–et–indicis 175, 185–186 infraclavicularis 142–143
extensor pollicis longus 175, 181, 188–189 infraglenoid muscle 149
extensor pollicis tertius 189 infraspinatohumeralis 145
extensor primi internodii hallucis 308–310 infraspinatus 145–146
external anal sphincter see sphincter ani externus infraspinatus minor 146
external intercostals 235–236 innermost intercostals 235, 237
external oblique 213–214 interarytenoideus see arytenoideus obliquus and arytenoideus
transversus
fbularis accessorius see fbularis quartus interclavicularis 138
fbularis brevis 315, 317–318 intercostales externi see external intercostals
fbularis digiti minimi 315, 320–321 intercostales interni see internal intercostals
fbularis longus 314–317 intercostales intimi see innermost intercostals
fbularis quartus 315, 318–320 interfoveolar muscle 214
fbularis tertius 306, 311–314 internal cricoid muscle 66
fbulocalcaneus externum see fbularis quartus internal intercostals 235, 236
fbulocalcaneus internus 298, 302–303 internal oblique 213, 215–216, 217
fbulocalcaneus medialis see fbulocalcaneus internus interossei dorsalis
fexor carpi radialis 168, 170–171 of the hand 193, 199–200
fexor carpi radialis brevis 168, 171 of the foot 336–338
fexor carpi ulnaris 168, 169–170 interossei palmaris 193, 197–199
fexor carpi ulnaris accessorius 170 interossei plantares 326, 335–336
fexor carpi ulnaris brevis 166 interspinales 242–244
fexor digiti minimi brevis intertransversarii 243, 244
of the hand 196, 201–202 intrinsic muscles of the tongue 49–50
of the foot 330, 334–335 ischiocavernosus 228, 231–232
fexor digitorum accessorius longus 298, 300–302 ischiococcygeus see coccygeus
fexor digitorum brevis 322, 323–324 ischiofemoralis 262
fexor digitorum longus 298–300 ischiopubicus 231
fexor digitorum profundus 160, 162–163
fexor digitorum superfcialis 163–165 lack of homeostasis model 2
fexor hallucis brevis 330–331 lateral cricoarytenoid see cricoarytenoideus lateralis
fexor hallucis longus 298, 303–304 lateral pterygoid 39, 41–43
fexor indicis profundus 162–163 lateral rectus see rectus lateralis
fexor pollicis brevis 196, 200–201 latissimus dorsi 131, 149–151, 159
fexor pollicis longus 159–161 levator anguli oris 22, 28–29
frontalis see occipitofrontalis levator ani 226, 227–228
levator claviculae 128–130
Gantzer’s muscle 160, 161–162 levator epiglottidis 51, 53
gastrocnemius 291–293 levatores costarum breves and longi 239, 241–242
gemellus inferior 276, 279–280 levator glandulae thyroideae 73, 78–79
gemellus superior 276, 278–279 levator labii superioris 22, 27
genioglossus 50–51 levator labii superioris alaeque nasi 22, 27–28
geniohyoideus (geniohyoid) 73, 81–82 levator palpebrae superioris 88–90
geniopharyngeus 51, 62 levator scapulae 122, 125–126
glenobrachialis 148 levator tendinis musculi latissimus dorsi 150, 151
gluteoperinealis 232 levator veli palatini 55, 57
gluteus maximus 261–263 logic of monsters model 1–6
gluteus medius 261, 263–264 longissimus 250, 251–252
gluteus minimus 264–265 longus capitis 100–101
gluteus quartus see scansorius longus colli 99–100, 101
gracilis 281–282 lumbricales (lumbricals)
gracillimus 89 of the foot 326, 328–330
of the hand 194–196
hepatodiaphragmaticus 234
hyoglossus 50, 51–52 malaris 22, 24
mandibulo–auricularis 54
iliacus 220, 221–222 mandibuloglossus see myloglossus
iliacus minimus 220, 221–222 masseter 38–40
iliacus minor 220, 222–223 medial pterygoid 39, 43–44
iliocapsularis see iliacus minor medial rectus see rectus medialis
iliococcygeus see levator ani mentalis 22, 35–36
iliocostalis 250, 252 mentohyoid 82–85
incisivus labii inferioris 22, 35 middle pharyngeal constrictor see constrictor pharyngis medius
incisivus labii superioris 22, 34–35 middle scalene see scalenus medius
incisurae mediae obliquus 66–67, 77 multifdus 243, 245–246
inferior oblique see obliquus oculi inferior musculus uvulae 55, 57–58
inferior pharyngeal constrictor see constrictor pharyngis inferior myloglossus 50, 53–54
Index 377
mylohyoideus (mylohyoid) 73, 79–81 pronator quadratus 164, 165–166

pronator teres 168, 171–172
nasalis 22, 33 psoas accessorius 225
psoas major 220, 223–224
oblique arycorniculate muscle 71 psoas minor 220, 224–225
oblique arytenoid see arytenoideus obliquus psoas quartus 220, 223–224
obliquus accessorius inferior 94 psoas tertius 223
obliquus capitis inferior 103, 105–106 pterygofascialis 45
obliquus capitis superior 103, 105 pterygoideus lateralis see lateral pterygoid
obliquus externus abdominis see external oblique pterygoideus medialis see medial pterygoid
obliquus externus abdominis profundus 213, 214–215 pterygoideus proprius 39, 45
obliquus internus abdominis see internal oblique pterygopharyngeus 61–62
obliquus oculi inferior 89, 94–95 pterygospinous 39, 44–45
obliquus oculi superior 89, 93–94 pterygotympanicus see pterygospinous
obturator externus 276, 277–278 puboanalis 227
obturator internus 226, 276–277 pubococcygeus see levator ani
occipitalis see occipitofrontalis puboperitonealis 216
occipitalis minor see platysma cervicale puborectalis see levator ani
occipitalis proprius 21 pyramidalis 217, 218–219
occipital platysma see platysma cervicale
occipitofrontalis 21–22, 46 quadratus femoris 276, 280–281
occipitoscapularis see rhomboideus occipitalis quadratus lumborum 220–221
omoclavicularis 76 quadratus plantae 326–328
omohyoideus (omohyoid) 73, 75–77
opponens digiti minimi radiocarpus 164, 166–167
of the foot 330, 335 radiocubitocarpien 166
of the hand 196, 202–203 rectus abdominis 216–218
opponens hallucis 330, 333–334 rectus abdominis lateralis 217, 218
opponens pollicis 196, 201 rectus capitis anterior 101
orbicularis oculi 22–23 rectus capitis lateralis 101–102
orbicularis oris 22, 30–31 rectus capitis lateralis longus 102
orbitozygomaticus 22, 24–25 rectus capitis minimus 101
rectus capitis posterior major 102–104
palatoglossus 55, 58–59 rectus capitis posterior minor 103, 104–105
palatopharyngeus 55, 59–60 rectus femoris 269, 271–272
palmar interossei see interossei palmaris rectus inferior 89, 91–92
palmaris brevis 193–194, 196 rectus lateralis 89, 92–93
palmaris longus 167–169 rectus medialis 89, 92
palmaris profundus 167, 169 rectus superior 89, 90–91
panniculus carnosus 134 retractor bulbi 91, 93
Passavant’s muscle 59–60, 62 retractor glandulae lacrimalis 89
pectineus 281, 285 rhomboatloideus 247–248
pectoralis abdominis see pectoralis quartus rhomboideus major (rhomboid major) 122–123
pectoralis intermedius 127–128 rhomboideus minimus 123–125
pectoralis major 131, 135–136 rhomboideus minor 122, 123–124
pectoralis minimus 135, 140–141 rhomboideus occipitalis 122, 124–125
pectoralis minor 135, 136–137 rhomboideus tertius 125
pectoralis tertius see pectoralis quartus risorius 22, 29–30
pectoralis quartus 135, 139 roos bands/muscles 143–144
pectorodorsalis 131, 140 rotatores longus and brevis 243, 245
peroneocalcaneocubcoideus see fbularis quartus
peroneocalcaneus see fbularis quartus sacrococcygeus anterior 226–227
peroneocuboideus see fbularis quartus sacrococcygeus posterior 243, 244
peroneoperoneolongus see fbularis quartus salpingopharyngeus 60–61
peroneotibialis 291, 295 salpingostaphylinus 57
peroneus quartus see fbularis quartus saphenous muscle 214, 271
peroneus digiti quinti see fbularis digiti minimi sartorius 269, 270–271
petropharyngeus 61, 63–64 scalenus anterior 95–96, 98
piriformis 226, 261, 265–267 scalenus medius 96–97, 98
pisiannularis 204 scalenus minimus 98–99
pisimetacarpeus 204 scalenus posterior 97–98
pisiuncinatus 204 scansorius 261, 267–268
plantar interossei see interossei plantares scapuloclavicularis 127, 129
plantaris 291, 295–297 scapulohumeralis digastricus singularis 146
platysma cervicale 37–38, 46 semimembranosus 286, 287–288
platysma myoides (platysma) 22, 36–37 semispinalis 246–247, 248
popliteus 291, 297–298 semitendinosus 285–287
posterior cricoarytenoid see cricoarytenoideus posterior serratus anterior 121–122
procerus 22, 32–33 serratus posterior inferior 239, 240–241
378 Index
serratus posterior superior 239–240 tensor fasciae latae 269–270

soleus 291, 293–295 tensor fasciae suralis 286, 290–291
soleus accessorius 294 tensor semivaginae articulationis humeroscapularis 141, 142
sphenotemporalis 39, 44 tensor trochleae 89, 90
sphincter ani externus 228–229 tensor tympani 48
sphincter colli profundus 36 tensor veli palatini 55–57
sphincter urethrae 228, 230–231 tenuissimus 261, 268–269
splenius capitis 247, 248 teres major 145, 147–148
splenius cervicis 247, 248 teres minor 145, 147
spinalis 249–251 thyreo–corniculatus 72
stapedius 48–49 thyroarytenoideus (thyroarytenoid) 71–72
sternalis 131–133 thyroepiglotticus 72
sternochondroscapularis 140–141 thyrohyoideus (thyrohyoid) 73, 78
sternoclavicularis 135, 138–139 thyroideus transversus anomalus 66–67
sternocleidomastoideus (sternocleidomastoid) 46, 106–107, 129 thyrotrachealis 66, 67
sternohyoideus (sternohyoid) 73–74 tibialis anterior 305–308
sternohyoideus azygos 74 tibialis posterior 298, 304–305
sternoscapularis 129, 130 tibialis secundus 305
sternothyroideus (sternothyroid) 73, 77–78 tibioastragalus anticus 307–308
straight arycorniculate muscle 70 tibiocalcaneus internus 291, 294
styloauricularis 50, 54–55 tibiofascialis anticus (tibiofascialis anterior) 307
stylochondrohyoideus 87 tibiotarsalis 294, 305, 325
stylopharyngeus 61, 62–63 tongue see intrinsic muscles of the tongue
styloglossus 50, 52–53 transpleuralis subclavius 233
stylohyoideus (stylohyoid) 73, 87–88 transversalis cervicis medius 95
stylohyoideus profundus 87 transversalis cervicis posticus minor 251
subanconeus 157 transverse arytenoid see arytenoideus transversus
subclavius 126–127 transversus abdominis 213, 216
subclavius posticus 127, 128 transversus glabellae 23
subcostales (subcostal muscles) 235, 237–238 transversus menti 31–32
subscapularis 145, 148–149 transversus nuchae see platysma cervicale
subscapularis minor 145, 149 transversus orbitis 89
subscapularis–teres–latissimus 148–149 transversus perinei profundus 228, 229–230
subscapulo–capsularis 148 transversus perinei superfcialis 228, 230
superfcial transverse perinei see transversus perinei superfcialis transversus thoracis 233, 234–235
superior cricoarytenoid 69 trapezius 46, 108–109, 129
superior oblique see obliquus oculi superior triceps brachii 145, 157–158
superior pharyngeal constrictor see constrictor pharyngis superior triticeoglossus 52
superior rectus see rectus superior
superior thyroarytenoideus (superior thyroarytenoid) 71, 72–73 ulnaris digiti quinti 173, 180–181
supinator 174–176 ulnocarpus brevis see fexor carpi ulnaris brevis
supinator radii longus accessorius 174 uvula see musculus uvulae
supraclavicularis proprius 129, 130–131
supracostalis anterior 233, 238 vastus intermedius 269, 274–275
supracostalis posterior 238–239 vastus lateralis 269, 273–274
supraspinatus 145, 146–147 vastus medialis 269, 272–273
syndesmopharyngeus 65 ventricularis 72
vocalis 71, 72
temporalis 39, 40–41
temporalis minor 40 zygobuccinator muscle 30
temporobuccinator band 30, 40 zygomatico–mandibularis 39
temporoparietalis 21 zygomatico–orbicularis see orbitozygomaticus
tensor capsuli tibiotarsalis anterior see tibialis secundus zygomaticus major 22, 26–27
tensor fascia colli see supraclavicularis proprius zygomaticus minor 22, 25–26

Handbook of Muscle Variations and Anomalies in Humans - A - Eve K. Boyle, Vondel S. E. Mahon, Rui Diogo - 2022 - CRC Press - 9780367538620 - Anna's Archive

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Handbook of Muscle Variations and

and by CRC Press

CRC Press is an imprint of Taylor & Francis Group, LLC

ISBN: 978-0-367-53862-0 (hbk)

Chapter 1 Introduction ...................................................................................................................................................... 1

Chapter 2 Head and Neck Muscles ...................................................................................................................................9

Chapter 3 Upper Limb Muscles .................................................................................................................................... 111

Chapter 4 Trunk Muscles..............................................................................................................................................207

Chapter 5 Lower Limb Muscles.................................................................................................................................... 253

References ......................................................................................................................................................................... 341

Typical Presentation ................................................................................................................................................... 35

Typical Presentation ...................................................................................................................................................44

Geniohyoideus (Geniohyoid) .......................................................................................................................................... 81

Rectus medialis (Medial rectus)......................................................................................................................................92

Posterolateral Neck Muscles..............................................................................................................................................106

FACIAL MUSCLES occipitalis is typically differentiated into a main portion

Typical Presentation Prevalence

Synonyms OrbitOzygOMaticus (figure 2.1)

sheet that extends between orbicularis oculi and orbicu- Prevalence

Clinical Implications Anomalies

Innervation Orbicularis oris (Figure 2.1)

Comparative Anatomy Synonyms

Comparative Anatomy Comparative Anatomy

Innervation Clinical Implications

Typical Presentation Prevalence

Typical Presentation Description

The upper fbers of mentalis interlaced with the inferior Variations

superfcial layer and three deeper layers of the masseter Variations

EXTRINSIC MUSCLES OF THE EAR Variations

stapeDius (nOt illustrateD)

Comparative Anatomy Clinical Implications

FIGURE 2.4 Muscles of the tongue in lateral view.

(enlargement), bifd tongue (anterior separation of the tongue Prevalence

Variations and the hyoglossus muscles. Furthermore, the right genio-

Anomalies MUSCLES OF THE SOFT PALATE

FIGURE 2.5 Muscles of the soft palate in posterior view.

Variations Musculus uvulae (figure 2.5)

Innervation Clinical Implications

FIGURE 2.6 Pharyngeal muscles in posterior view.

Innervation side and inserted deep into hyoglossus (Pettersen 1979). In an

Comparative Anatomy Description

Variations LARYNGEAL MUSCLES

Description cricOthyrOiDeus (cricOthyrOiD) (figure 2.7)

FIGURE 2.7 Anterior laryngeal muscles in anterior view.

Anomalies ceratOcricOiD (figure 2.8)

Synonyms Typical Presentation

FIGURE 2.8 Posterior laryngeal muscles in posterior view.

laryngeal nerve (Turner 1860; Hetherington 1934; Sharp Anomalies

Comparative Anatomy In a fetus with craniorachischisis, the posterior cricoary-

Variations Typical Presentation

Typical Presentation connect with the thyroarytenoid or the lateral cricoary-

FIGURE 2.9 Laryngeal muscles in superior view.

Synonyms (the ventricular muscle), a muscle situated in the vestibular

See also: Thyroarytenoideus

Innervation INFRAHYOID MUSCLES

FIGURE 2.10 Suprahyoid and infrahyoid muscles in anterior view.

Comparative Anatomy males and present in 29 out of 230 sides (12.61%) in

Synonyms OMOhyOiDeus (OMOhyOiD) (figure 2.10)

thyrOhyOiDeus (thyrOhyOiD) (figure 2.10) Anomalies