2018 A Concise Dictionary of Paleontology

Robert L.
Carlton
A Concise
Dictionary of
Paleontology
Technical Editor:
Brittney A. Oleniacz
A Concise Dictionary of Paleontology
Robert L. Carlton
A Concise Dictionary
of Paleontology
Technical Editor: Brittney A. Oleniacz

Robert L. Carlton
Salem, Oregon, USA
ISBN 978-3-319-73054-7 ISBN 978-3-319-73055-4 (eBook)

https://doi.org/10.1007/978-3-319-73055-4
Library of Congress Control Number: 2017963120
© Springer International Publishing AG 2018

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For Patricia, Morgan, Melissa, Brendan,
and Cayden
Acknowledgments
This book owes its existence chiefly to my patient and forbearing Patricia. But I
must also express my gratitude to my excellent editor, Brittney Oleniacz, and to the
many other paleontologists whose advice and encouragement helped the project
along. I am especially grateful to Paul Selden, whose detailed advice greatly
improved an earlier draft. Thanks also to Frankie Jackson, David Varrichio, Ben
Creisler, William Schopf, Kathleen Ritterbush, Tony Ekdale, Frank DeCourten, Phil
Debarros, Tom Deméré, Ellen Thomas, Jonathan Payne, Demir Altiner, and Walter
Obermiller. Special thanks to the librarians who helped me rifle the collections at
the Berkeley, San Diego, and Merced campuses of the University of California; the
Salem Public Library; Chemeketa College Library; Willamette University Library;
and the Valley Library at Oregon State University. And finally, for actually bringing
this book to life, thanks to Sherestha Saini and the very capable Springer staff.
vii
Contents
A�� 1
B �� 31
C�� 43
D�� 71
E �� 89
F �� 103
G�� 109
H�� 121
I�� 137
J�� 143
K�� 147
L �� 153
M�� 165
N�� 185
O�� 193
P �� 205
Q�� 245
R�� 247
S �� 257
T �� 279
U�� 299
ix
x Contents
V�� 301
W�� 305
X�� 309
Y�� 311
Z �� 313
Further Reading�� 315

Introduction
This little book is a moderately comprehensive reference handbook covering the

most important paleontological taxa, terms, concepts, and localities. It is written
primarily for beginning students in the field and general readers. My professional
experience lies in technical writing and in teaching literature and history, with a
lifelong interest in paleontology inspired by Professor Charles Higgins at U.C. Davis
and by studying the geology collection in U.C. Berkeley’s beautiful Bacon Hall
before it was demolished in 1961. Using the essays of Loren Eiseley, Stephen
Gould, and E.O. Wilson in my composition classes in the 1980s, I made critical
thinking and the evolution of life on earth the center of my teaching. An interesting
technical writing project in the early 1990s showed me the lack of a dictionary like
this, and I began the groundwork for a good reference work as I directed student
research in the history of life in my writing courses. Retiring from teaching in 2014,
I have widened my understanding of paleontology with intensive study and have
worked systematically to complete the dictionary in the last three years.
There are many science dictionaries and several purporting to cover natural sci-
ence or earth science, but I have found them all very inadequate for paleontology.
Even the larger works like the McGraw-Hill Dictionary of Scientific and Technical
Terms, now in its seventh edition, have minimal and often outdated coverage of the
field, with very little detail on relationships and significance. So I believe there is a
need for a more comprehensive dictionary in print.
Online reference sites like the Paleobiology Database, the World Register of
Marine Species, and Palaeos are excellent for some basic information needed by
specialists, but they are of little use to the general reader. Wikipedia is good for its
detailed articles and up-to-date coverage, and I found some information in it for
about three-fourths of my entries. But it has surprising gaps and mistakes, and it
often reflects individual opinions that are not completely trustworthy.
The lexicon of paleontology is of course too large to allow really comprehensive
coverage of taxa, concepts, or terminology in a small dictionary. Here I try to include
the most significant taxonomic groups down to the family level, as well as the most
commonly mentioned genera in the literature. But in addition to covering these
traditionally important taxa, I have included up-to-date information on several other
xi
xii Introduction
categories. First, I focus on taxa associated with historical turning points such as
mass extinctions and first appearances of derived characters. Second, in addition to
these transitional groups, I have included many entries that present phylogenetic
problems because they do not fit neatly into established categories, thus inviting
further research. Third, I pay detailed attention to Konservat-Lagerstätten that pro-
vide comprehensive information on the fossil communities and ecosystems of par-
ticular time frames, bringing together information from many sources on the most
significant Lagerstätten and other collecting localities. Fourth, I have also tried to
present the most current ideas about early hominin evolution.
With the aid of my excellent editor, Brittney Oleniacz, I have tried to maintain a
mid-level tone, keeping the language accessible but including enough technical
specificity to be useful for the serious student. To narrow the scope, I have kept the
references to theoretical concepts to a minimum. Now let me explain my
procedures.
In the entries for taxa, I state the classification and other relationships first, then
a physical description of notable traits, and sometimes an indication of significance.
I supply translations of names only when the etymology aids in identifying the
taxon. Length measurements are nose-to-tail unless otherwise specified. The entries
end with information on collecting localities and chronostratigraphic position. The
references to geographic distribution indicate only the collecting localities of speci-
mens, not necessarily the actual distribution unless so noted. The term “global dis-
tribution” excludes Antarctica, and I use the term “cosmopolitan” for taxa with
widely scattered and relatively sparse occurrences in appropriate environments.
All taxa listed as entries are extinct unless noted as extant. Most dates assigned
to organisms and geological periods are approximate, but I indicate greater precise-
ness and certainty when possible. For dates and capitalization of the divisions of the
time scale, I follow the Geologic Time Scale and the International Chronostratigraphic
Chart (2017), as published by the International Commission on Stratigraphy. The
latter provides the current names and dates of the most broadly accepted subdivi-
sions of geological periods and epochs.
The entries for descriptive terms generally include only terms used in this dic-
tionary, and they refer only to the paleontological application of the term. Geological
and biological references are made only where necessary for their paleontological
significance.
I have attempted to provide the most generally accepted taxonomic relationships,
generally following the classifications given by the Paleobiology Database, and also
its distribution information where it is given and not contradicted by other sources.
But since paleontological description and taxonomy is by its nature a work in
progress, with frequent changes in classifications as a result of ongoing work by
specialists, there is no question of full coverage or even of up-to-the-minute accu-
racy. Newly described taxa increase every day both the range of first and last appear-
ances and the range of geographic occurrences. So the names, classifications, and
dates given here are not to be taken as set-in-stone determinations but rather as
handles to provide access to further information about the taxa being discussed.
With this in mind, I have tried to give at least two references to related taxa for
Introduction xiii
each taxon entry. Of course, mistakes and omissions are inevitable in such an
ambitious undertaking as this dictionary, and I will welcome all suggestions for
improvement.
I have tried to steer a middle course between strict neo-Linnaean taxonomy and
the extremes of modern cladistic systematics. As Colin Tudge so eloquently demon-
strates in The Variety of Life, cladistic analysis is crucial in improving the accuracy
of our picture of the tree of life. However, systematic taxonomy may blur the impor-
tant distinctions between large groups such as dinosaurs and birds, or eurypterids
and arachnids. At least for my purposes, when referring to reptiles in general, the
paraphyletic classification Reptilia is far preferable to “non-avian, non-mammalian
amniotes.” Also, the idea that each node in the branching of the tree deserves a new
rank is a leap too far, even for most professionals, and I have chosen to ignore sub-
groups like microclass, parvorder, and sub-tribe.
The final section of the book, “Further Reading,” includes all of the important
monographs that I have consulted, as well as many of the more important journal
articles. But because writing a dictionary requires checking many print and online
sources for each entry, each entry contains information from several sources, and I
have not attempted to include references to all that I have consulted.
A
Aachenosaurus: name given to a piece of petrified wood found in the Aachenian

deposits near Moresnet, Belgium, and mis-identified in 1888 as part of a duckbill
dinosaur’s jaw.
Aaptoryctes: “unapproachable digger,” a monospecific genus of insectivorous sori-
comorph mammals in the family Palaeoryctidae; 4–5 inches long and known only
from three specimens found in Wyoming, the shrew-like A. ivyi had a stronger
crushing dentition than its cousin Palaeoryctes; United States, late Paleocene,
57–56 Ma.
Aardonyx celestae: a species of anchisaurian sauropod dinosaurs described in
2010. Known from disarticulated fragments of two individuals dated at 195 Ma, it
is estimated to have been about 40 feet long. Aardonyx seems to be intermediate
between the earlier prosauropods, which were generally smaller and often ran on
their hind legs, and the later, more ponderous sauropods. Spion Kop, South Africa;
Early Jurassic.
abathochroal: an unusual type of compound eye found only in the early trilobite
families of the suborder Eodiscina, comprised of a small number of lenses (about
70) with individual corneas.
Abavornis: a proposed genus of primitive birds of the Late Cretaceous, about
85 Ma; known only from a single bone in the shoulder process, it is tentatively clas-
sified as enantiornithine; it was found in Uzbekistan in 1998 and given the species
name A. bonaparti.
Abelisauridae: a long-lived family of ceratosaurian theropod dinosaurs with two
subfamilies, Majungasaurinae and Carnotaurinae; known almost exclusively from
Gondwana, they were perhaps the most successful large predators there in the
Cretaceous; they are characterized by strong hindlimbs, almost vestigial forelimbs,
and a variety of knobs and horns on a short skull; see also Arcovenator,
© Springer International Publishing AG 2018 1

R.L. Carlton, A Concise Dictionary of Paleontology,
https://doi.org/10.1007/978-3-319-73055-4_1
2 A
Eoabelisaurus, and Rugops; India, Africa, Madagascar, South America, and France;
Middle Jurassic to Late Cretaceous.
Abelisaurus: a genus of theropod dinosaurs discovered in Patagonia, the type spe-
cies of the family Abelisauridae; known from a 35-inch incomplete skull but prob-
ably bipedal, A. comahuensis is estimated at 25–30 feet long and 16 tons; Argentina,
Late Cretaceous, about 80 Ma.
Abrictosaurus: a genus of basal ornithopod dinosaurs in the family
Heterodontosauridae; the size of a large dog, about 5 feet long and nearly 100
pounds, it had a long tail and was bipedal and herbivorous; it had high-crowned
teeth but lacked the canine-like teeth on the lower jaw that were typical of heter-
odontosaurids; known from fragments of two individuals; Lesotho, South Africa,
Early Jurassic.
Abrocomidae: a still-extant family of caviomorph South American rodents in the
suborder Hystricomorpha; commonly known as rat chinchillas, the Abrocomidae
diverged from the ctenomyids in the late Miocene; two extant species can be found
in the central Andes.
Abydosaurus: a genus of herbivorous saurischian dinosaurs in the suborder
Sauropodomorpha, discovered in 2010 at Dinosaur National Monument; remark-
able because four skulls have been found, two of them entire, very unusual for the
relatively delicate head of a sauropod; related to Brachiosaurus, Abydosaurus lived
about 100 Ma; the size is unclear because all four specimens are from juveniles,
which were about 25 feet long when they died; Utah, Early Cretaceous.
Acanthochaetetes wellsi: see chaetetids.
Acanthodidae: one of the three largest families of acanthodian fish; characterized
by only one dorsal fin, they were generally edentulous, adapted to filter-feeding;
North America, Europe, and Africa; Devonian to Permian.
Acanthodii: a paraphyletic class of primitive bony fishes in the infraphylum
Gnathostomata. The first organisms to develop an efficient, moveable lower jaw, the
acanthodians possessed characteristics of both chondrichthyans and osteichthyans.
They are characterized by stout spines supporting numerous fins, usually two dorsal
fins and a strongly heterocercal tail fin. The Acanthodii are generally considered to
include the families Acanthodidae, Climatiidae, Diplacanthidae, Gyracanthidae,
and Ischnacanthidae. Early Silurian to late Permian.
Acanthostega: a genus of stem-tetrapods intermediate between the lobe-finned
fish of the Middle Devonian and the amphibians which began to colonize the land
later in the period. The salamander-shaped Acanthostega, dated at 365 Ma, was
6 feet long and famously had 8 digits on its forelimbs and 7 on its hindlimbs. See
Tetrapoda. Greenland, Late Devonian.
A 3
Acanthothiris: an obscure genus of articulate brachiopods in the still-extant order

Rhynchonellida and extinct family Acanthothirididae; characterized by numerous
hollow spines; Europe, Asia, and Africa; Early to Late Jurassic.
Acanthothoraci: “spine chest,” an order of primitive placoderms, closely related
to the rhenanids and also similar to the ptyctodonts except for having somewhat
heavier armor; characterized especially by a pair of large spines projecting from the
chest; North America, Europe, Asia, and Australia; Early to Middle Devonian.
Acari: a still-extant subclass of arthropods in the class Arachnida that includes the
mites (Actinotrichida and Anactinotrichida) and the ticks (Ixodida); by far the most
numerous group of arachnids, more than 30,000 species of mites are known, with a
far greater number still undescribed; global distribution, Early Devonian to Recent.
Acervularia: a genus of cerioid corals in the order Rugosa; characterized by dome-
shaped colonies with tightly packed polygonal corallites; Europe and North
America, middle Silurian to Middle Devonian.
Achaenodon: a genus of pig-like artiodactyl mammals in the family Dichobunidae;
formerly called Protelotherium; North America and Africa, Eocene.
Achelousaurus: a monospecific genus of medium-sized centrosaurine ceratopsian
dinosaurs, A. horneri is known from three skulls found in the Two Medicine
Formation of Montana and Alberta; about 20 feet long and weighing about 3 tons, it
is notable for a large skull and frill, with two small horns at the back of the frill;
North America, Late Cretaceous, 74 Ma.
Achillobator: a genus of large maniraptoran theropod dinosaurs in the family
Dromaeosauridae; similar to Deinonychus but at least a third larger — the type spe-
cies A. giganticus was very large for a dromaeosaur, up to 19 feet long; Mongolia,
Middle to Late Cretaceous.
Acipenseridae: the sturgeons, a family of “living fossils,” bony fish in the order
Actinopterygii. Their ancestry reaches back to the Triassic, with true sturgeons
appearing in the Late Cretaceous. They are notable for their size (the largest are
more than 20 feet long and weigh more than a ton) and individual longevity (a cen-
tury). Also notable for their longevity as a taxonomic family, being very little
changed since the Cretaceous. Some species, such as the pallid sturgeon,
Scaphirhynchus albus, are endangered because they are anadromous and often can-
not reach their spawning grounds. North America, Europe, and Asia; Triassic to
Recent.
acritarchs: “uncertain origin,” not a taxonomic group but a large and varied group
of non-carbonate and non-siliceous organisms whose remains cannot be easily clas-
sified. Nevertheless, they are so abundant and diverse through the Precambrian and
up to the Devonian that they are extremely important as biostratigraphic markers,
especially for times when other fossils are not abundant. At least some of the acri-
tarchs are thought to be the resting cysts of phytoplanktonic algae, and they may be
ancestral to the dinoflagellates. Dating back to 1400 Ma (even possibly more than
4 A
3000 Ma), the collection was grouped informally in form taxa in the 1990s on
morphological criteria and wall structure, but it awaits classification on a more natu-
ral basis. The acritarchs seem to have declined markedly at the time of the Cryogenian
Glaciation, about 650 Ma, and also during other periods of widespread glaciation.
The Doushantuo Formation in Southern China contains an important collection of
acritarchs from 590 to 565 Ma, during the Cryogenian Period.
Acrodelphidae: an extinct family of long-snouted cetaceans in the suborder
Odontoceti; North America and Europe, early Miocene to early Pliocene.
Acrosalenia: a genus of Mesozoic sea urchins in the still-extant family
Acrosaleniidae; characterized by keeled teeth and solid spines; Europe and Africa,
Early Jurassic to Early Cretaceous.
Acrotretoidea: a superfamily of inarticulate brachiopods in the class Lingulata
and order Acrotretida; characterized by conical pedicle valves; cosmopolitan, early
Cambrian to Devonian.
Actinistia: a subclass of sarcopterygian fish that includes the living coelacanths
and several extinct families of lobe-finned fish.
Actinocerida: a subclass of middle Paleozoic nautiloid cephalopods; generally
straight, most were about 3 feet long, with a few somewhat larger; found in shallow-
water strata, they probably alternated between swimming and resting on the bottom;
Early Ordovician to Devonian.
Actinopterygii: a large subclass of ray-finned fishes in the class Osteichthyes that
includes most of the living bony fishes; the earliest actinopterygians arose in the
Devonian.
Actinostromariidae: a family of reef-building Mesozoic demosponges; formerly
considered stromatoporoids; Europe, Asia, and Mexico; Late Jurassic to Late
Cretaceous.
Actinotrichida: see Acari.
Adapidae: a family of early primates in the suborder Strepsirrhini, probably
diverging from the Plesiadapiformes in the late Paleocene, about the same time as
the Omomyidae; the type genus is Adapis; they are generally considered to be
related to the North American notharctids, including Notharctus and Smilodectes,
and are possibly ancestral to the lemuroids of Madagascar; Europe, North America,
and Asia; late Paleocene to late Miocene.
adaptive radiation: a sudden increase in diversity within a group of organisms,
resulting from environmental changes that open new ecological niches in a geo-
graphically isolated area. The marsupials of Australia and the birds of the Galápagos
Islands are the most widely known examples, and one of the most scrupulously
described is E.O. Wilson’s ground-breaking study in the 1950s of ant speciation on
the island of New Caledonia.
A 5
advanced: of organisms: a state derived from a more primitive form, more modified
and specialized than primitive ancestral species.
Aechminidae: a Paleozoic family of ostracods in the suborder Beyrichicopina;
characterized by lobate shells with a single dorsal spine; Ordovician to early
Permian.
Aeduellidae: a family of chondrostean actinopterygian fish; North America and
Europe, Pennsylvanian to early Permian.
Aegyptocetus: a genus of semi-aquatic archaeocete whales in the family
Protocetidae; notable for cranial features that gave it the ability to hear underwater,
but also for spinal features similar to those of other terrestrial weight-bearing proto-
cetids such as Rodhocetus; Egypt, middle Eocene.
Aegyptopithecus: a genus of early anthropoid primates in the suborder Catarrhini,
closely related to the smaller Propliopithecus; 2 to 3 feet long, it was probably her-
bivorous and arboreal; Fayum, Egypt, late Eocene to early Oligocene.
Aegyptosaurus: a genus of titanosaurian sauropod dinosaurs, about 50 feet long
and weighing 12 tons; it had a long neck and tail, and had relatively long legs for a
sauropod; the only known specimens were discovered by Ernst Strömer but were
lost during World War II; North Africa, Cretaceous, 106–94 Ma.
Aelurognathus: “cat jaw,” a genus of small therapsids in the family Gorgonopsidae,
related to Rubidgea; about 2 feet long, it is characterized by very long canine teeth;
known from several specimens from the Karoo Basin; South Africa, late Permian.
Aepycamelus: a genus of giraffe-like artiodactyl mammals in the family Camelidae;
about 10 feet tall, with long legs and a long neck, they browsed in open grassland;
North America, early to late Miocene.
Aepyornis: a genus of giant ratite flightless birds in the still-extant infraclass
Palaeognathae and extinct order Aepyornithiformes; commonly called “elephant
bird,” Aepyornis was about 10 feet tall and weighed almost 900 pounds; it is known
only from Madagascar and became extinct after humans arrived on the island;
restricted to Pleistocene.
Aerosteon riocoloradensis: a species of carnivorous dinosaurs from 85 Ma, nota-
ble for breathing the same way as modern birds do, with a bellows-like pulmonary
system; Argentina, Late Cretaceous.
Aethophyllum: an unranked monospecific genus of herbaceous conifers; low-
growing and spreading, A. stipulare differs from most fossil conifers, which were
trees; Europe and China, Middle Triassic.
Aetiocetidae: a family of basal toothed mysticete whales, notable for representing
one of the transitional steps between toothed and baleen whales. From 10 to 26 feet
long, aetiocetes were toothed but also possessed nutrient foramina and therefore
probably some form of baleen, a synapomorphy of baleen whales. See also
6 A
Mammalodontidae. British Columbia, Washington, Oregon, Mexico, and Japan;

early to late Oligocene.
Aetodactylus halli: a species of ornithocheirid pterodactyloid pterosaur from the
Early Cretaceous; it had a wingspan of 9 feet and a long, narrow snout lined with
sharp, fish-catching teeth; see also Cimoliopterus; Texas, 97 Ma.
Aetosauria: “eagle lizard,” a monofamilial order of crocodilian archosaurs, con-
sisting of the family Stagonolepididae; generally heavily armored, they ranged from
3 to 10 feet long, and several genera bore backward-projecting spines to protect the
back; some are important Triassic index fossils; although known only from the Late
Triassic, they probably originated in the Middle or even Early Triassic; global dis-
tribution, Late Triassic.
Afrosoricida: a proposed order of mammals to include the tenrecs and golden
moles (Chrysochloridae); Africa and Madagascar, Oligocene to Recent.
Afrotarsius: a genus of haplorrhine primates in the family Tarsiidae; known from
several fragments found in North Africa and Burma; Africa and Asia, Eocene to
Oligocene.
Afrotheria: a large clade that includes the clade Tethytheria (proboscideans, sire-
nians, and desmostylians) as well as other genera that evolved in Africa, such as the
aardvarks, tenrecs and hyraxes; see also Eutheria; Paleocene to Recent.
Afrovenator: a monospecific genus of theropod dinosaurs in the family
Megalosauridae, known from a single, relatively complete skeleton; 27 feet long
and similar to Allosaurus, A. abakensis was smaller than Tyrannosaurus rex but an
efficient predator with strong forelimbs and sickle claws; Niger, Middle Jurassic,
193 Ma.
Agate Fossil Beds National Monument: a site of lower Miocene volcaniclastic
deposits in northwestern Nebraska, notable for specimens of several mammal spe-
cies that are slightly different from similar species of the Oligocene White River
Group fauna of the same area; one of the most striking is the helical burrow of
Palaeocastor, q.v., and others are Amphicyon, Miohippus, and the chalicothere
Moropus; United States, early Miocene.
Agathoxylon: a genus of large gymnosperm trees in the family Araucariaceae;
Europe, South and North America, Africa, and Antarctica; Middle Jurassic to late
Miocene.
Agilisaurus: a monospecific genus of ornithischian dinosaurs, probably related to
Heterodontosaurus; about 4 feet long and lightly built, A. louderbacki was option-
ally quadrupedal but agile and a fast runner bipedally; known from a single com-
plete skeleton dated between 168 and 161 Ma, it has the typical ornithischian
beak-like predentary structure; China, Middle Jurassic.
Agilodocodon: a monospecific genus of Jurassic therapsids in the suborder
Docodonta, found in the Tiaojishan Formation and dated at about 160 Ma. Described
A 7
in 2015 as only 5–6 inches long, the shrew-like A. scansorius is the earliest-known

tree-climbing mammaliaform animal. It had spade-shaped teeth, long claws, and
flexible elbows and ankles, and probably fed on sap by gnawing through tree bark.
China, Late Jurassic.
Aglaspidida: an order of arthropods, generally considered to be related to the che-
licerates but now placed in the class Artiopoda; they are superficially similar to a
trilobite with the elongate tail spine of a horseshoe crab; North America, Europe,
Asia, and Australia; early to late Cambrian.
Agmata: a proposed phylum; erected in 1977 to include the enigmatic fossils
Salterella and Volborthella of the early Cambrian; reaffirmed in 2003 but still
controversial.
Agnatha: a class of jawless fish, a still-extant paraphyletic superclass of verte-
brates; dating back as early as 530 Ma, the agnathans include the extinct ostraco-
derms and conodonts, as well as the modern lampreys and hagfish; see also
Gnathostomata; early Cambrian to Recent.
Agnosphitys: a monospecific genus of early dinosaurs, placed variously in the
families Guaibasauridae and Silesauridae; dated about 206 Ma, A. cromhallensis is
known only from partial remains but seems to be a stem-dinosaur; England, Late
Triassic.
Agnostida: a primitive order of small early trilobites; isopygous and generally
eyeless, they had only two or three thoracic segments; some genera are thought to
be planktic, but most are benthic like most other trilobites; global distribution, early
Cambrian to Late Ordovician.
Agnostus: a genus of small trilobites in the order Agnostida and family Agnostidae;
one of the earliest trilobites, especially prominent in Cambrian deposits; North
America, Europe, and Asia; early Cambrian to Early Ordovician.
Agoniatitida: an early order of primitive ammonoids, ancestral to all later ammo-
noids; also known as Anarcestida; global distribution, Early to Middle Devonian.
Agorophius: a genus of early odontocete whales in the family Agorophiidae; first
named by E.D. Cope in the 1870s, the family was long in question because the
holotype of the genus was lost, but a 2016 description of a skull of Agorophius pyg-
maeus substantiates its position as a basal odontocete; North America, known only
from the late Oligocene, 28–24 Ma.
Agriochoeridae: “wild pig,” a family of oreodont artiodactyl mammals in the sub-
order Tylopoda. Superficially pig-like or sheep-like and as large as a small horse,
they are now generally thought to be related to camels. Less advanced than their
cousins the Merycoidodontidae, they had a selenodont dentition and claws instead
of hooves, rare for artiodactyls. Widespread in North America but especially well-
known from specimens in the Turtle Cove fauna of the John Day Fossil Beds. North
America, late Eocene to early Miocene.
8 A
Agriotherium: “hunting beast,” a genus of early bears in the subfamily

Agriotheriinae; formerly called Hyaenarctos; mainly carnivorous and about 9 feet
long, it was among the larger bears of the Pliocene; North America, Europe, Asia,
and Africa; late Miocene to early Pleistocene.
Aigialosauridae: a family of semi-aquatic varanoid lizards in the order Squamata
and extinct superfamily Mosasauroidea. The type genus is Aigialosaurus (formerly
called Opetiosaurus). About 3 feet long, the aigialosaurs show some aquatic spe-
cializations — a slightly flattened tail and reduced limbs — but were not structur-
ally different from terrestrial ancestors. Considered basal mosasaurs, they are
important as the earliest-known ancestors of larger and more specialized mosasaurs,
which appeared about 92 Ma. Known from a small number of fragments from
Europe, Late Jurassic to Late Cretaceous (Cenomanian).
Aïstopoda: an order of snake-like amphibians in the subclass Lepospondyli; rang-
ing from 2 inches to 3 feet in length, some had reduced limbs but most were limb-
less; they appeared 20 million years after the first tetrapods; North America and
Europe, Early Carboniferous to middle Permian.
Ajacicyathus: a genus of early colonial or solitary archaeocyathan sponges; global
distribution, early Cambrian.
Alamosaurus: “cottonwood lizard,” a monospecific genus of titanosaurian sauro-
pod dinosaurs in the family Saltasauridae. Known from many fragmentary speci-
mens and a nearly complete juvenile skeleton found in 2002, A. sanjuanensis was
among the last surviving dinosaurs and possibly the largest North American dino-
saur. It was first found in the Kirtland Formation of the San Juan Basin, New
Mexico, and other specimens have been found in Utah and Texas. Some estimates
claim that it may have weighed 80 tons and been 100 feet long. North America, Late
Cretaceous, 70–66 Ma.
Albanerpetontidae: a long-lived family of salamander-like lissamphibians in the
extinct order Allocaudata; fossorial, with a robust head and neck; North America
and Europe, Middle Jurassic to Pliocene.
Albertosaurus: a genus of large tyrannosaurid theropod dinosaurs, 27 feet long
and 2–3 tons in weight. Albertosaurus is characterized by two digits on the very
small manus and a skull more than 3 feet long with about 60 teeth. The most signifi-
cant find of Albertosaurus was made by Barnum Brown in 1910, in a bone bed on
the Red Deer River; excavations there over the next century have uncovered more
than 1100 bones from two dozen individuals. North America, Late Cretaceous,
70–66 Ma.
Aletopelta: a monospecific genus of armored dinosaurs in the family
Ankylosauridae; known only from a partial skeleton found in Southern California,
A. coombsi was about 20 feet long; named “wandering shield” because the forma-
tion in which it was found is a part of a tectonic plate that was several hundred miles
to the south in the Late Cretaceous; United States, Late Cretaceous, 85–71 Ma.
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allochthonous: referring to a fossil assemblage consisting of organisms removed

from their place of origin before burial, as opposed to an autochthonous assemblage
buried in situ; see also taphocoenosis.
Allodesmus: a genus of pinnipeds in the extinct family Desmatophocidae; 8 feet
long and about 800 pounds but sexually dimorphic (females were smaller);
Allodesmus had large eyes and was adapted to hearing underwater sounds; North
America and Asia, Miocene.
Allosaurus: a genus of dinosaurs that were very similar to the later Tyrannosaurus
rex; ranging from 20 to 32 feet long, the allosaurs were generally slightly smaller
than T. rex, but one of the later species (known only from a skull recently discovered
in Wyoming) was at least as large; mostly known from Western North America, but
also found in Portugal and possibly in Tanzania; Late Jurassic, 155–150 Ma.
Altirhinus: a monospecific genus of iguanodont ornithischian dinosaurs; charac-
terized by a tall beak on a rounded snout, A. kurzanovi was 21–26 feet long; it was
optionally quadrupedal and had five digits on the manus, including a thumb spike;
its arched snout is similar to that of Muttaburrasaurus, and the two share several
other similarities; Mongolia, Early Cretaceous, 125–100 Ma.
altriciality: of animals’ young: born relatively undeveloped and needing parental
care.
Alum Shale Formation: an upper Cambrian formation in Scandinavia; see Orsten
Lagerstätten.
Alveolites: a genus of colonial favositid corals in the order Tabulata; characterized
by compressed, thin-walled corallites in massive or branching bodies; Silurian to
Devonian.
Alveugena: a genus of non-placental mammals in the family Cimolestidae, possi-
bly ancestral to the taeniodonts; known only from two partial specimens; North
America, Early Paleocene, 66–63 Ma.
Amaltheus: a genus of ammonoids in the order Ammonitida and family
Amaltheidae; involute and up to 9 inches in diameter, it has strong ribbing and a
prominently braided keel; occurring boreally and only between the strata of its rela-
tives Prodactylioceras and Dactylioceras, it is an important index fossil in the
Pliensbachian; global distribution, Early Jurassic, 186–184 Ma.
Amargasaurus cazaui: a species of large sauropod dinosaurs from about 130 Ma,
about 30 feet long and 9 feet tall; known only from an almost complete skeleton
discovered in 1984, it is notable for a double row of long spines extending from the
vertebrae at the base of the skull along the entire long neck; Argentina, Early
Cretaceous.
amber: fossilized tree resin, resulting in an unusual sedimentary deposit. Although
they have the drawback of not being able to be handled, amber fossils of insects,
arachnids, and plants preserve exceptional detail. Amber deposits are the principal
10 A
source of spider fossils, and even though very few vertebrate fossils occur in amber,
bird feathers and even a nestling enantiornithine in amber are known from about
100 Ma. Since amber occurs near trees, sometimes far from the more common
aquatic depositional environments, it provides a unique sampling of the biota. The
earliest amber fossils are known from the Triassic, occurring in small amounts glob-
ally; but the Baltic amber of the Eocene has produced the greatest concentration of
fossils, with important discoveries also in Cenozoic Dominican and Burmese amber
deposits.
Ambiortus: a controversial genus of primitive birds; generally classified in the
Ornithomorpha, it is poorly known and phylogenetic relationships are unclear;
Early Cretaceous, about 130 Ma.
Amblypoda: a discarded term for a polyphyletic grouping of extinct ungulates
including the uintatheres and pantodonts.
Amblypygi: “blunt rump,” an order of chelicerate arthropods in the class
Arachnida; up to an inch long, they are known as whip spiders or tailless whip scor-
pions; with a flattened body and posture, they have long, slender legs and no silk
spinnerets; they have no tail or venomous fangs, but their spiny pedipalps can punc-
ture their prey; dating back to the Late Carboniferous (Graeophonus), about 160
species are presently known in tropical and subtropical areas; cosmopolitan, Late
Carboniferous to Recent.
Ambondro: a monospecific genus of early mammals in the family Henosferidae,
A. mahabo is probably close to the lineage of monotremes; Madagascar, Middle
Jurassic, 167 Ma.
Amborella: a still-extant genus of Early Cretaceous plants in the family
Amborellaceae, considered possible ancestors of angiosperms; global distribution,
130 Ma to Recent.
Ambulocetus: “walking whale,” a genus of archaeocete whales in the family
Ambulocetidae. About 10 feet long and more than 600 pounds, it was an important
transitional form between ancestral land-dwelling mammals and fully marine mam-
mals, one or more of which evolved into modern whales. Ambulocetus is character-
ized by such aquatic adaptations as a long tail and strong, paddle-like feet, but it still
has land-mammal characters such as hooves on its toes and powerful jaws with
shark-like teeth. It is related to Indocetus and Dorudon, and is probably ancestral to
Rodhocetus and Basilosaurus. Asia, Eocene.
Amebelodon: a genus of large shovel-tusked proboscideans in the family
Gomphotheriidae; about 10 feet high, Amebelodon was similar to Platybelodon,
another shovel-tusker; see also Gnathobelodon; Africa, Asia, and North America;
late Miocene.
Amia calva: a carnivorous freshwater fish, commonly called bowfin, once distrib-
uted globally but now known only in the Great Lakes and Mississippi Valley; char-
acterized by primitive ganoid scales; Paleocene to Recent.
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ammonite: a general term for ammonoid but sometimes also used to refer to the
later ammonoids of the order Ammonitida.
Ammonitida and Ammonitina: see Ammonoidea.
Ammonoidea: a subclass of externally shelled cephalopod mollusks, notable for
their chambered and coiled shells, generally with a ventral siphuncle. The ammo-
noids appeared in the Early Devonian and became important predators in the
Paleozoic and Mesozoic seas, the last becoming extinct toward the end of the
Cretaceous. The shells of almost all ammonoids follow a planispirally coiled pat-
tern, but in the course of their 350 million years of existence they developed an
unparalleled diversity of patterns and ornamentation, especially in the late Mesozoic.
Spreading into all parts of the oceans, they were generally good swimmers, nektic
or nektobenthic and preying on smaller marine organisms. The ammonoids are the
most important Mesozoic index fossils because of their wide geographic occur-
rence, their rapid diversification, and the persistence of a few genera through the
Permian and Triassic extinctions.
Ammonoids’ classification in orders and suborders is highly disputatious; the
version given here attempts to avoid controversy. They may be divided into eight
orders, in order of appearance: (1) the Agoniatitida, Middle to Late Devonian,
ancestral to all later ammonoids; (2) the Clymeniida, Middle to Late Devonian; (3)
the Goniatitida, Middle Devonian to Late Permian; (4) the Prolecanitida, Late
Devonian to Late Triassic; (5) the Ceratitida, Permian to Triassic; (6) the
Phylloceratida, Early Triassic to Late Cretaceous; (7) the Lytoceratida, Late Triassic
to Late Cretaceous; and (8) the Ammonitida, Early Triassic to Late Cretaceous.
These eight groups may also be placed in three large orders, based mainly on their
increasingly complex suture patterns: the Goniatitida, Ceratitida, and Ammonitida.
Two large suborders of the Ammonitida, the Ammonitina and Ancyloceratina,
are especially important biostratigraphically because of their generally short life-
spans and wide geographic range. The Ammonitina appeared very early in the
Jurassic and include 1000 known genera, organized in about 15 superfamilies. They
typically follow the regular planispiral coiling pattern, with just a few heteromorph
genera. The Ancyloceratina appeared in the Late Jurassic and persisted to the end of
the Cretaceous, as the Ammonitina declined. They include many heteromorph gen-
era of the Cretaceous, such as Bostrychoceras, Hamites, Nipponites, Scaphites, and
Turrilites. Most were relatively short-lived, and they were probably weak swimmers
because of their shape.
The earliest ammonoids evolved from unknown nautiloids near the beginning of
the Devonian, but collection failure (scarcity of specimens of nautiloids at the end
of the Silurian and in the Early Devonian, and of the early ammonoids in the
Devonian) leaves the connections unclear. The Phylloceratida (sometimes consid-
ered a suborder of the Ammonitida) probably gave rise to all post-Triassic ammo-
noids, such as the Lytoceratida. The Cretaceous ammonoids declined toward the
end of the period, and the last disappeared in the end-Cretaceous extinction. See
also Nautiloidea and aptychus.
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amniotes: animals (reptiles, birds, and mammals) which evolved ways of

p rotecting eggs inside the mother or at least inside a protective case; the basal amni-
otes were tetrapods that appeared in the Carboniferous about 312 Ma, descended
from some group of reptiliomorphs such as the diadectomorphs or seymouria-
morphs; since they made reproduction independent of water, the appearance of
amniotes was important in the development of terrestrial animal life.
Amphechinus: a genus of hedgehogs, small placental mammals in the still-extant
family Erinaceidae; it appeared in Europe and Asia in the Oligocene and spread
worldwide in the Miocene, becoming extinct at the end of the period.
Amphibamus: a genus of temnospondyl tetrapods in the superfamily
Dissorophoidea and family Amphibamidae; considered by some to be close to the
ancestry of modern amphibians, but the connection is in dispute; North America and
Europe, Middle Pennsylvanian.
Amphibia: a diverse class of vertebrates, including two extinct groups, the
Temnospondyli and the Lepospondyli, and one still-extant group, the Lissamphibia,
including the frogs, toads, salamanders, and other smaller groups.
Amphichelydia: an obsolete term for a suborder of cryptodire turtles.
Amphicyonidae: a family of dog-like carnivorous mammals in the superfamily
Arctoidea. Possibly related to the Ursidae, they generally ranged from 10 to 200
pounds, but the largest species, Amphicyon ingens, is estimated to have weighed
more than 1000 pounds. The amphicyonids achieved a holarctic distribution in the
Oligocene. North America, Europe, Asia, and Africa; Eocene to early Pliocene.
Amphilestidae: an enigmatic family of mammals in the order Eutriconodonta,
related to the Morganucodontidae and Gobiconodontidae; Africa, Asia, Europe, and
North America; Jurassic to Late Cretaceous.
Amphimerycidae: a family of artiodactyl ruminants in the suborder Tylopoda;
Europe, late Eocene to middle Oligocene.
Amphioxiformes: see Cephalochordata.
Amphipithecidae: a family of early primates in the suborder Haplorrhini; includes
Pondaungia and the more recently discovered Myanmarpithecus and Siamopithecus;
middle to late Eocene.
Amphipithecus: see Pondaungia.
Amphisbaenia: a group of limbless reptiles in the order Squamata, commonly
called worm lizards; global distribution except Australia, Cretaceous to Recent.
Amphissites: a large genus of straight-backed palaeocopan ostracods in the family
Amphissitidae; global distribution, Carboniferous to Permian.
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Amphistium: a genus of flatfish showing the movement, over millions of years, of

an eye first toward the top of the skull and then over to the upper side of the body;
Europe, early to middle Eocene; see also flatfish.
Amphitheriida: a poorly known order of early pantothere mammals in the super-
order Dryolestoidea; scansorial insectivores, they were probably in the line of evo-
lution toward placentals and marsupials; England, Middle Jurassic.
Amphiura plana: an extinct species of burrowing brittle stars in the still-extant
order Ophiurida and family Amphiuridae, known from the late Cenomanian to the
end of the Cretaceous; Europe, Late Cretaceous.
Amphoracrinus: a genus of crinoids in the family Amphoracrinidae, characterized
by a flat, truncate base; North America and Europe, Carboniferous.
Amynodontidae: a family of hornless perissodactyl mammals in the superfamily
Rhinoceratoidea; some genera resemble hippopotamuses in habit and appearance,
while others have a tapir-like appearance; North America, Europe, and Asia; late
Eocene to Pleistocene.
Anactinotrichida: see Acari.
Anagale: a genus of early eutherian mammals in the order Anagaloidea; rabbit-like
in appearance, it was probably insectivorous, burrowing for beetle and worms;
Mongolia, known only from the early Oligocene.
anagenesis: a concept briefly described as evolution within a lineage, while clado-
genesis is evolution leading to a splitting of the lineage; anagenesis is said to result
in change in a species through a gradual accumulation of genetic and physical dif-
ferences, without a branching-off of a new species (cladogenesis); anagenesis is
considered the basic process involved in phyletic gradualism, while cladogenesis is
basic to punctuated equilibrium.
analogous structures: body parts of different organisms that perform similar
functions but are not necessarily built on the same anatomical plan. For example,
wings of both insects and birds perform the same function and are analogous, but
they are not homologous because they have no basic structural parts (such as bones)
in common. See also homologous structures and vestigial structures.
Anancus: a genus of large proboscideans in the family Gomphotheriidae and sub-
family Anancinae; a forest browser 10 feet tall and weighing about 5 tons, with
strikingly long tusks in the upper jaw, as long as 13 feet; Africa, Europe, and Asia;
late Miocene to early Pleistocene.
anapsid: a term used to refer to reptiles that lack fenestrae behind the eye sockets
in the skull; traditionally, anapsids have been considered one of five main subclasses
of reptiles and as ancestral to diapsids and synapsids; the group is now recognized
as paraphyletic or even polyphyletic, and the term is used only informally.
14 A
Anarcestes: a genus of small ammonoids in the order Agoniatitida and family

Anarcestidae; an important index fossil in the Eifelian Stage of the Middle Devonian;
Africa, Europe, Asia, and North America.
Anarcestida: see Agoniatitida.
Anasca: a still-extant polyphyletic group formerly considered a suborder of bryo-
zoans in the order Cheilostomata; characterized by the absence of an ascus (a water-
filled sac with a hydrostatic function) in each zooid, but being reclassified into
separate orders on the basis of other characters; Late Jurassic to Recent.
Anaspida: an order of small, jawless, fish-like vertebrates usually considered
agnathid ostracoderms; although they have no more than a cartilaginous skeleton
and only a weak headshield, some have an armor of bony scales; also characterized
by a reversed heterocercal tail fin; early Silurian to Early Devonian.
Anatolepis: a possible early jawless fish, proposed as a heterostracan vertebrate,
but known only from tooth and carapace fragments and incertae sedis; late Cambrian
to Early Ordovician.
Anatosuchus minor: a small species of Cretaceous crocodiles, about 3 feet long
and with a pointed, duckbill-like snout; Niger, 110 Ma.
Anchiornis huxleyi: a species of small, bird-like theropod dinosaurs. Probably
flightless, it was less than 2 feet long, with long legs, 4 toes, and proportionally large
forelimbs. It was almost covered in feathers, some colored, according to researchers
using melanosomes, cells that contain the pigment melanin. The color itself is not
visible in the fossil, but electron-microscope analysis of follicle structure gives indi-
rect evidence of color. Extrapolating from colors associated with the shape of the
melanosomes in modern birds, researchers believe Anchiornis was covered with
black and white plumage but also had a reddish crest of feathers. China, Late
Jurassic, 161 Ma.
Anchisaurus: a genus of small basal sauropodomorph dinosaurs in the family
Anchisauridae; formerly called Megadactylus and Amphisaurus; from 7 to 8 feet
long and 60 to 80 pounds, it browsed on small palm trees and large shrubs; known
definitively only from North America, Early Jurassic, 190–174 Ma.
Anchitherium: a genus of horses in the extinct subfamily Anchitheriinae, related
to Mesohippus and Sinohippus; about 2 feet high at the shoulder, it originated in
North America in the Miocene and migrated to Asia, where it became extinct in the
Pliocene.
Ancodonta: a discarded term for a polyphyletic grouping of some primitive artio-
dactyl mammals.
Ancyloceratina: “bent horn,” a suborder of mostly heteromorph ammonoids in the
order Ammonitida; see Ammonoidea.
A 15
Ancylopoda: a suborder of perissodactyl ungulate mammals characterized by

long, curving, cleft claws; it includes the eomoropids and the chalicotheres in the
superfamily Chalicotherioidea; holarctic distribution, Eocene to early Pleistocene.
Ancylotherium: a genus of very large chalicotheres in the subfamily Schizotheriinae;
it was about 7 feet tall and weighed as much as 1000 pounds; Europe, Asia, and
Africa; late Miocene to early Pleistocene.
Ancyrodella rotundiloba: a species of conodonts whose first appearance marks the
beginning of the Late Devonian at 382.7 Ma; its extinction, along with that of the
conodont genus Ozarkodina, marks the end of the Frasnian Age of the Late Devonian
at 372.2 Ma.
Andalgalornis: a monospecific genus of large, flightless predatory birds in the
family Phorusrhacidae; A. steulleti was about 90 pounds and 4–5 feet tall, and had
a large, heavy skull; related to the larger Kelenken; northwestern Argentina, Miocene
to Pliocene.
Andrewsarchus: a monospecific genus of very large carnivorous or scavenging
artiodactyl mammals; known only from a single skull almost 3 feet long, A. mongo-
liensis was found by Roy Chapman Andrews, who estimated the body to have been
about 12 feet long; it was formerly classed as a mesonychid but is now considered
an artiodactyl, most closely related to hippopotamuses and whales; Mongolia, mid-
dle Eocene.
Anfesta stankovskii: an obscure species of trilobozoans; similar in shape to
Tribrachidium, it is circular and less than an inch in diameter, with three straight
radiating ridges; Russia, Ediacaran.
angiosperms: “vessel seed,” the flowering plants, characterized principally by
seeds enclosed in an ovary. Their ancestors have obscure roots in the Triassic (see
Sanmiguelia), and among the groups of early plants considered possible ancestors
are the Nympheales, Amborellaceae, and the disputed order Illiciales. The first
definitive angiosperms appeared in the Early Cretaceous and quickly spread world-
wide. By the beginning of the Late Cretaceous, they were developing showy petals
and layers of fruit to attract the insects and herbivores that were diversifying along-
side the flowering plants that they fed on. As a result of this symbiotic lifestyle, the
angiosperms have been the dominant plants globally since 100 Ma. See also
Amborellaceae, coevolution, Gnetales, and Archaefructus.
Anhanguera: a genus of pterodactyloid pterosaurs that flourished in Northern
Brazil about 112 Ma; with a wingspan of up to 15 feet, Anhanguera is a close rela-
tive of Ornithocheirus, and like some other ornithocheirids it had a pair of rounded
crests at the top and bottom of a long beak; known only from the Araripe Basin of
Brazil, Early Cretaceous.
Animantarx: “animated citadel,” a monospecific mid-Cretaceous genus of ankylo-
saurian dinosaurs in the family Nodosauridae; about 10 feet long, A. ramaljonesi
16 A
was heavily armored with bony scutes and horns but no tail club; known from only
a single set of remains; Utah, Early to Late Cretaceous, 104–98 Ma.
anisograptid fauna: see Graptolithina.
Ankarapithecus meteai: a species of fruit-eating apes that inhabited Asia Minor
about 10 Ma; about 60 pounds, Ankarapithecus is known from a fragmentary male
specimen found in 1950 and an almost complete face discovered in the 1990s; with
some similarities to Sivapithecus, the finds are important because they are among
the few primate fossils of the period between 18 Ma and 5 Ma; Turkey, late Miocene.
Ankylosauria: a suborder of squat, heavily armored ornithischian dinosaurs that
contains the families Nodosauridae and Ankylosauridae; see also Polacanthus;
global distribution, Late Jurassic to Late Cretaceous.
Ankylosauridae: a family of medium-sized ornithischian dinosaurs in the subor-
der Ankylosauria. Mainly herbivorous, squat, and quadrupedal, some grew to
20 feet long. Ankylosaurs are characterized by encasing armor plates, a fringe of
pointed spines around the midriff of the body, and a tail club; some species had
plates even over the eyelids. North America, Europe, Asia, and Australia; Cretaceous,
125–66 Ma.
Annelida: the phylum of segmented worms, including the subclasses Oligochaeta
(earthworms) and Hirudinea (leeches), and the paraphyletic class Polychaeta. The
latter are marine invertebrates with a segmented body and many bristles; their chi-
tinous jaws, called scolecodonts, are a widespread fossil from the Cambrian to the
present.
Anodontosaurus: “toothless lizard,” a monospecific genus of ankylosaurid dino-
saurs in the subfamily Ankylosaurinae; dated between 73 and 67 Ma, A. lambei is
known from partial specimens found in the Horseshoe Canyon Formation of Alberta;
medium-sized, armored, and herbivorous, it has a wide, pointed tail club; Canada,
Late Cretaceous.
Anomalocarididae: a family of soft-bodied animals of the Cambrian and
Ordovician in the order Radiodonta. Anomalocaris, at 3–4 feet the largest predator
in the Burgess Shale assemblage, has been a contentious stem-arthropod for decades,
but is now well-known from widespread specimens, including some larger speci-
mens in China and some from Kangaroo Bay in Australia. Both its mouth and its
body were at first incorrectly identified as separate animals, “Peytoia” and
“Laggania” respectively, but Peytoia is now accepted as the name of another anom-
alocaridid, related to Hurdia. North America, Europe, Asia, and Australia; middle
Cambrian to Early Devonian.
Anomphalus: a genus of marine aspidobranch snails in the superfamily Trochoidea
and family Anomphalidae; type species A. rotalus; global distribution, Early
Devonian to Jurassic.
A 17
Anoplotheriidae: a family of artiodactyl ungulate mammals in the suborder

Tylopoda and extinct superfamily Anoplotheroidea; related to camels, they were
medium-sized forest browsers; Europe, Eocene to Oligocene.
anoxia: a chemical state characterized by a low level of oxygen, sometimes a com-
plete lack of oxygen; anoxia favors the preservation of fossils.
Anseriformes: an order of waterfowl in the superorder Galloanserae; it includes
many modern species as well as extinct groups such as the Gastornithidae and the
earliest-known genus, Vegavis; Cretaceous to Recent.
Antarctodolops: a genus of primitive marsupials tentatively placed in the family
Polydolopidae; the only marsupial yet known from Antarctica, it is represented by
the single species A. dailyi; Eocene to possibly Oligocene.
Antetonitrus: a monospecific genus of sauropodomorph dinosaurs that lived
between 221 and 210 Ma, found in 1981 but only in 2003 recognized as one of the
oldest sauropods known; about 30 feet long and almost 2 tons, A. ingenipes was the
largest land animal of its time, although later sauropods came to weigh as much as
100 tons; found in the Elliot Formation along with the prosauropod Melanorosaurus;
South Africa, Late Triassic.
Anthocyathea: a class of solitary, occasionally colonial, organisms placed in the
Archaeocyatha; characterized by a roughly cylindrical cup and numerous bars, rods,
and tabulae; early to late Cambrian.
Anthozoa: “flower animals,” one of the three classes of cnidarians, including prin-
cipally the corals and sea anemones; the class contains the subclasses Hexacorallia,
Octocorallia, and Ceriantharia.
Anthracosauria: an obscure group of early amphibian tetrapods, probably para-
phyletic, including the Embolomeri, Seymouriamorpha, and Diadectomorphs; their
classification is widely debated, but they are generally considered a stem group in
the lineage leading to the amniotes; Late Devonian or Middle Mississippian to Early
Triassic.
Anthracosia: see Carbonicola.
Anthracotheriidae: an extinct family of artiodactyl ungulates, pig-like but related
to hippopotamuses and possibly whales; North America, Europe, Asia, and Africa;
middle Eocene to late Pliocene.
anthropoid: an informal term referring to the monkeys, apes, and humans; gener-
ally replaced by more specific terms; see Haplorrhini, Strepsirrhini, Catarrhini,
and Platyrrhini.
Anthropornis: a genus of giant penguins almost 6 feet long; New Zealand and
Antarctica, late Eocene to early Oligocene.
Antiarchi: an order of small, heavily armored freshwater placoderm fish; the ante-
rior body is armored while the posterior is fish-like, characterized by a pair of small,
18 A
moveable pectoral appendages; the order is well-known from the genera Bothriolepis
and Yunnanolepis; global distribution, early Silurian to Late Devonian.
Antilocapridae: “goat antelope,” a still-extant family of ruminant artiodactyl
mammals. Similar in size and shape to antelopes, they are distinguished from other
artiodactyls by their branching horns; there is a permanent bony core like that of
bovid horns, and the horny sheath of the horn is shed annually. Four genera are
known from the early Miocene to the end of the Pleistocene, but the only living spe-
cies is Antilocapra americana, ubiquitous in Western North America since the
Pleistocene. The Miocene genus Ramoceros had long, cervid-like forked horns with
3 or 4 tines, and other genera had a variety of horn configurations. Restricted to
North America, early Miocene to Recent.
Anura: “tailless,” the frogs, an enigmatic order of amphibians whose evolutionary
history is unclear because of the paucity of the fossil record; the order includes the
family Bufonidae, the toads; see Proanura; global distribution, Jurassic to Recent.
Anzu wyliei: a species of Late Cretaceous saurischian dinosaurs in the family
Caenagnathidae; the largest oviraptorosaur of North America, it weighed about 650
pounds and was about 11 feet long and 10 feet high; it had a toothless beak and
prominent crest, and was probably feathered; see also Gigantoraptor; Hell Creek
Formation, North and South Dakota; late Maastrichtian, 66 Ma.
Apatemyidae: a family of small Paleocene mammals in the extinct order Cimolesta
and suborder Apatotheria; characterized by long, hooked teeth, they were probably
insectivorous and arboreal; North America and Europe, Paleocene to Eocene.
Apatornis: “deceptive bird,” a genus of flying birds in the family Apatornithidae;
generally considered neornithine and possibly close to the Anseriformes, but known
from only one fragment found in Kansas; North America, Late Cretaceous, 84 Ma.
Apatornithidae: a contentious family of very early flying birds, generally placed
in the order Ichthyornithiformes, the Cretaceous toothed birds; its exact affinities,
however, are still in question because of a paucity of specimens; it seems to have
had a well-developed keel and other flight features; North America, Late Cretaceous.
Apatosaurus: a genus of very large saurischian dinosaurs in the infraorder
Sauropoda and family Diplodocidae; with a long neck and a long, whip-like tail,
they are estimated to have been up to 75 feet long and to weigh as much as 30 tons;
see also Brontosaurus; North America, Late Jurassic, 152–151 Ma.
Aphrosalpingoidea: an enigmatic superfamily of sponge-like sessile animals gen-
erally classified as archaeocyathans; they had conical or cylindrical cups 1–2 inches
wide; Europe, late Silurian.
Apidium: see Parapithecidae.
Apinae: the bees, a still-extant subfamily of hymenopteran insects in the super-
family Apoidea. The taxonomic classification of bees is widely debated, but they are
generally accepted as monophyletic, with Apinae including all genera of bees as
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well as several lesser-known insects. The earliest specimens are known from Early
Cretaceous amber fossils (see Melittosphex), with possible bee nests found in
paleosols as far back as the Late Triassic. Along with other pollinating insects, bees
spread globally alongside angiosperms in the early Late Cretaceous.
Apiocrinites: a genus of crinoids in the still-extant order Millericrinida, character-
ized by a long stem and a rigid, pear-shaped calyx; Apiocrinites is found now only
in an abyssal form, although it was common in the Jurassic in a shallow-water envi-
ronment; Middle Jurassic to Late Cretaceous.
Aplacophora: “bearing no shell,” a still-extant class of benthic, deep-ocean mol-
lusks; mostly small, microscopic to 2 inches, but a few grow up to several inches;
cylindrical and generally without shells (but see Kulindroplax), and their body is
covered with small aragonitic spicules; they are carnivorous, burrowing in the sub-
strate or feeding on cnidarians; global distribution, early Cambrian to Recent.
apomorphy: a derived trait, a novel specialized feature of an organism; see also
synapomorphy.
Apsaravis ukhaana: a species of primitive birds in the suborder Theropoda, a link
in the chain of descent from dinosaurs to birds; Mongolia, Late Cretaceous,
80–75 Ma.
apterygote: referring to wingless insects.
Apthoroblattina: a genus of cockroaches in the extinct family Archimylacridae,
about 2 inches long and 1.5 inches wide; North America, Europe, and Russia;
restricted to Pennsylvanian.
aptychus: a curved calcitic or horny plate, usually one of a pair of aptychi; often
found with ammonoids, sometimes attached to the body at the aperture, perhaps
combining the functions of an operculum and a jaw; often found singly and referred
to as an anaptychus.
Arabidopsis thalianai: thale cress, a living member of the mustard family, the
Brassica, which includes cabbages and radishes. A model plant for genetic research,
it is significant for having a small genome and for its genetic similarity to many
other flowering plants, including corn, wheat, and rice. Arabidopsis offers further
proof that the genes that control cell division are similar in plants and animals, sup-
porting the inference that plants and animals originate from a common ancestor,
probably a one-celled organism from about 1.5 billion years ago. It was originally
named Pilosella siliquosa, and its name has changed several times since.
Arachnida: a large class of arthropods in the subphylum Chelicerata; the most
familiar order is the Araneae, the spiders, and others are scorpions, ticks, mites, and
the extinct trigonotarbids; all arachnids have 8 legs, although in some families one
pair has been converted to other functions; early Silurian to Recent.
Araeoscelis: a genus of small, lizard-like diapsid reptiles in the order Araeoscelidia;
2 feet long, it was one of the earliest diapsids but was exceptional in that both parts
20 A
of one pair of temporal fenestrae were closed with bone; similar to Petrolacosaurus
of the Late Pennsylvanian; North America, early Permian.
aragonite: a polymorph mineral of calcium carbonate, chemically identical to cal-
cite but with an orthorhombic crystal structure, different from calcite’s rhombohe-
dral structure; aragonitic mollusk shells fossilize less readily than calcite shells
because of their unstable structure, resulting in gaps in the fossil record.
Arandaspis: a genus of jawless heterostracan fish in the order Pteraspidiformes;
dated at 465 Ma, it was perhaps the oldest known vertebrate; it was 6–8 inches long
and finless, and was armored with large bony plates and straps rather than with the
tiles seen in its cousin Astraspis; Australia and South America, Middle Ordovician.
Araneae: the true spiders, an extant order of arachnids with eight legs, spinnerets
for web-weaving, and cheliceral venom glands; several early genera date back to the
Carboniferous, and the order diversified greatly in the Jurassic, but 90 percent of the
known spider fossils are in amber from the Paleogene; the oldest known spider web
fragments (140 Ma) are encased in a piece of amber from Sussex, England; global
distribution, Devonian to Recent.
Araneoidea: the larger of the two still-extant superfamilies of orb-weaving spiders
in the order Araneae; araneoids weave webs of adhesive threads and share several
morphological characters for spinning; see also Deinopoidea; middle Paleozoic to
Recent.
Araripemys: a genus of early turtles in the order Testudines and extinct family
Araripemydidae or Pelomedusidae; type species A. barretoi, about 2 feet long but
not well-known; Brazil, Early Cretaceous.
Araripe Plateau and Basin: a region in Northeast Brazil that has produced a large
and diverse fossil assemblage of plants and animals from the Early Cretaceous,
before South America split off from Africa. See Crato and Santana Formations.
Araripesuchus gomesii: the type species of Araripesuchus, a notosuchian croco-
dyliform of Gondwana; a short-snouted crocodilian about 2 feet long, found in the
Santana Formation of the Araripe Basin in Brazil; Early Cretaceous.
Araripesuchus rattoides: the second African species of Araripesuchus to be found;
3 feet long, characterized by buckteeth which it used to root in soil for grubs and
plants; probably spent most of its time on land; Morocco, Early Cretaceous.
Araripesuchus tsangatsangana: the geologically most recent Araripesuchus,
dated at 66 Ma; Madagascar, Late Cretaceous.
Araripesuchus wegeneri: the first African species of the genus; it ran on long legs
like a dog and swam well with a long crocodilian tail; about 3 feet long and omnivo-
rous; Niger, Early Cretaceous.
Araucaria: a genus of plants in the still-extant family Araucariaceae, distantly
related to pines. Several genera have existed almost unchanged for hundreds of
A 21
illions of years and are considered “living fossils,” such as the Norfolk Island pine
m
(A. heterophylla) and the monkey puzzle tree (A. araucana), which is known from
as far back as 250 Ma. Araucaria appeared in the Early Triassic and diversified
rapidly worldwide, creating forests of trees as tall as 250 feet. Triassic to Recent.
Araucariaceae: a still-extant family of coniferous trees in the class Pinopsida and
order Pinales; appearing in the Early Carboniferous, they spread worldwide in the
Mesozoic but disappeared in Laurasia at the end of the Cretaceous; a notable site for
them is Petrified Forest National Park in the Southwestern United States; global
distribution, Carboniferous to Recent.
Archaea: one of the three basic domains of life forms, a group of prokaryotes,
single-celled organisms without a defined nucleus; distinguished from bacteria, the
other prokaryotic domain, by some eukaryotic genes and metabolic characteristics;
notable for tolerance of extreme environments, many of them living near hydrother-
mal vents in the ocean floor; the earliest-known fossilized Archaea date back to 3.5
billion years ago, but genetic research indicates that they probably originated closer
to 4 billion years ago; see also Bacteria and Eukaryota.
Archaefructus liaoningensis: the best-known of three species of a genus of early
(125 Ma) flowering plants in the Jehol biota, proposed as stem-group angiosperms
but still poorly known; China, Early Cretaceous.
Archaeocalamites: a genus of the earliest-known horsetails in the order
Calamitales (q.v.); Pennsylvanian to Permian.
Archaeoceratops: a genus of basal neoceratopsian dinosaurs in the family
Archaeoceratopsidae; only 3 feet long, it had no horns or neck frills but a large skull
and a small bony frill near its tail; the type species A. oshimai, the oldest known
neoceratopsian, is dated at about 125 Ma; China, Early Cretaceous.
Archaeoceti: a paraphyletic suborder of ancestral cetaceans; fish-eating and
dolphin-like, they grew up to 33 feet long; characterized by vestigial hindlimbs and
an elongated snout; the earliest are known from the island continent of India in the
early Eocene (see Pakicetus and Ambulocetus), and they diversified rapidly world-
wide; early Eocene to late Oligocene.
Archaeocidaris: a genus of early euechinoids in the extinct order Cidaroida; global
distribution, Late Devonian to late Permian.
Archaeocopida: a problematic early Paleozoic order of ostracods, characterized
by a thin carapace and a shell with high-chitin content, only slightly calcified;
Cambrian to Ordovician.
Archaeocyatha: “ancient cups.” The archaeocyathans were sessile and sponge-
like marine organisms, mostly solitary but some colonial. Their classification has
shifted considerably over time, with the current consensus placing them with the
hypercalcified sponges. Among the earliest reef-builders, they were solitary or colo-
nial, generally cup-shaped or conical, with two layers of perforated calcareous
22 A
walls, mostly between 1 and 6 inches tall. Prominent in the Tommotian biota, they
were worldwide and highly diverse index fossils for the early Cambrian but began
to decline by the middle Cambrian. Most genera had disappeared by the end of the
Cambrian, and the last-known species, Antarcticocyathus webberi, became extinct
just before the end of the period. After the disappearance of Archaeocyatha, there
were almost no reef-building animals until the rise of the Late Ordovician reef-
builders such as the stromatoporoids and stenolaemates 50 million years later.
Archaeocyon: a genus of small early dogs in the extinct subfamily Borophaginae;
up to 2 feet long and weighing 3–4 pounds; Western North America, early to late
Oligocene.
Archaeodontosaurus descouensi: a species of sauropod dinosaurs with
prosauropod-like teeth, found in 1999 and dated at 168 Ma; it is known only from
bone fragments and teeth and estimated to have been 40 feet long; Madagascar,
Middle Jurassic.
Archaeogastropoda: a large and still-extant paraphyletic group of sea snails and
limpets, prosobranch gastropods that first appeared in the Cambrian and have been
very widespread and diverse since a mid-Paleozoic radiation; most are extinct, but
limpets and many sea snails remain; there are several competing classification sys-
tems of the group.
Archaeognatha: a still-extant order of apterygote insects; also called Microcoryphia
and previously considered part of the now obsolete order Thysanura; among the
most primitive insects, they appeared about 390 Ma; characterized by a long, slen-
der body, arched back, and two cerci and a medial extension on the tail; cosmopoli-
tan, Middle Devonian.
Archaeolynthus: a genus of primitive archaeocyathans in the order Monocyathida;
solitary or colonial and branching, the small, single-walled Archaeolynthus was one
of the simplest archaeocyathans; Cambrian.
Archaeopteris: “ancient feather,” a genus of progymnosperm trees in the order
Archaeopteridales; growing up to 25 feet high, Archaeopteris created the first large
forests in the Devonian and was an important ancestor of later plants; Archaeopteris
had fern-like true leaves and was similar to modern trees in its branching growth
pattern; its significant characteristics include heterospory and dense gymnospermic
secondary wood; global distribution, Late Devonian to Early Carboniferous.
Archaeopteryx: a famous genus of feathered theropods, generally considered tran-
sitional between dinosaurs and modern birds; known principally from a dozen spec-
imens from the Solnhofen limestone quarry in Central Bavaria and dated at 150 Ma;
modern bird physiology seems to have first appeared in later feathered dinosaurs
found in China, about 130 Ma; about the size of a raven, Archaeopteryx was a weak
flyer; Europe, Late Jurassic.
“Archaeoraptor liaoningensis”: one of several fake fossil specimens produced in
China in recent decades, supposedly a turkey-sized feathered dinosaur capable of
A 23
flight. Publicized in 1999 by National Geographic, it was soon discovered to be a

composite of bones from different animals; the magazine published a full retraction
later, but the hoax has joined Piltdown Man as one of the most famous frauds in the
history of science. Many genuine feathered dinosaurs have been found since then,
mostly in China, now undergoing a much more thorough investigation than in the
1990s.
Archaeornithomimus: a genus of coelurosaurian theropod dinosaurs in the family
Ornithomimidae; about 11 feet long and probably omnivorous, Archaeornithomimus
is known from disarticulated skeletal specimens of several individuals; Asia, Late
Archaeotherium: a genus of artiodactyl mammals in the family Entelodontidae;
known only from North America, Archaeotherium was carnivorous but mainly a
scavenger; the size of a bison but looking more like a giant warthog, Archaeotherium
closely resembled its later relative Daeodon; North America, late Eocene to late
Oligocene.
Archaeothyris: a genus of small, lizard-like synapsids in the family
Ophiacodontidae; less than 2 feet long, Archaeothyris resembled a small iguana; the
three known specimens are dated between 311 and 307 Ma, and it is the earliest
accepted synapsid, but see also Echinerpeton; Canada, United States, and Czech
Republic; Late Pennsylvanian.
Archanodon: a genus of large bivalves in the order Unionoida; the earliest-known
freshwater bivalves; North America, Middle Devonian.
Archelon: a monospecific genus of marine turtles in the still-extant suborder
Cryptodira and extinct family Protostegidae; the largest known turtle, about 13 feet
long and 16 feet wide flipper to flipper; instead of the solid shell of most modern
turtles, A. ischyros had a skeletal framework that supported a leather carapace;
Archeria: a genus of primitive anthracosaurians in the family Archeriidae; about
7 feet long, it was an aquatic eel-like predator; notable for being one of the few
almost complete skeletons of its type; Texas and Oklahoma, early Permian.
Archicebus achilles: the earliest-known haplorrhine primate, an almost complete
specimen discovered in China around 2005 and dated on the cusp of the Paleocene
to the Eocene; about 9 inches long, including a 5-inch tail, it weighed only an ounce
but is characterized by a distinctive heel bone and by marmoset-like teeth; 55 Ma.
Archimedes: a genus of stenolaemate bryozoans in the order Fenestrida and family
Fenestellidae; very different in appearance from other fenestellids, Archimedes is
named for its shape, which closely resembles an Archimedes Screw; growing in
upright colonies up to 3 feet high, with a fenestrate network at the edges of the
screw; global distribution, Middle Mississippian to middle Permian.
24 A
Archipolypoda: a superorder of millipedes in the subclass Chilognatha, possibly

the earliest terrestrial animals; Europe and North America, Silurian to Late
Carboniferous.
Archonta: a proposed superorder that would include primates, Plesiadapiformes,
tree shrews, and dermopterans.
Archosauria: a clade of advanced diapsid reptiles including modern crocodiles
and birds as well as the extinct thecodonts, pterosaurs, and dinosaurs. The earliest
archosaurs were the Proterosuchidae, which appeared near the end of the Permian
and diversified rapidly in the Early Triassic. The archosaurs were widespread and
dominant throughout the Mesozoic, but only a few families survived the mass
extinction at the end of the Cretaceous. Global distribution, Permian to Recent.
Arcovenator: a monospecific genus of ceratosaurian theropod dinosaurs in the
family Abelisauridae; A. escotae is known only from a partial skull but seems to
have most of the abelisaurid characters; dated at 74 Ma, it was probably part of a
northward migration of the Gondwanic abelisaurs toward the end of the Cretaceous;
France, Late Cretaceous.
Arctocyonidae: a possibly polyphyletic family of primitive, unspecialized mam-
mals; ranging in size from raccoon-like (Chriacus) to bear-like (Mentoclaenodon
and Arctocyon), the group presents a combination of carnivore and ungulate charac-
teristics; North America and Europe, Late Cretaceous to early Eocene.
Arctodus: a genus of large and very long-limbed bears in the subfamily
Tremarctinae, which became extinct near the end of the Pleistocene; A. simus, the
Great North American short-faced bear, was larger than the modern grizzly, 4 to
5 feet high at the shoulder and weighing almost a ton; North America, Pleistocene.
Arctolepis: a genus of early arthrodire placoderm fish in the family Phlyctaeniidae;
less than a foot long, it is characterized especially by its long and stout pectoral fins
and by its heavy anterior armor; also called Acanthaspis; North America and Europe,
Early to Middle Devonian.
Arctotypus: a genus of griffinflies in the order Meganisoptera, with a compact and
relatively heavy body, and a wingspan of about 12 inches; Europe and Asia,
Carboniferous to Permian.
Arcyzonidae: a family of paleocopan ostracods in the superfamily Kirkbyoidea;
characterized by valves with a central pit; known only from China, Early Devonian.
Ardipithecus: Ar. ramidus is a species of hominin known principally from a partial
female skeleton found in 1992 in the Middle Awash region of Ethiopia, dated about
4.4 Ma. It is closely related to the last ancestor shared by chimpanzees and humans,
and may be a direct ancestor of the australopithecines. It was omnivorous, not just a
frugivore, and had a flexible palm and possibly bipedal feet, although it may have
nested in trees. The discovery in a more heavily forested area than most hominid
skeletons has led Tim White and others to question the open-savannah theory of
A 25
early hominids’ development of bipedalism. Other paleontologists, however,

question whether Ardipithecus lived in a closed forest setting; others question its
classification as a hominid, and still others question whether it was bipedal. A par-
tial skeleton whose skull was in coin-sized pieces is kept in the National Museum of
Ethiopia. A second possible specimen, dated between 5.8 and 5.2 Ma and named Ar.
kadabba, is known from teeth and several skeletal bones. Miocene to Pliocene.
Argentavis: a monospecific genus of very large predatory and scavenging birds in
the family Teratornithidae; weighing up to 170 pounds, A. magnificens had a wing-
span of about 23 feet, similar to the smaller seabird Pelagornis sandersi; Argentina,
late Miocene.
Argentinosaurus huinculensis: one of the largest land animals known, a species
of sauropod dinosaurs in the clade Titanosauria; although only a few vertebra and
other bones have been found, A. huinculensis may have weighed more than 80 tons
and reached more than 130 feet in length, with a long neck and tail; Patagonia, Late
Arkanites: a genus of goniatitic ammonoids in the family Reticuloceratidae,
related to Retites and Quinnites; subglobose, with prominent ribbing and sometimes
nodular bases; known only from the Midwestern United States, Early Pennsylvanian.
Arkarua: an Ediacaran genus tentatively placed in Echinodermata but still of
uncertain relationships; somewhat similar to the trilobozoans; known only from a
cast half an inch in diameter, it has a raised center with pentameral symmetry;
Australia, Ediacaran.
“Arpylorus antiquus”: a supposed species of protists long considered a possible
early dinoflagellate but rejected in 2012, being designated as part of a more complex
organism.
Arsinoitherium: a genus of large paenungulate mammals in the extinct order
Embrithopoda and family Arsinoitheriidae, known mainly from a complete skeleton
found in the Fayum Depression; growing up to 10 feet long, they are related to ele-
phants despite having a very different skull, which bears two very large bony horns
extending forward from its nose; possibly related specimens have also been found
in Europe and Mongolia but are still being investigated; Egypt and Arabian
Peninsula, Eocene to Oligocene.
Arthrodira: “joint-necked,” a large order of placoderm fish; characterized espe-
cially by a ball-and-socket joint behind the head that allowed it to raise its head,
enlarging its gape; generally armored and at first only a few inches long, many
became very large during the Devonian, reaching a length of 30 feet in Dunkleosteus;
the arthrodires were among the dominant predators of the Devonian until their
extinction at the end of the period; Silurian to Late Devonian.
Arthropleura: a genus of Paleozoic millipedes in the extant subphylum Myriapoda
and the extinct subclass Arthropleuridea; arthropleurids were 1–7 feet long and up
26 A
to a foot wide, the larger species being the largest terrestrial invertebrates known;
North America and Europe, Carboniferous to early Permian.
Arthropoda: “jointed foot,” the largest phylum of animals, including about 80
percent of all described extant species; characterized by a segmented body, jointed
appendages, and an exoskeleton (a chitinous cuticle); among its subphyla are trilo-
bites, crustaceans, myriapods, chelicerates, and hexapods (which includes insects);
early Cambrian to Recent.
articulate: of brachiopods and bivalves, using teeth or sockets to hold valves
together, as opposed to using muscles to secure the joint in the manner of inarticu-
late species.
Artiodactyla: an order of herbivorous hoofed mammals with an even number of
toes, the most successful group of large mammals. Evolved from condylarths, the
group includes most large land mammals, such as the pig, sheep, hippopotamus,
pronghorn, deer, cow, giraffe, camel, llama, and bison, and dozens of species of
antelope. Since the cetaceans (whales, dolphins, and porpoises) evolved from early
even-toed ungulates, they are also considered to belong with artiodactyls in the
clade Cetartiodactyla. Early Eocene to Recent.
Asaphida: a large order of trilobites in the subclass Librostoma, containing about
20 percent of the known trilobite species; although the order includes a great num-
ber of disparate shapes and sizes, its members are generally isopygous and have a
prominent ventral suture; the superfamily Asaphoidea is also characterized by
prominent eye-stalks; the asaphids declined at the end of the Ordovician, with only
one of the six superfamilies, the Trinucleoidea, surviving into the Silurian; global
distribution, middle Cambrian to late Silurian.
Asaphus: a genus of large trilobites in the order Asaphida and family Asaphidae;
isopygous and rounded, they grew up to 2 feet long; Northern Europe, Early to
Middle Ordovician.
Ascidiacea: a paraphyletic class of soft-bodied animals in the subphylum Tunicata;
because of their scanty fossil record, there are only a few reliably identified fossils;
the oldest reported is from the early Cambrian Chengjiang biota of China; among
the later definitive fossil specimens are some specimens from the Jurassic and the
still-extant genus Cystodytes, known from the Eocene; early Cambrian to Recent.
Ascomycota: “sac fungus,” the largest division of the Kingdom Fungi, generally
characterized by a spore-producing ascus; known from Devonian fossils but believed
to be much older.
Asilisaurus: a genus of very early archosaurian dinosauromorphs in the clade
Dinosauriformes and family Silesauridae; quadrupedal, 3–10 feet long, and weigh-
ing as much as 70 pounds, Asilisaurus displays many dinosaurian traits; Tanzania,
Middle Triassic, 245 Ma.
A 27
Asioryctitheria: a proposed order of basal eutherian mammals; insectivorous and

probably the earliest modern mammals; includes the genera Asioryctes and
Kennalestes and the proposed genera Sasayamamylos and Ukhaatherium; China
and Japan, Early to Late Cretaceous.
Aspidella: an enigmatic genus of discoidal fossils of the late Precambrian, well-
known from the Ediacaran fauna of Canada and Europe. Generally 3–4 but up to
7 inches wide, it has a concentric ring structure and seems to be the imprint of an
anchoring disk of an organism with fronds. Aspidella terranovica is the name of the
first Ediacaran organism to be described, in Newfoundland in 1872. See also
Charnia.
Aspidorhynchiformes: a Mesozoic order of actinopterygian fish in the subclass
Neopterygii; about 2 feet long with an elongated and tooth-lined jaw, they superfi-
cially resemble the modern gars but are not related; North America, Europe, and
Asia; Middle Jurassic to Late Cretaceous.
Asteraceae: the largest family of angiosperm plants; also called Compositae, they
are characterized by composite flowers with flowering heads surrounded by bracts;
Late Cretaceous to Recent.
Asteroidea: the sea stars or starfish, a class of asterozoan echinoderms; character-
ized by radial symmetry, the central body extending into five or more arms, not
sharply marked off from the body; since sea stars disintegrate quickly after death,
the fossil record is weak; global distribution, Ordovician to Recent.
Asterozoa: a subphylum of echinoderms; see Asteroidea and Ophiuroidea.
astogeny: the growth and development of colonial organisms such as corals; see
also ontogeny.
Astrapotheria: an order of hoofed mammals in the superorder Meridiungulata; up
to 10 feet long, the larger species resembled a small, tapir-like mastodon, with long,
tusk-like canines; Trigonostylops is a smaller Eocene version, and the Miocene
Astrapotherium is the type genus; South America and Antarctica, Paleocene to
Miocene.
Astraspis: a genus of jawless fish in the subclass Astraspida; 8 inches long, armored
with bony tiles, which were somewhat star-shaped on its head; related to Arandaspis;
North and South America, Middle Ordovician.
astrophic: of brachiopods, without a well-defined hinge line, having the two
valves hinged at a point.
Atacamatitan chilensis: name given in 2011 to a species of titanosaurian sauropod
dinosaurs, possibly related to the Nemegtosauridae; originally called Domeykosaurus,
it is known from several skeletal fragments first described in 2003 and dated near
the end of the Cretaceous; Northern Chile, Late Cretaceous.
28 A
Atelostomata: a superorder of irregular echinoids characterized by the absence of

a lantern; they are microphagous feeders, finding particles of food on the substrate.
Athyridida: a long-lived order of articulate brachiopods in the class Rhynchonellata;
characterized by biconvex, smooth, and impunctate shells with laterally directed
spires; global distribution, Late Ordovician to Jurassic.
Atlanthropus mauritanicus: see Ternifine Man.
Atlasaurus imelakei: a species of sauropod dinosaurs; known only from an almost
complete skeleton found in Morocco, it is estimated at about 50 feet long and 26
tons; Atlasaurus is related to the larger Brachiosaurus but has a shorter neck and
comparatively larger skull; its spoon-shaped teeth have denticles; Morocco, Middle
Jurassic.
Atrypa: a genus of articulate brachiopods in the order Atrypida and family
Atrypidae; characterized by a round, impunctate shell with many fine ridges; the
pedicle valve is convex except that it may transition to concave toward the anterior
edge; the brachial valve is convex; cosmopolitan, early Silurian to Late Devonian.
Attercopus: a monospecific genus of arachnids in the order Uraraneida; A. fimbri-
unguis, the oldest known proto-spider, produced silk but probably used it only to
line its burrows; see also Permarachne; North America, Middle Devonian.
Aturia: a genus of nautiloids in the order Nautilida, the only genus in the family
Aturiidae; it is discoidal, involute, and smooth-shelled, and has a very complex
suture pattern; cosmopolitan, early Paleocene to late Miocene.
Augustynolophus: a monospecific genus of hadrosaurid dinosaurs in the subfam-
ily Saurolophinae; known from a skull, vertebrae, and limb bones, A. morrisi was
the first dinosaur found in California, near Fresno; it was first identified as
Saurolophus morrisi; late Maastrichtian.
Aulacothyris: a genus of punctate articulate brachiopods in the superfamily
Zeillerioidea; characterized by a long loop and a convex pedicle valve with a strik-
ingly large fold along the posterior to anterior plane; Triassic to Jurassic.
Aulacocerida: the oldest coleoids, an order of cephalopods in the subclass
Belemnoidea; global distribution, Devonian to Late Jurassic.
Aulolepis: a genus of teleost neopterygian fish in the small extinct order
Ctenothrissiformes; characterized by a short, deep body and a large dorsal fin;
Europe, Late Cretaceous, late Cenomanian, about 95 Ma.
Aulophyllum: a genus of solitary rugose corals in the extinct superfamily
Zaphrenticae; closely related to Dibunophyllum; Great Britain, Early Carboniferous.
Aulopora: an order of colonial tabulate corals in the extinct family Auloporidae;
characterized by cylindrical corallites in a branching tubular structure, encrusting a
hard substrate; Ordovician to Permian.
A 29
Australopithecus: a genus of primates in the family Hominidae. With specimens

spanning the period from 4.4 to 1.5 Ma, the australopithecines are among several
possible ancestors of the genus Homo. They exhibit both human and ape character-
istics, with a cranial capacity of around 600 cc. Some authorities have classified
them in two types, “robust” and “gracile,” but the continuing variety exhibited by
newly found specimens indicates that there was a wide range of types of australo-
pithecines and other early hominins living in East Africa at the time. In addition to
the five species mentioned here, other proposed species named so far include, Au.
boisei, Au. garhi, Au. paranthropus, Au. prometheus, and Au. robustus.
Australopithecus afarensis: the best-known species of australopithecines, known
from remains of several individuals, including the famous specimen named Lucy.
Found in East Africa, particularly near the border of Ethiopia and Tanzania, the
remains have generally been dated as being between 3 and 4 million years old.
Research indicates that Au. afarensis could walk upright because the foot bone
exhibits an arch similar to that of modern humans. See also Middle Awash.
Australopithecus africanus: a well-known species of australopithecines, known
from several sites in Southern Africa. Although anatomically similar to Au. afaren-
sis, it had a larger brain and smaller teeth. Au. africanus is probably not a direct
ancestor of H. sapiens. Since most of the specimens were found in caves, its dating
is problematic, but it probably lived between 3.3 and 2.1 Ma.
Australopithecus anamensis: an early australopithecine, known from specimens
found in the Middle Awash and in Kenya; about 4.1 Ma.
Australopithecus bahrelghazali: a species of australopithecine hominins, known
from several skull and dental fragments dated between 3.5 and 3.0 Ma, about the
same age as Au. deyiremeda and Kenyanthropus platyops; Chad.
Australopithecus deyiremeda: a proposed new hominin species, based on a 2015
description of a jaw found in Ethiopia, dated between 3.5 and Ma.
Australopithecus sediba: a possible human relative from around 2 Ma, known
from two almost complete skeletons found in 2008 near the Malapa Cave, north of
Johannesburg, South Africa. A significant find for being possibly the only hominin
specimen known between the Au. afarensis known as Lucy (about 3.2 Ma) and the
Homo erectus known as Turkana Boy (about 1.5 Ma). The disputed dates proposed
for the more recent discovery H. naledi vary between 2.5 and 0.9 Ma.
Austroptyctodus gardineri: See Gogo Formation.
autapomorphy: a distinctive derived character possessed uniquely by a particular
lineage and not possessed by others in comparison; see also synapomorphy.
autochthonous: referring to an assemblage of fossils preserved undisturbed in
their life position and location; see also taphocoenosis.
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Avalofractus: an Ediacaran genus of rangeomorphs; 2 inches long, its alternating

fern-like fronds branched off from a central stalk; known only from the Avalon
Peninsula, Newfoundland, Canada.
Aves: a group of winged vertebrates, almost all capable of flight to some degree.
Although their origins and line of descent are still debated, the current consensus is
that birds evolved from feathered dinosaurs. The Enantiornithiformes are the earli-
est large group of primitive birds, but others are Confuciusornis (130–125 Ma),
Enaliornis (100–94 Ma), Ichthyornis (95–85 Ma), Hesperornis (84–78 Ma). See
also Protoavis.
Aysheaia: a genus of Cambrian soft-bodied animals superficially resembling a cat-
erpillar; one of the enigmatic animals of the Burgess Shale assemblage, possibly
related to the modern Onychophora; its body consisted of 10 segments, each with a
pair of appendages; North America and possibly China, early to middle Cambrian.
Azhdarchidae: a family of pterosaurs in the suborder Pterodactyloidea. Although
several partial specimens may date back to the Early Cretaceous, it is known well
only toward the end of the period. The family is known for very large species,
including the leading candidates for largest pterosaur, Quetzalcoatlus and
Hatzegopteryx, both of which had wingspans of about 40 feet. But a 2009 discovery
of a cat-sized azhdarchoid in British Columbia (between 80 and 70 Ma) shows that
some small pterosaurs were still competing with birds at the end of the Cretaceous.
Global distribution, Early to Late Cretaceous.
B
background extinction see extinction.

Bacteria: the major domain of prokaryotes, single-celled organisms without a
defined nucleus. They are abundant in every environment from the Mariana Trench
to the Himalayas, and there is even some evidence of bacteria on the surface of
Mars. On Earth these microbes constitute a greater biomass than all other animals
and plants combined. The earliest evidence of bacteria is cyanobacteria-like fossils
3.5 billion years old. See also Archaea and Eukaryota.
Bactrites: a genus of nautiloid cephalopods in the order Bactritida; generally
straight-shelled, it may have been ancestral to the ammonoids and coleoids, and
was also one of the few nautiloids that survived the end-Permian extinction; global
distribution, Early Devonian to Early Triassic.
Baculites: “walking stick,” an unusual genus of large orthocone ammonoids in the
order Ammonitida and superfamily Turrilitoidea. Widespread in the Late Cretaceous,
Baculites evidently lived in large shoals, although not so numerous as the earlier
orthocerid nautiloids that form key Ordovician beds worldwide. It is very similar in
appearance to the orthocerids but is differentiated by a relatively simple suture pat-
tern. From 6 inches to 6 feet long and sexually dimorphic, the male baculitids aver-
age less than half the size of the females. In cross-section it is oval and often found
so because the fossil shell fractures easily. The genus contains many index fossils
for the Western Interior of North America, such as B. clinolobatus of the Pierre
Shale (69 Ma). Cosmopolitan, Cretaceous.
Bahariasaurus: a genus of large but swift ceratosaur dinosaurs about the same size
as Tyrannosaurus rex and Carcharodontosaurus; discovered in 1924 by Ernst
Strömer but lost with his other specimens in 1944; see also Spinosaurus and
Deltadromeus; North Africa, Late Cretaceous, about 95 Ma.

32 B
Balaenidae: a still-extant family of whales in the suborder Mysticeti, including the

right and bowhead whales; reaching lengths over 50 feet and weighing up to
100 tons, the balaenids are widespread in temperate and polar waters; Miocene to
Recent.
Balaenoptera: a still-extant genus of mysticete whales in the family
Balaenopteridae; its eight living species include the blue, fin, and minke whales;
several fossil species are known, including B. cephalus, which was formerly thought
to be in the gray whale’s genus (Eschrichtius); Miocene to Recent.
Balcoracania: an unusual genus of redlichiid trilobites in the family Emuellidae;
it is less than an inch long, with a semicircular cephalon; its most striking feature is
a high number of thoracic segments (almost 100 in B. dailyi), some with pleural
spines as long as the two genal spines; Australia and Antarctica, early Cambrian.
Balizoma variolaris: the type species of the genus, phacopid trilobites in the fam-
ily Encrinuridae; with strikingly coarse tubercles on the cephalon, it is one of sev-
eral “strawberry-headed” genera; Europe and North America, restricted to Silurian.
Baltica: the name given to the land mass that is now Northern Europe when it was
a separate continent from the Cambrian to the Middle Ordovician.
Baltoniodus triangularis: a species of conodonts in the family Balognathidae; its
first appearance marks the beginning of the Middle Ordovician at 470 Ma; China.
Baluchitherium: see Paraceratherium.
Bambiraptor feinbergi: a species of Late Cretaceous dromaeosaurs, related to
velociraptors; 3 feet long and delicately boned, it was bipedal and is notable for its
long, wing-like arms; known only from a single complete skeleton found in 1994 at
Choteau, Montana, possibly a juvenile; United States 75 Ma.
Banffia: a genus of primitive animals placed tentatively in the phylum Vetulicolia;
although it is known from hundreds of specimens from the Burgess Shale, its clas-
sification is unsettled; about 4 inches long, its oval body and flat tail are twisted;
Canada, middle Cambrian, 508 Ma.
Bangiomorpha pubescens: a species of red algae in the order Bangiales; the earli-
est-known complex multicellular organism; also the earliest known example of
sexual reproduction, as evidenced by its differential spore and gamete formation;
Canada, Mesoproterozoic, 1200 Ma.
Bankia: a genus of shipworms, marine bivalve mollusks in the family Teredinidae;
global distribution, Early Jurassic to Recent.
Baragwanathia: a genus of primitive plants in the order Drepanophycales; the old-
est known lycopod club moss and one of the first vascular plants; known best from
Australia but global distribution, late Silurian to Early Devonian.
Barbourofelis: a genus of nimravid carnivorous mammals in the family
Barbourofelidae; muscular, leopard-sized predators, the Barbourofelidae were one
B 33
of the groups known as false sabertooth cats; they were generally smaller and more
primitive than the true felids; Miocene to Pliocene, Western North America.
barnacles: see Cirripedia.
Barychilina: a genus of ostracods in the order Metacopida; Silurian to
Carboniferous.
Barylambdidae: a small family of pantodont mammals in the extinct order
Cimolesta; they were herbivorous and tapir-like, the largest the size of a small horse;
Paleocene to early Eocene.
Baryonyx: “heavy claw,” a genus of theropod dinosaurs in the family Spinosauridae,
very similar to Suchomimus. About 25 feet long and weighing more than a ton, it
was characterized by a large claw on its first finger and a triangular nasal crest. The
baryonychines differed from most spinosaurids in having no dorsal fin or sail, and
their long, narrow gharial-like snout indicates that they were at least partly fish-
eaters. England and Iberian Peninsula, Early Cretaceous, 130–125 Ma.
Basilosaurus: a genus of archaeocete whales in the family Basilosauridae, closely
related to the slightly earlier protocetids. Fully aquatic, Basilosaurus is character-
ized by a completely flexible backbone for efficient propulsion and by vestigial
external hindlimbs too small to be used for swimming. It reached lengths of up to
40 to 65 feet and was the largest marine animal of the late Eocene. Africa and North
America, Eocene.
Batesian mimicry: the name given to camouflaging structural mimicry used for
protection from predation. The mimicking organism (usually uncommon but good-
tasting) closely resembles another that predators have learned to avoid because,
although abundant, it is foul-tasting, venomous, or stinging. First analyzed and
explained by Henry W. Bates in the 1860s, it is observed throughout the plant and
animal domain but is most common in invertebrates. A related kind of deception
called Müllerian mimicry involves two distasteful or poisonous species that have
evolved to resemble each other. A famous example of Müllerian mimicry is the case
of Danaus plexippus (monarch butterfly) and Limenitis archippus (viceroy), which
were long thought to be in a Batesian relationship until the viceroy was found to be
also unpalatable to predators; research continues, however, with conflicting reports.
Mimicry is difficult to identify in the fossil record because it usually involves sur-
face coloration, scent, or sound.
Bathornithidae: a family of birds in the order Cariamiformes; generally flightless,
the family also includes at least one genus capable of flight, Neocathartes; closely
related to the Phorusrhacidae and to the still-extant seriemas; North America, mid-
dle Eocene to early Miocene.
Batoidea: a superorder of chondrichthyan fish in the subclass Elasmobranchii,
including the skates, rays, and sawfish; generally benthic, except for the pelagic
manta rays; characterized by expanded pectoral fins which, especially in the skates
34 B
and rays, resemble wings; an example of convergent evolution compared to the

unrelated rhenanid placoderms (e.g., Gemündina), which also had expanded pectoral
fins; global distribution, Early Triassic to Recent.
Batrachia: the clade that includes the salamanders and the modern frogs, the
Anura.
Beecher’s Trilobite Bed: a Konservat-Lagerstätte near Rome, New York. It is in a
stratum of the deep-ocean Frankfort Shale, dated at about 445 Ma, that contains a
remarkable assemblage of about thirty Ordovician animals, many pyritized (replaced
by iron sulfide). It is notable for preserving many soft-tissue features of trilobites
(especially Cryptolithus, Triarthrus, and larval Cornuproetus) and other marine ani-
mals. Extensively explored by Charles Beecher in 1893, the site was thoroughly
excavated by a large team in 1989. North America, Late Ordovician.
Beelzebufo: a genus of large toads in the still-extant family Leptodactylidae,
related to the Ceratophryidae of South America. Many specimens have been found
on Madagascar in strata dated around 70 Ma, after Africa and India had begun to
separate from South America, isolating Beelzebufo from its ceratophryid cousins.
Up to 16 inches long and among the largest of all frogs, it probably bore bony scutes
on its head. Madagascar, Late Cretaceous.
Beipiaosaurus inexpectus: a genus of feathered theropod dinosaurs in the super-
family Therizinosauroidea, known from two individuals of about 120 Ma. From 6
to 7 feet long with 4-inch claws and stiff, narrow proto-feathers, it probably did not
fly. Although a theropod, it seems to have adapted to a herbivorous diet. It is one of
several feathered dinosaurs in the Jehol biota; China, Early Cretaceous, 120 Ma.
Beishanlong grandis: a species of ostrich-shaped dinosaurs in the family
Deinocheiridae; one of the largest ornithomimosaurs known, more than 25 feet long
and weighing 1.5 tons, it had digging claws and powerful forelimbs; southern Gobi
Desert, Early Cretaceous, 110 Ma.
Belantsea: a genus of chondrichthyan fish in the family Petalodontidae. It was
2–3 feet long, deep-bodied with large pectoral and ventral fins but a small tail fin.
Belantsea’s body plan suggests it was a slow swimmer. It is characterized by a bul-
bous shape and an unusual petalodont dentition which enabled it to eat crinoids and
encrusting organisms like sponges. Montana, Early Carboniferous, about 350 Ma.
Belemnitida: “dart-shape,” a large order of Mesozoic squid-like cephalopods in
the still-extant subclass Coleoidea and extinct superorder Belemnoidea. Belemnites
were from 4 to 8 inches long and had ten arms; like other cephalopods, they had an
ink sac, beak, tentacles, and large eyes. Unlike other cephalopods, they had a bullet-
shaped calcitic internal structure called a rostrum, which was often fossilized. The
belemnites appeared in the Triassic and were widespread in the Jurassic and Early
Cretaceous. All belemnoids became extinct at the end of the Cretaceous, but they
are closely related to modern cuttlefish.
B 35
Bellinurus: an extinct genus of freshwater arthropods in the order Xiphosurida;

though less than 2 inches long, it is related to modern horseshoe crabs (see Limulus);
Europe, Devonian to Carboniferous.
bellerophontids: a large group of sea snails, gastropods in the order
Archaeogastropoda (or possibly monoplacophorans); they were generally coiled
symmetrically; up to 2 inches long, the type genus is Bellerophon; Ordovician to
Triassic.
Bennettitales: a group of small cycadeoid or anthophyte trees, growing to about
7 feet high; their leaves resemble cycads, but they are probably not related; wide-
spread Triassic to Cretaceous and surviving sparsely to the Oligocene.
benthic: living at the lowest level of a body of water, from eulittoral to abyssal, as
opposed to living in the water column; generally referring to oceans but also to lakes
and rivers; see also nektic and nektobenthic.
Bernissartia: a genus of small, semi-aquatic crocodylomorphs in the family
Bernissartiidae; similar to modern crocodiles, it is considered a neosuchian; it was
less than 3 feet long, and its dentition was adapted to crushing shellfish and crusta-
ceans but also to catching fish; named for specimens found at the Bernissart mine in
Belgium; North America and Europe, Early Cretaceous, 135–125 Ma.
Bernissart site: a Belgian coal mine where many significant fossils of the Early
Cretaceous have been found, especially important for the specimens of iguanodonts
found there in the 1870s.
Berriasella jacobi: a species of ammonoids whose first appearance marks the
beginning of the Early Cretaceous at 145.0 Ma; Europe.
Bertie Waterline: name given to several upper Silurian deposits in New York that
contain well-preserved arthropod and plant fossils, known especially for Eurypterus
remipes; United States, late Silurian.
Bethia serraticulma: a species of brachiopods placed variously in the orders
Orthida and Strophomenida, known only from one specimen found in a carbonate
nodule (see Herefordshire Nodules); the digital reconstruction of Bethia shows
excellent soft-part morphology; England, Silurian.
Beyrichicopina: a suborder of dimorphic ostracods in the extinct order
Palaeocopida; North America and Europe, Ordovician to Early Mississippian.
Biarmosuchia: a suborder of basal therapsids, including Biarmosuchus and
Eotitanosuchus; generally similar to sphenacodonts, with long canine teeth; Russia,
early Permian.
Bighorn Basin: an arid area in northern Wyoming that is an important study site
for the transition from Paleocene to Eocene; about 10,000 square kilometers in area,
notable for the plant life and the large numbers of mammal species recorded in its
varied strata.
36 B
bilateralism: having a definite and symmetrical left and right side, as opposed to
amorphous animals like sponges and jellyfish.
bilaterians: the bilaterally symmetrical animals, excluding sponges, cnidarians,
placozoans, and ctenophores; characterized by bilateral symmetry and an anterior-
posterior body orientation; the oldest known specimen may be microfossil embryos
from an Ediacaran deposit in China (see Vernanimalcula guizhouena), thought to
be 580 million years old; these are also the earliest-known embryos of complex
animals; Ediacaran to Recent.
Billingsellida: an order of articulate brachiopods in the class Strophomenata; one
of the first and most primitive articulate brachiopods, it is characterized by a wide
hinge-line; middle Cambrian to Middle Ordovician.
biocoenosis: an assemblage of organisms living together at one time, an ecological
community.
biochron: a period of time represented by a relatively small biostratigraphic zone,
one characterized by a particular fauna or flora.
biosphere: 1. the area favorable for the occupation of living organisms, including
the lithosphere, hydrosphere, and atmosphere. 2. All the living organisms of the
Earth and its atmosphere.
biostratigraphy: the science of correlating geographically separated rock strata
by identifying the biotic content and time relations of the strata; it is based on faunal
succession, the process in which species change over time.
biostratinomy: a subsection of taphonomy, dealing with the orientation of the
remains of an organism and what it says of the processes the organism experienced
between its death and its burial, such as transportation and chemical change.
biota: the total fauna and flora living or preserved as fossils within a certain area
or period of time.
biozone: a stratigraphic unit of specific geological strata defined by various com-
binations of taxa present in the strata.
bird-hipped dinosaur: general term referring to the ornithischian dinosaurs.
birds: see Aves.
Birgeria americana: a species of large predatory fish in the family Birgeriidae,
known from a partial skull described in 2017; a bony fish about 7 feet long, it lived
in water warmer than previously thought possible for the development of eggs;
Nevada, United States, Early Triassic, 251 Ma.
Bison priscus: a species of large-horned bison of North America; its extinction in
the late Pleistocene was probably related to human activity.
Biston betularia: the peppered moth, a still-extant species of lepidopterans in the
family Geometridae. Two forms (B. b. f. typica and B. b. f. carbonaria) have evolved
B 37
in the Holocene and have been studied in detail since the 1850s. Their variation in
color over two centuries, caused by industrial melanism, results in differential
predation by birds and is now generally accepted as a clear example of Darwinian
natural selection in action. Global distribution, Recent.
bitheca: a smaller theca that appears in many graptolites, alternately on both sides
of the autothecae.
Bithecocamara: an enigmatic genus of pterobranch encrusting graptolites in the
order Camaroidea; the only camaroid with bithecae; Baltic, Ordovician.
bivalves: mollusks that typically have shells with two hinged valves, not related to
the two-valved brachiopods, which are different organisms that have developed a
similar shell because of a similar lifestyle, an example of convergent evolution. The
two valves of a bivalve are usually symmetrical, but there are several exceptions.
The rudists, for example, are horn- or tube-shaped bivalves whose external mor-
phology seems to bear no similarity to other bivalves. And shipworms (see Bankia
and Teredo) have only a pair of very small shells, at the anterior end of a tube-like
body, with which they bore through wood. Global distribution, early Cambrian to
Recent.
Bjuvia: a genus of short trees with a crown of broad lanceolate leaves atop a squat,
branchless trunk; Europe, Early Triassic.
Blackberry Hill: a Lagerstätte in Wisconsin consisting of several quarries; it is
best-known for its extensive trace fossils, mostly trackways left in a shallow-water
sandstone formation; notable also for body fossils of a phyllocarid and a euthycar-
cinoid (for the latter, see Mosineia), considered by some to be the earliest body
fossils of a land animal; United States, middle or late Cambrian.
Blastoidea: a class of sessile pelmatozoan echinoderms in the subphylum
Blastozoa; they are sometimes called sea buds because of their nut shape; com-
monly stacked, they were attached to the sea floor on a stem; global distribution,
Ordovician to late Permian.
Blastozoa: an extinct subphylum of echinoderms; characterized by hydrospires,
small folds beneath the lancet plates; Cambrian to Permian.
Blattodea: the order of insects that includes cockroaches and termites; the ances-
tors of cockroaches are known from the Carboniferous, and modern roaches
appeared in the Triassic; termites probably evolved in the Permian, but specimens
are known only as far back as the Early Cretaceous; also called Blattoptera.
Blikanasaurus: a proposed genus of sauropodomorph dinosaurs, known only from
one limb bone; South Africa, Late Triassic, 210–202 Ma.
Bobasatraniidae: a small family of deep-bodied chondrostean fish, in the crown
group of the Actinopterygii; characterized by a diamond-shaped body, a forked tail
fin, and long, thin pectoral fins; North America, Europe, and Africa; late Permian to
Early Triassic.
38 B
body plan: see Cambrian and Ediacaran biota.

bone bed: a fossil site that contains a large number of animal macrofossils. Of
special importance are bone beds where many animals died abruptly, leaving whole-
body fossils. Those where whole-body fossils of many different species are found
together raise questions about the cause of the deaths, from flash floods to accumu-
lations of poisonous substances. The Messel Shale Pit, for instance, contains many
different types of animals, including birds and bats, and also turtles that died in the
act of copulating; poisonous gases have been the most common cause suggested.
Toxic algae have been proposed as a possible cause of the deaths of several layers
of fossils in the Maevarano site in Madagascar. No conclusive evidence has been
found in either case.
bony fish: see Osteichthyes.
Bootherium nivicolens: a species of musk ox in North America whose extinction
in the late Pleistocene may have been caused by humans.
Borealopelta: a monospecific genus of nodosaurid dinosaurs, known from a
whole-body specimen described in 2017. Recovered from an oil sands mine in
Alberta, B. markmitchelli is the best-preserved specimen known of an ankylosaur,
with soft tissue as well as its armor (many sharp-pointed osteoderms and two long
shoulder spines) in life-position. Like other nodosaurids, it has no club at the end of
the tail. The keratin in skin and scales indicates countershading, with a dark reddish-
brown coloration above and a lighter underside. Canada, Early Cretaceous, 110 Ma.
Borealosuchus: a genus of crocodilians widespread in North America from the
Late Cretaceous up to the middle Eocene; about 9 feet long, it is one of several
genera of crocodilians that survived the end-Cretaceous extinction.
Boskovice Furrow: a large site in Moravia that contains a series of deposits in
upper Carboniferous and lower Permian strata; important for tetrapods from the
Permian strata, especially in the family Discosauriscidae, q.v.; Czech Republic.
Bostrychoceras: a genus of heteromorph ammonoids in the order Ammonitida and
family Nostoceratidae; loosely coiled but slightly tighter than its cousin Nostoceras,
its shell is covered with fine, dense ribs and forms a long, slightly open helical spire;
Europe, Africa, and North America; known only from the Campanian Stage near the
end of the Cretaceous.
Bothriocidaris: an unresolved genus of echinoderms, generally considered an
echinoid; it was globular and was covered with spines and tube feet; known only
from Upper Ordovician deposits in Estonia, Scotland, and North America.
Bothriolepis: a genus of heavily-armored placoderm fish in the order Antiarchi,
bottom-dwelling and mainly freshwater rather than marine. Generally 1 foot long,
with some almost 3 feet, Bothriolepis had a slender and somewhat flattened body,
and was notable for a pair of long, spine-like pectoral fins with small spines on
them. Its eyes were located on the top of the headshield, along with a light-receptive
B 39
pineal eye. It had heavy anterior armor but no posterior armor, not even scales.
Widespread, abundant, and diverse in the Middle to Late Devonian, it became
extinct with all other placoderms at the end of the Devonian.
Botryococcus: a still-extant genus of one-celled freshwater algae that can also
grow in brackish water environments; notable for a high hydrocarbon content in and
outside its thick, waxy cell walls, held together by a lipid biofilm; useful in inter-
preting paleoenvironments; Triassic to Recent.
Bovidae: a family of horned, cloven-hoofed, ruminant mammals in the order
Artiodactyla, including cattle, sheep, bison, goats, buffaloes, and antelopes; distin-
guished from the Cervidae by their hollow, non-deciduous, unbranched horns, all
male bovids and two-thirds of the females have two or more horns; the earliest-
known genus is Eotragus, which appeared in Eurasia around 20 Ma; global distribu-
tion, early Miocene to Recent.
bowfin: see Amia calva.
Brachiopoda: “arm foot,” a phylum of solitary marine invertebrates characterized
by a dorsal (brachial) and a ventral (pedicle) valve; a pedicle, a muscular extension
of the body wall used by most genera to attach to the substrate; and an internal
lophophore, which is bilobate and typically horseshoe-shaped but often in long
loops. In some articulate brachiopods, the two lobes of the lophophore are sup-
ported by calcareous support structures called brachidia, which are attached to the
brachial valve (see Spiriferida).
Called lampshells, brachiopods are superficially similar to bivalves but are not
mollusks; the two halves of their shells are not mirror-images of each other, as in
bivalves, but symmetrical, just in a different plane. The internal body parts are rela-
tively sparse compared to the those of the bivalves. Most fossil brachiopods are
1–3 inches wide, and the largest known is about 8 inches wide. See also punctate.
Several genera were epi-planktic and a few were nektic, swimming in the style
of scallops. They were widespread in the Paleozoic, but only one-third of the genera
survived the end-Permian extinction. Although the bivalve mollusks have become
dominant, brachiopods have continued to increase slightly in number in the
Cenozoic, with about 250 species of brachiopods still extant, the most famous being
Lingula, which has remained essentially unchanged in shape since the Cambrian.
Extinct brachiopods are significant index fossils in the Paleozoic because of their
broad distribution and their abundance and diversity at several taxonomic levels,
especially for the Devonian and Carboniferous. Cambrian to Recent.
Brachiosaurus: a genus of titanosaurian sauropodomorph dinosaurs in the family
Brachiosauridae; estimated at 85 feet long and 32 tons, it is one of the largest sau-
ropods; its forelimbs are relatively elongated, and its neck is very long; United
States, Europe, and Africa; Jurassic.
Brachyaspidion: a genus of small trilobites in the order Ptychopariida; B. microps
(1 cm) and B. sulcatum (4 cm) are among the dozens of trilobites in the Wheeler
Shale Lagerstätte; North America, middle Cambrian.
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Brachyceratops: an obscure genus of ceratopsian dinosaurs, known only from

p artial juvenile specimens found in Campanian strata in Montana and Alberta;
North America, Late Cretaceous.
brachydont dentition: low-crowned teeth with long, narrow roots; they do not
grow throughout life in all animals, but do in some, such as rodents.
Brachylophosaurus: a genus of hadrosaurid ornithischian dinosaurs in the sub-
family Saurolophinae; the type species is B. canadensis, but the genus is known
from many skeletons and casts showing soft-tissue detail, including tendons and
hundreds of thin tracks that have been interpreted as evidence of stomach parasites;
Montana and Alberta, Late Cretaceous, 80–75 Ma.
Brachysaurus: synonym: Prognathodon.
Brachythoraci: the largest suborder of armored arthrodires; the early Brachythoraci
were relatively small, like the agnathans, but the best-known was the gigantic
Dunkleosteus, which grew to 30 feet long; like all other placoderms, they became
extinct at the end of the Devonian.
Bradgatia: a monospecific genus of Ediacaran fossils, possibly a rangeomorph;
dated at about 570 Ma, B. linfordensis is a small, bush-like petalage, a cluster of 6–8
petals a few inches high; found especially in the Ediacaran assemblages at Mistaken
Point and Charnwood Forest; Canada and England, Ediacaran.
Bradoriida: an order of basal crustaceans widespread in the early Paleozoic; they
were small, with a bivalved carapace over the head and body; global distribution,
Cambrian to Ordovician.
bradyodont: having paired hard, enameled toothplates in the upper and lower jaw,
used to crush prey; characterizes several groups of fish that flourished in the
Carboniferous and are sometimes referred to as Bradyodonti; Late Devonian to
Permian.
Branchiopoda: “gill foot,” a still-extant class of small, generally freshwater, crus-
taceans. The class includes several living groups like the fairy shrimp (Anostraca),
which are as long as 7 inches. Branchiopods have a long (see Lepidocaris rhynien-
sis) but incomplete fossil record, and they may be more closely related to insects
than to other crustaceans. They are thought to have originated in a marine environ-
ment but to have adapted to freshwater environments because of competition. Late
Cambrian to Recent.
branchial: referring to gills.
branchiostegal: having covered gills.
Branchiostomiformes: see Cephalochordata.
Brontosaurus: a contentious genus of sauropod dinosaurs; long considered a
junior synonym of Apatosaurus, Brontosaurus is now considered by some to be a
separate genus; for description, see Apatosaurus.
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Brontotheriidae: “thunder animal,” a large family of herbivorous perissodactyl

mammals, a sister group to the Equidae in the suborder Hippomorpha. Also called
titanotheres, they were almost elephant-sized, ranging in height from 2 to 9 feet.
They are marked especially by a variety of bony frontal horns, even with a pair of
blunt nasal horns in Megacerops. Although similar in appearance to rhinoceroses,
they are more closely related to horses. The earliest brontothere, the dog-sized
Eotitanops, appeared in the early Eocene. North America, Europe, and Asia; early
to late Eocene.
bryophytes: “moss plant,” a common name for the group of primitive land plants
that includes the mosses (clade Bryophyta), hornworts (clade Anthocerotophyta),
and liverworts (clade Marchantiophyta). Not vascular and lacking true roots and
leaves, the bryophytes reproduce by spores. Molecular phylogenetic research indi-
cates that the bryophytes were the earliest land plants, probably appearing in the
Ediacaran. The earliest definitive bryophytic fossils (such as Sporogonites) are
dated Early Devonian. See also Protosalvinia.
Bryozoa: “moss animals,” a still-extant phylum of small colonial animals that
secrete calcified skeletons around the individual organisms’ bodies, a process seen
also in reef-building corals. Bryozoa is the only phylum of animals that are wholly
colonial. The bryozoans are the only phylum with an extensive fossil record to have
arisen later than the Cambrian — the earliest-known bryozoan fossils date from the
Early Ordovician. Bryozoans are one of the dominant groups of Paleozoic fossils,
with tens of thousands of named species. Only a few families survived the end-
Permian extinction, but there are still about 6,000 living species.
Bubalus cebuensis: an extinct species of dwarf water buffalo, less than 3 feet high
and weighing about 350 pounds; descended from much larger buffalo, they experi-
enced island dwarfism when stranded on the small Philippine island of Cebu by a
rising sea level between 100,000 and 20,000 years ago; see also B. mindoroensis.
Bubalus mindoroensis: the tamaraw, a still-extant species of dwarf water buffalo
found on the Philippine island of Mindoro; an example of island dwarfism, being
descended from larger buffalo stranded on the island by rising sea levels about
20,000 years ago.
Buchia: a genus of edentulous bivalves in the superfamily Buchioidea; they have a
slender shape, with the left valve larger than the right valve; known around the
Pacific Rim, Jurassic to Cretaceous.
Bumastus: a genus of large (up to 6 inches) bottom-dwelling trilobites in the order
Corynexochida and family Styginidae; notable for a smooth, rounded, isopygous
body and well-developed compound eyes; global distribution, Early Ordovician to
late Silurian.
bunodont dentition: low-crowned, conical or rounded teeth with ridged cusps;
separate, often quadrate in shape; typical especially of early artiodactyl and suiform
mammals.
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Burgess Shale: name given to one of the most important Konservat-Lagerstätten,

a calcareous black shale formation dated at 508 Ma. Located near Kicking Horse
Pass in the Rocky Mountains of eastern British Columbia, the formation is unique
in containing a surprisingly well-preserved assemblage of soft-bodied animals from
the middle Cambrian. The fossils are allochthonous, probably having been removed
from their living location by a series of mudslides. The assemblage evidently repre-
sents a complete marine community that includes both nektic and benthic animals.
The great diversity and excellent preservation of these very unusual fossils — dis-
covered in 1909 by Charles Walcott — provides a rare view of this period of rapid
diversification of animal species. Several specimens are the only known representa-
tives of phyla that are unlike any modern phyla. See also Chengjiang biota. Canada,
middle Cambrian.
byssus: a bundle of tough, hairlike filaments commonly used by bivalve mollusks
to attach themselves to objects, usually rocks or other hard substrate but sometimes,
as with Modiolus, to detritus in sediment.
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Caelifera: the grasshoppers and locusts, a suborder of chewing herbivorous insects

in the order Orthoptera; mostly ground-dwelling and stridulatory, with strong hind-
legs and wings; global distribution, Early Triassic to Recent.
Caenolestidae: a family of small South American marsupials in the order
Paucituberculata; commonly called shrew opossums and rat opossums, they were
once abundant but are now confined to small regions in the Andes Mountains; late
Oligocene to Recent.
Cainotheriidae: a family of small, herbivorous artiodactyl mammals in the subor-
der Tylopoda and extinct superfamily Anoplotheroidea; characterized by a rabbit-
like, strongly curved back, they resemble modern hares and rabbits but are unrelated;
they diversified rapidly in the Eocene, with only a few persisting into the Oligocene,
but then diversified again before disappearing by the end of the period; Europe,
Eocene to Oligocene.
Calamitales: an order of arborescent sphenopsid plants; resembling giant horse-
tails, they were reed-like plants with horizontal rhizomes and segmented and
grooved stems; up to 100 feet high and 12 inches in diameter; global distribution,
Late Devonian to Permian.
Calcarea: a class of small marine sponges; the only sponges with spicules of cal-
cium carbonate, they are found mainly in shallow tropical waters but have been
found as deep as 4,000 feet; global distribution, early Cambrian to Recent.
Calcaronea: a still-extant subclass of sponges in the class Calcarea, with three- or
four-rayed spicules; they may have appeared as early as the Cambrian, were espe-
cially prominent in the Cretaceous, and still exist worldwide.
Calceola: a genus of solitary rugose corals in the family Goniophyllidae; known as
slipper coral because of its shape, Calceola was semicircular and rested on the sea
floor, convex side down; Northern Europe, Early to Middle Devonian.

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Calcinea: a still-extant, probably paraphyletic subclass of sponges in the class

Calcarea; a generally spreading habit, usually yellow or red, with regular three- or
four-rayed spicules; global distribution, Permian to Recent.
Calclamna: a genus of sea cucumbers, echinozoans in the class Holothuroidea and
extinct family Calclamnidae; Europe and Asia, Early Triassic to Cretaceous.
Callavia: a monospecific genus of early Cambrian olenellid trilobites; C. broegeri
is known from several sites in northeastern North America.
Callistophytales: an order of vinelike pteridosperms widespread in the late
Paleozoic; North America, Europe, and Asia; Carboniferous to Permian.
Callograptus salteri: a species of irregularly branching graptolites in the order
Dendroidea; characterized by a fine net of branches with many small thecae; Europe,
Asia, and North America; late Cambrian to Early Carboniferous.
Calsoyasuchus: see Goniopholis.
Calymene: a genus of phacopid trilobites; some well-preserved specimens of
Calymene and its relative Flexicalymene show appendages and soft body parts;
specimens are frequently found enrolled, rolled into a ball in the manner of the
modern terrestrial isopods; global distribution, Ordovician to Devonian.
calyx: primarily, the central cup-shaped body of crinoids, resting on a stem, with
arms extending outward and upward; also, the cup formed by the sepals of a flower.
Camarasaurus: “chambered lizard,” a genus of very large herbivorous dinosaurs,
the most common sauropod of North America in the Late Jurassic. The largest spe-
cies was about 75 feet long and weighed about 52 tons, with a relatively short neck.
Camarasaurus is named for air sacs in the vertebrae that were connected to the
lungs. Many specimens have been found in the Morrison Formation, some well-
preserved. North America, Europe, and Africa; Late Jurassic to Early Cretaceous.
Camaroidea: an order of sessile encrusting graptolites known mainly from the
Ordovician of the Baltic Sea area; characterized by thecae with a balloon-shaped
base and vertical chimney; the best-known genus is Bithecocamara; Baltic and
Siberia, late Cambrian to Early Ordovician.
Cambaytherium: a genus of early placental mammals in the family
Cambaytheriidae, formerly placed in the Anthracobunidae; now considered a basal
perissodactyl. It is known from several fragmentary specimens found in strata dated
54.5 Ma, when the Indian island continent was nearing Asia after splitting off from
Gondwana. The size of a small pig, it has five digits and seems close to the ancestry
of horses, rhinoceroses, and tapirs. Its discovery has led to the conclusion that peris-
sodactyls evolved in Asia and spread to North America during the Eocene. Gujarat,
India, early Eocene.
Cambrian: the first period of the Paleozoic Era, 541 to 485 Ma. Its macrofossil
record is primarily marine, with just a few traces of early terrestrial invertebrates.
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The marine environment was dominated at first by reef-building Archaeocyatha and

other small animals, and then increasingly by primitive deuterostomes. Some of the
basic animal body plans had appeared in the Ediacaran, but most are known first
from the early Cambrian, probably between 535 and 530 Ma. These early species
diverged morphologically in a short period of time in what is known as the Cambrian
Explosion. The first primitive marine vertebrates (see Myllokunmingia,
Haikouichthys, and Zhongjianichthys) appeared near the end of this brief time.
Camelidae: an extant family of even-toed ungulates in the suborder Tylopoda. The
first camelids (see Protylopus) appeared in North America about 45 Ma and evolved
into several distinct genera, especially in the late Eocene and early Miocene. They
were known only in North America until the early Pliocene, 4 to 5 Ma, when some
families crossed the new land bridge to South America, developing there into the
modern guanacos, llamas, vicuñas, and alpacas. Then in the early Pleistocene, North
American camelids crossed the newly dry Bering Strait into Asia, where they devel-
oped into the modern camels. Camelids disappeared from North America soon
afterward.
Camelops: a late genus of camels, one of a dozen North American camelids that
became extinct at the end of the Pleistocene.
Camerata: a large extinct subclass of stalked crinoids. Its two main orders
appeared in the Middle Ordovician; the Diplobathrida disappeared in the
Carboniferous, while the Monobathrida continued up to the end-Permian extinc-
tion; the small order Cladida is restricted to the Devonian. All are characterized by
a long, flexible stem, rigidly sutured calyx, and more than ten free arms. Middle
Ordovician to late Permian.
Cameroceras: see Endoceratidae.
Camptosaurus: a genus of mid-sized herbivorous dinosaurs in the order
Ornithopoda, formerly called Camptonotus. About 26 feet long, it had a very small
head and small, flat teeth, and is estimated to have weighed up to 1 ton. Clawless
and basically bipedal, it had four toes on each pes and was a good runner. North
America and Europe, Early Jurassic.
Caninae: a subfamily of carnivorous mammals in the order Carnivora and family
Canidae; it includes the dogs, wolves, foxes, coyotes, dingoes, and jackals, as well
as their most recent ancestors, such as Leptocyon; Oligocene to Recent.
canine: conical mammal teeth between the incisors and premolars, adapted for
grasping and piercing prey; generally two pairs, upper and lower, but vestigial in
many herbivores.
Caninia: a genus of large, solitary rugose corals in the family Cyathopsidae; living
mainly in shallow lagoons, it was often affected by storms; global distribution,
Devonian to Permian.
Canobius: a genus of small palaeoniscid actinopterygian fish. Three inches long, it
is notable for its vertical jaw suspension, as opposed to the diagonal arrangement of
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other palaeoniscids. Its larger gape allowed it to swallow great volumes of plankton,
catching them on its very small teeth, which grew even on its gills. Scotland, Early
Carboniferous.
Cantius: a genus of early strepsirrhine adapiform primates in the family
Notharctinae; lemur-like, it is known from numerous teeth and jaw fossils; North
America and Europe, early Eocene.
Capinatator: “seizing swimmer,” a genus of large arrow worms in the phylum
Chaetognatha; C. praetermissus is known from specimens found at the Burgess
Shale site and described in 2017; it was 4 inches long and is characterized by two
dozen mobile grasping spines on each side of its mouth, with which it captured
prey; Canada, middle Cambrian, 508 Ma.
Capitosauridae: a family of temnospondyl tetrapods, generally considered primi-
tive amphibians; cosmopolitan, Early to Late Triassic.
Captorhinida: a paraphyletic order of stem reptiles of the late Paleozoic; formerly
called cotylosaurs; see Pareiasauridae and Procolophonia.
Captorhinus: a genus of small, lizard-like reptiles in the family Captorhinidae; it
was specialized, characterized by a hooked snout with procumbent front teeth; its
skull was 5 inches long and its limbs were slender; global distribution, early
Permian.
carapace: the hard, protective dorsal body covering of many arthropods and some
reptiles, such as turtles; it is composed of chitin or bony substances.
Carbonicola: a genus of vagrant freshwater clams in the family Anthracosiidae;
formerly called Anthracosia; several short-lived species are index fossils in the
British Coal Measures; Europe, Carboniferous.
Carboniferous: the fifth period of the Paleozoic Era, from 359 to 299 Ma. For
North American strata, the Carboniferous is divided into two sub-periods: the
Mississippian, from 359 to 323 Ma, and the Pennsylvanian, from 323 to 299 Ma;
both the Mississippian and the Pennsylvanian are divided into Early, Middle, and
Late epochs. The period saw the culmination of a long period of increasing levels of
oxygen in the atmosphere, reaching as high as 35 percent. The Carboniferous is
characterized globally by coal-bearing beds resulting from large forests. It was also
marked by several periods of widespread glaciation, especially in the last half of the
period and extending into the Permian. The seed ferns continued to be an important
part of the flora, the first gymnosperms appeared and spread rapidly, and the coni-
fers and ancestors of the cycads appeared near the end of the period. An important
development for vertebrates was the evolution of the amniotes. Late in the
Carboniferous, these later tetrapods diversified rapidly, one branch producing the
first reptiles. Terrestrial arthropods and flora also increased greatly in diversity.
Carcharodon: a still-extant genus of large white sharks in the family Lamnidae;
synonym: Carcharocles; the largest shark known to have lived was C. megalodon,
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which may have grown to a length of more than 60 feet and became extinct only
around 50,000 years ago; cosmopolitan, Miocene to Recent.
Carcharodontosaurus saharicus: a species of theropod dinosaurs in the family
Carcharodontosauridae. Named for the resemblance of its serrated teeth to those of
Carcharodon, it reached a length of almost 40 feet and was a top predator. It was as
large as Tyrannosaurus rex but slightly smaller than Spinosaurus. Tunisia and
Egypt, Late Cretaceous, 90 Ma.
Cardiocephalus: a genus of microsaurian lepospondyl amphibians in the family
Gymnarthridae; it was lizard-like, with four short legs and a long tail, and is named
for its heart-shaped head; North America, early to middle Permian.
Cardioceras: a genus of nektic ammonoids in the order Ammonitida and super-
family Stephanocerataceae; up to 10 inches across, it is notable for its strong ribs,
evolute circular shell, and prominent crest on the margin; the first appearance of C.
redcliffense defines the beginning of the Late Jurassic at 163.5; global distribution,
Jurassic.
carnassial teeth: upper and lower cheek teeth in most carnivorous mammals,
adapted for shearing flesh, as opposed to crushing or tearing.
Carnivora: a diverse order of placental mammals in the superorder Laurasiatheria.
The primary suborders are the Caniformes, Feliformes, and Pinnipedia. The earliest
were the Miacidae, which appeared in North America in the late Paleocene, and
their descendants radiated rapidly throughout the world. Classified on the basis of
dentition, some members of the order are omnivorous while others are herbivorous.
Paleocene to Recent.
carnosaurs: traditionally, a polyphyletic group of many large carnivorous sauris-
chian dinosaurs but now restricted to the allosaurs; a large head, a short neck, and
generally short forelimbs are characteristic structures of the group; global distribu-
tion, Jurassic to Cretaceous.
Carnotaurus: a genus of large carnivorous dinosaurs in the family Abelisauridae,
characterized by a large head, two short but stout horns above the eyes, relatively
small teeth, and very small vestigial forelimbs; C. sastrei is the type species, known
from a well-preserved skeleton found by Paul Sereno in Patagonia; Argentina, Late
Carnufex carolinensis: an ancestral crocodile, among the earliest crocodylo-
morphs, known from a North Carolina specimen described in 2015, with a skull,
spine, and forelimbs. It was about 9 feet long; judging from size and weight distri-
bution, it may have stood on its hind legs. Its diet consisted of early mammals and
small armored reptiles. North America, Late Triassic, 231 Ma.
carpoids: see Homalozoa.
cast: casts are formed when original parts of an organism are dissolved and the
resulting space is filled with sediment or other mineral matter.
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Castor canadensis: the North American beaver, a genus of rodents in the family
Castoridae; 3 feet long and almost extinct after centuries of trapping for its pelt, it
has recovered; it has many subspecies but is not related to Aplodontia, the “moun-
tain beaver”; North America, Miocene to Recent.
Castorocauda lutrasimilis: a fish-eating species of Jurassic semi-aquatic mamma-
liaforms in the order Docodonta. It was similar to beavers and otters but not related
to them. Recorded as the earliest semi-aquatic mammal, dated between 170 and
165 Ma, it is the largest known mammal of the Jurassic at 1–2 pounds and almost
2 feet long. It had webbed feet; a flat, beaver-like scaly tail; otter-like vertebrae; and
teeth like those of a seal. Northern China, Middle Jurassic.
Catarrhini: an infraorder of anthropoid primates that includes men, apes, and
Old-World monkeys (see also Platyrrhini). The Catarrhini are characterized by
nostrils that face downwards; most have tails, but they are non-prehensile. They
diverged from the Platyrrhini in the late Eocene or early Oligocene yet remained
primarily alike because of parallel evolution. The Catarrhini are subdivided into
Cercopithecidae (Old-World monkeys), Hominidae (humans, gorillas, chimpan-
zees, orangutans, and australopithecines), and Hylobatidae (gibbons). Eocene to
Recent.
catastrophism: see uniformitarianism.
Catenipora: a large genus of tabulate corals in the family Halysitidae; Baltic, Late
Ordovician to early Silurian.
Cathaymyrus: a genus of early Cambrian cephalochordates; similar to Pikaia, it is
long and slender (1–2 inches) and seems to have a notochord in the depression along
the center of its back; Chengjiang biota, China, 530–515 Ma.
Catopithecus: possibly the earliest-known primate; a squirrel-sized monkey
known from several finds in the Fayum Depression, including tooth and skull frag-
ments found in 1995; characterized by shovel-shaped upper front teeth, a flattened
face, forward-looking eye sockets, and a fused forehead bone; the latter is a key
difference between lemurs and the higher primates; all known specimens are dated
at about 36 Ma; Egypt, late Eocene.
Caudipteryx: “tail feather,” a genus of theropod dinosaurs with considerable plum-
age, including a large tail fan. Although it was bird-like and the size of a peacock,
its body was covered with only short feathers and it probably lacked the aerody-
namic qualities required for flight. Dated at 125 Ma, it is one of several feathered
dinosaurs in the Jehol biota. China, Early Cretaceous.
Cavellinidae: a family of Paleozoic ostracods in the order Platycopida and subor-
der Cytherelliformes; characterized by sexual dimorphism and a ridged shell sur-
face; Silurian to Permian.
Caytoniales: an order of Mesozoic seed ferns, important because of their possible
transitional relationship to the angiosperms; their four-part palmate leaves (form
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genus Sagenopteris) are similar to some later flowering plants, and they bear a
carpel-like case containing seeds (form genus Caytonia); a direct relationship is still
unclear, however, and they may be gymnosperms; global distribution, Late Triassic
to Early Cretaceous.
Cebochoerus: a genus of basal artiodactyl mammals in the extinct superfamily
Dichobunoidea and family Cebochoeridae; pig-like but unspecialized, the cebocho-
erids may have an ancestry in the little-known Helohyidae; Europe, middle Eocene
to early Oligocene.
cell division: see mitosis.
Cenophyticum: term sometimes used for a paleobotanic division of time roughly
corresponding to the Cenozoic Era and characterized by the plant life of the
Cenozoic; the abundant angiosperm floras of the Late Cretaceous are also generally
considered cenophytic.
Cenozoic Era: the third division of Phanerozoic geologic time, following the
Paleozoic and Mesozoic and extending to the present.
Centronellidina: a suborder of articulate brachiopods in the order Terebratulida;
characterized by a primitive loop structure and an incurving ventral beak; North and
South America, Early Devonian to late Permian.
Centrosaurinae: a subfamily of horned ornithischian dinosaurs in the family
Ceratopsidae; medium-sized, averaging less than 20 feet long, these herbivores
ranged across Western North America from Alaska to Arizona, with at least one spe-
cies, Sinoceratops, found in China in 2010; important especially for several embryos
found in nest sites; North America, 84–66 Ma.
Centrosaurus: a genus of ceratopsian dinosaurs in the subfamily Centrosaurinae;
the type species C. apertus was found in southeastern Alberta, where great numbers
of the genus have since been found; a medium-sized ceratopsian, it had a large nasal
horn and several small horns on its frill, two of them pointing forward; the frill had
large fenestrae; not related to Kentrosaurus, a stegosaur of the Late Jurassic; Canada,
Late Cretaceous, 77–76 Ma.
Cephalaspis: “head shield,” a genus of freshwater ostracoderm fish in the extinct
order Osteostraci; these primitive armored fish were about 1 foot long and charac-
terized by a flattened headshield and large pectoral fins; with a mouth located under
its head, Cephalaspis probably fed on detritus on the bottom of streams; Early
Devonian.
Cephalochordata: the least numerous subphylum of the chordates, characterized
by a stiff notochord throughout life, as well as a dorsal nerve cord and gill slits.
They are generally fish-like marine animals with a notochord the length of the body,
which ranges from 2 to 6 inches long. Because they have no hard parts, the fossil
record is sparse. However, it extends from Pikaia gracilens of the Burgess Shale to
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the modern lancelets, including dozens of modern genera in the Branchiostomiformes

(also called Amphioxiformes). See also Tunicata. Middle Cambrian to Recent.
Cephalodiscidae: the largest family of small, worm-like animals in the class
Pterobranchia, q.v.; Late Ordovician to Recent.
Cephalogale: “helmet head,” a genus of carnivorous mammals in the family
Ursidae and subfamily Hemicyoninae; among the earliest ancestors of bears, it was
dog-like and the size of a raccoon, and later species became omnivorous;
Cephalogale appeared in Eurasia in the middle Oligocene and in North America in
the early Miocene, and was replaced in the Miocene by more advanced ursids.
cephalon: the front section of an arthropod; in trilobites the cephalon consists of
specialized parts, such as the eyes and glabella, and also the free cheeks in most
genera.
Cephalopoda: a still-extant class of marine mollusks that includes ammonoids,
nautiloids, and coleoids (squids, octopuses, the Spirulida, and cuttlefish); probably
descended from a monoplacophoran such as Knightoconus, q.v. Global distribu-
tion, late Cambrian to Recent.
Ceramoporidae: a family of bryozoans in the extinct order Cystoporata; the early
Ceramoporinae were simple encrusting organisms, while the later Crepiporinae had
branching colonies; Ordovician to Late Devonian.
Ceratiocarididae: a family of phyllocarid crustaceans in the order Archaeostraca;
a new species, Ceratiocaris winneshiekensis, was reported in 2016 as the oldest
known representative of the family; North America, Middle Ordovician to Silurian.
Ceratitida: an order of ammonoids generally characterized by smooth saddles and
frilled lobes. This ceratitic suture pattern consists of strongly indented and some-
times interlocking loops; it is more complex than the earlier goniatitic pattern but
less complex than the ammonitic pattern characteristic of most Jurassic and
Cretaceous ammonoids. The Ceratitida include most of the Triassic ammonoids
(about 400 genera). Global distribution, Permian to Triassic.
Ceratogaulus: a genus of fossorial horned rodents in the family Mylagaulidae, the
only known horned rodent. About a foot long and generally marmot-like, it is nota-
ble for a pair of sturdy, sharp horns on the snout, between the nostrils and the eyes.
The function of the horns is disputed; they may have aided in digging but were prob-
ably used mainly for defense. Ceratogaulus is an example of convergent evolution
with the unrelated Peltephilus of South America, the only other known fossorial
horned mammal. Originally called Epigaulus. Western North America, late Miocene
to early Pleistocene.
Ceratomorpha: a suborder of perissodactyl mammals that incudes the Tapiroidea
and Rhinocerotoidea; characterized by a strong transverse crest on the molars;
global distribution, late Paleocene to Recent.
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Ceratopsia: “horned face,” a large suborder of herbivorous ornithischian dinosaurs

that originated in the Late Jurassic and thrived in the Cretaceous across the Northern
Hemisphere. Only 3 to 7 feet long, the early ceratopsians (such as the basal Yinlong
of the late Jurassic and the Psittacosauridae of the Early Cretaceous) were hornless
and bipedal. The later genera developed a great variety of horns and frills on the
head and neck, and became quadrupedal as they grew larger. The first-named speci-
mens were given the names Ceratops and Proceratops, but these are now consid-
ered dubious names since the specimens have all been assigned to later genera or
discarded as too fragmentary. North America, Europe, and Asia; Late Jurassic to
Late Cretaceous.
Ceratosauria: a large clade of theropod saurischian dinosaurs that arose around
225 Ma; carnivorous, generally bipedal, and at first quite small, by the Middle
Jurassic they had evolved into larger species up to 9 feet long and by the Late
Jurassic others as long as 19 feet; ceratosaurs gave rise to the carnosaurs of the Late
Jurassic, a group of large carnivores that included the very widespread allosaurs;
Late Triassic to Late Cretaceous.
Cercidiphyllum: a still-extant genus of riparian trees that was common in the early
Cenozoic in the northern areas of Asia, Europe, and North America; a relative of the
modern Katsura tree, now found only as two species in Eastern Asia.
Cercopithecoidea: the Old-World monkeys, including macaques, mandrills, and
baboons; classified with humans and great apes in the infraorder Catarrhini; early
Miocene to Recent.
Ceriantharia: a small subclass of anthozoan cnidarians, consisting of only one
order of the same name; called tube-dwelling anemones, they are classified as a
sister-group of the corals; they are almost absent from the fossil record because they
have no hard parts, but an early Cambrian microfossil named Eolympia pediculata
was described in 2010 and classified as a ceriantharian.
cerioid: of corals, having generally polygonal individual corallites packed tightly
together.
Cervalces alaskensis: the stag-moose, a large ancestor of the modern moose;
Alaska, late Pleistocene.
Cervidae: a family of cloven-hoofed ruminant ungulate mammals in the order
Artiodactyla; distinguished from the Bovidae by their solid, deciduous, generally
branching antlers; a probable early ancestor is Diacodexis (middle Eocene), and one
of the earliest cervids is Leptomeryx (Eocene to Miocene); global distribution,
Oligocene to Recent.
Cetacea: a still-extant order of marine artiodactyl mammals that includes the mod-
ern whales, porpoises, and dolphins. Long thought to be descended from the large
group of protoungulate land mammals called mesonychids, the earliest ancestral
whales are now generally considered more closely related to the hippopotamuses.
The direct ancestors of Pakicetus and other early whales are still unclear, however.
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For the evolution of marine mammals, see Aegyptocetus, Ambulocetus,

Basilosaurus, Indocetus, Indohyus, Pakicetus, Pezosiren, Protocetus, and
Rodhocetus. Global distribution, early Eocene to Recent.
Cetartiodactyla: the clade including cetaceans and other artiodactyls.
chaetae: chitinous bristle-like setae found on some insects, especially characteris-
tic of polychaete annelid worms.
Chaetetes: a genus of demosponges in the family Chaetitidae, widespread from
Ordovician to Jurassic.
chaetetids: small tubular organisms formerly classified as anthozoans, specifically
tabulate corals, or even as algae; now considered a polyphyletic group of demo-
sponges. They generally formed globular colonies composed of fused tubular skel-
etons, in mounds sometimes as large as 10 feet. Appearing in the Ordovician and
widespread through the Permian, they had been thought to be extinct at the end of
the Jurassic, but some extant species (such as Acanthochaetetes wellsi) have been
discovered off the northeastern coast of Australia.
Chaetognatha: a phylum of arrow worms known from the Cambrian to the pres-
ent; obscure because of a paucity of whole-body fossils, but a significant zooplank-
tonic component in modern oceans; see also Capinatator and Eognathacantha;
Cambrian to Recent.
Chalicotherioidea: a superfamily of clawed perissodactyl mammals in the subor-
der Ancylopoda, comprising two families, the widespread Chalicotheriidae and the
Eomoropidae, which are known mainly from Asia. Some genera were about the size
of a horse, probably browsers but their habits are not completely known. Their
claws, often long, made movement awkward, but some may have been able to raise
the claws enough to walk on their pads. They are related to brontotheres, horses,
rhinoceroses, and tapirs. Holarctic distribution, early Eocene to Pleistocene.
Chamitataxus: a genus of caniform carnivores in the family Mustelidae and sub-
family Taxideinae; known only from a skull found in New Mexico, Chamitataxus
existed alongside Taxidea, the modern American badger, in the late Miocene.
Champsosaurus: a genus of diapsid reptiles in the order Choristodera and family
Champsosauridae, widespread in the Late Cretaceous and Paleocene; superficially
similar to the modern gharial, Champsosaurus was almost completely aquatic; its
jaws were adapted to catching fish, with powerful muscles attached to its broad
skull; it was generally about 5 feet long, although the largest species, C. gigas, grew
to 12 feet; North America and Europe, Late Cretaceous to Eocene.
Chañares Formation: an important Early Carnian site in northwestern Argentina;
notable for tetrapods and archosaurians, including possible ancestors of the earliest
dinosaurs; the Santa Maria Formation of Southern Brazil also has outcrops of the
same strata; Late Triassic, 235–233 Ma.
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Chancelloriidae: a poorly known group of sponge-shaped Cambrian organisms,

possibly related to Halkieria; sessile and probably filter-feeders, sometimes found
attached to other organisms; they had a bag-shaped body covered with spines and
sclerites, 1–8 inches long and narrow at the bottom; North America, Asia, and
Australia; early to late Cambrian.
character: a distinctive heritable feature or property, usually morphological, that
defines a taxon, differentiating it from other taxa.
Charassognathus: a monospecific genus of small therapsids in the suborder
Cynodontia; the earliest-known cynodont, C. gracilis is known only from a skull
and one leg but is estimated to have been 1–2 feet long; South Africa, late Permian,
260 Ma.
Charnia: an enigmatic genus of Ediacaran animals. It was benthic, probably in a
deep-water environment, and was superficially similar to sea pens. The central stalk
rose from a bulbous holdfast set in sediment. The long, narrow fronds display a
series of fine ridges along the entire length, up to 6 feet. It is found in association
with Aspidella and may have been attached to it. Named for Charnwood Forest in
England, where it was first found; Europe and North America, 579–555 Ma.
Charniodiscus: a feather-shaped Ediacaran animal, the tallest more than a foot
long. Like its relative Charnia, it was benthic and sessile, with a central branch ris-
ing from a bulbous or disc-shaped holdfast. It was a filter-feeder with about 20
secondary branches on each side of the central shaft, with polyps on each branch.
Charniodiscus is morphologically similar to the pennatulaceans of the Cambrian,
but its affinities are uncertain. It has been found as a trace fossil in ferruginous
impressions in the upper surfaces of Ediacaran paleosols in Australia. Global distri-
bution, 565–555 Ma.
Charophyceae: a class of freshwater green algae in the division Charophyta;
Silurian to Recent.
Charophyta: a division of calcareous freshwater green algae; the gyrogonites of
some charophytes are stratigraphically important; early Silurian to Recent.
Chasmatosuchus: synonym: Proterosuchus.
Chasmosaurus: a genus of medium-sized ceratopsid dinosaurs in the subfamily
Chasmosaurinae; up to 17 feet long and herbivorous, characterized by a short nasal
horn and very large fenestrae in the large neck-frill; North America and Asia, Late
Chaunograptus: a possible genus of graptolites, the earliest known, common in
the Burgess Shale, 508 Ma; C. sphaericola has been identified in a Devonian deposit
in Estonia; North America and Europe, middle Cambrian to Devonian.
Cheiracanthus: a small genus of freshwater acanthodian fish in the family
Acanthoessidae; deep-bodied and about a foot long, it is notable for having only one
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dorsal fin; whole-body specimens are known from Europe, and its distinctive scales
have been found as far away as Antarctica; global distribution, Middle Devonian.
Cheirolepis: a basal genus of freshwater actinopterygian fish, the only genus in the
family Cheirolepidae; almost 2 feet long, it is characterized by a heterocercal tail
fin, a moveable jaw that gave it a wide gape, and ganoid scales locked together in a
peg-and-socket design; see also Moythomasia; North America and Europe, Middle
to Late Devonian.
Chelicerata: a class of arthropods that includes the arachnids, scorpions, eurypter-
ids, horseshoe crabs (Xiphosurida), and possibly the sea spiders (Pycnogonida), all
of which have living representatives; several other animals as far back as the Burgess
Shale fauna have been proposed as chelicerates, but the earliest unchallenged ones
(eurypterids) appeared in the Middle Ordovician; named “claw horn” because of
their principal synapomorphy, jointed appendages (chelicerae) which are usually
pincers; Ordovician to Recent.
Chelonian: see Testudines.
Chelydridae: a small family of cryptodiran turtles; it was once known throughout
the Northern Hemisphere, but its range has shrunk so that only two genera of snap-
ping turtles of North and South America are still extant; North America, Europe,
and Asia; Late Cretaceous to Recent.
Chengjiang biota: an important assemblage of early Cambrian fossils found in
several Konservat-Lagerstätten in the Maotianshan Shales. Dated between 530 and
515 Ma, at least 7 million years earlier than the Burgess Shale, the strata are mostly
finely bedded mudstones and preserve a great variety of soft-tissue fossils. In con-
trast to the organisms of the Burgess Shale, those in the Chengjiang sites are gener-
ally autochthonous, preserved in their living position. One of the largest groups is
the arthropods, many of them bivalved crustaceans (see Kunmingella). Several,
such as Waptia and Anomalocaris, are congeneric with the Burgess biota, and spe-
cies of other organisms, such as the lobopods Hallucigenia and Aysheaia, are also
closely related. See also Yunnanozoon. China, early Cambrian.
Chesaconcavus: a genus of barnacles in the still-extant family Balanidae and sub-
family Concavinae; North America and Europe, Miocene to Pliocene.
chevrotain: see Tragulidae.
Chileata: one of the three extinct classes of articulate brachiopods in the subphy-
lum Rhynchonelliformea; characterized by a large perforation in the ventral valve,
they have either strophic or astrophic hinges; Cambrian to Silurian.
Chilobolbinidae: a family of dimorphic ostracods in the suborder Beyrichicopina
and superfamily Hollinoidea, characterized by a frill and a straight hinge; Ordovician
to Silurian.
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Chilopoda: a still-extant class of myriapods that includes the centipedes;

c arnivorous and venomous, they have only one pair of legs on each body segment;
Silurian to Recent.
Chindesaurus: a genus of theropod dinosaurs in the family Herrerasauridae; one
of the first true dinosaurs, it was up to 7 feet long, bipedal, and carnivorous; it is
known primarily from several partial skeletons found in the Chinle Formation of the
Southwestern United States, dated between 221 and 216 Ma; North America, Late
Triassic.
Chiniquodontidae: a family of advanced carnivorous therapsids in the suborder
Cynodontia; closely related to the immediate ancestors of mammals; South America
and possibly Europe and Africa, Late Triassic.
Chiosella timorensis: a species of conodonts whose first appearance is generally
taken (but challenged in 2016) to mark the beginning of the Middle Triassic at
247.2 Ma; Romania and China.
Chippewaella patellitheca: a species of primitive snails, the earliest-known gastro-
pod mollusk; known only from specimens found in Wisconsin, it has a round, cap-
shaped shell about an inch in diameter; see also Strepsodiscus; United States, late
Cambrian, 500 Ma.
Chirodus: a genus of actinopterygian fish in the extinct suborder Platysomoidei
and family Chirodontidae; deep-bodied with very short dorsal and pectoral fins and
a crushing dentition; Carboniferous to Triassic.
Chirognathidae: a family of conodonts in the suborder Prioniodinina, among the
earliest of the suborder; characterized by multiple denticles on the teeth; Ordovician.
Chiroptera: “hand wing,” the bats; the order is divided into two suborders,
Megachiroptera (large frugivorous bats that do not use echo-location) and
Microchiroptera (small echo-locating bats, generally insectivorous); the classifica-
tion of bats is disputed, but they are generally considered closer to primates than to
rodents; early Eocene to Recent.
chitinozoans: an enigmatic group of middle Paleozoic microfossils. Distinctively
flask-shaped, they may be eggs or a juvenile stage of unknown marine animals, pos-
sibly orthocone cephalopods. Occurring worldwide and especially abundant in
Ordovician and Silurian strata, they are important index fossils because of their
wide distribution and rapid evolution. Early Ordovician to early Mississippian.
chitons: see Polyplacophora.
Choeropotamus: a genus of pig-like entelodont artiodactyl mammals, the only
genus in the family Choeropotamidae; related to the Cebochoeridae and possibly
descended from the little-known Helohyidae; late Eocene.
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Choia: a genus of demosponges in the order Monaxonida; about an inch wide, its
conical body was surrounded by a thicket of spicules; the fossils of some species
resemble stars; it was originally thought to have rested on the sea floor but is now
believed to have floated, attached to the substrate by three long spines; Canada,
United States, China, and Africa; Cambrian to Ordovician.
Chondrostei: a still-extant, paraphyletic subclass of primarily cartilaginous fishes
in the class Actinopterygii. The chondrosteans flourished in the late Paleozoic and
the Triassic, declining in the Cenozoic. Some Mesozoic families have essentially
ossified skeletons and other similarities to the teleosts, so that the modern chondro-
steans’ cartilaginous condition is probably derived. Cosmopolitan, Silurian to
Recent.
Chonetes: a long-lived genus of articulate brachiopods in the extinct order
Productida and superfamily Chonetoidea; Chonetes was characterized by a row of
short, angled spines along its margin that it used to anchor itself; Late Ordovician to
Jurassic.
Chordata: the phylum of deuterostome organisms that includes the subphyla
Vertebrata, Cephalochordata, and Tunicata; characterized by a dorsal nerve cord
and pharyngeal apertures (gill slits); early Cambrian to Recent.
Choristodera: an order of semi-aquatic diapsid reptiles, possibly in the
Archosauromorpha; similar to modern gharials, the choristoderans are represented
principally by the families Champsosauridae and Simoedosauridae, which appeared
in the Cretaceous and survived the end-Cretaceous extinction into the middle
Eocene; champsosaurs are significant especially for specimens found near Ellesmere
Island, high in the Canadian Arctic; North America, Europe, and Asia; Middle
Jurassic to Miocene.
Chromogisaurus: a monospecific genus of sauropodomorph dinosaurs in the fam-
ily Guaibasauridae; about 7 feet long and dated at 231 Ma, it is one of the earliest
dinosaurs; C. novasi is known from one specimen, a partial disarticulated skull and
skeleton found in a middle Carnian layer of the Ischigualasto Formation; Argentina,
Late Triassic.
chronocline: a time series of fossil specimens showing transitional morphological
changes in a taxon in successive strata.
Chronoperates paradoxus: “puzzling time-wanderer,” a problematic Paleocene
fossil jaw; originally thought to be an extremely late survivor of the cynodonts, the
therapsid ancestors of mammals which became extinct 100 million years earlier,
midway through the Cretaceous; it is now considered a late-surviving symmetro-
dont mammal; Alberta, late Paleocene.
Chrysochloridae: a family of insectivorous burrowing mammals in the order
Afrosoricida; called golden moles, they are so like the marsupial moles of Australia
that some researchers have claimed they are related; Southern Africa, early Miocene
to Recent.
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Cidaroida: a still-extant order of primitive sea urchins in the subclass

Perischoechinoidea, the only order of the subclass to survive into the Cenozoic Era;
characterized by large and widely separated primary spines; early Permian to
Recent.
Cimolesta: a proposed order of eutherian mammals; related to several other early
groups of mammals such as the creodonts, dinoceratans, and pantodonts; North
America, Europe, Asia, and Africa; Cretaceous to Eocene.
Cimoliopterus: a genus of pterodactyloid pterosaurs with a wingspan of about
16 feet; the two species are C. cuvieri (England) and C. dunni (Texas), and they are
closely related to a slightly older pterosaur from Texas, Aetodactylus halli; Early
Cretaceous, 94 Ma.
cinctans: an enigmatic group of slow-moving early Paleozoic animals thought to
be non-pentameral basal echinoderms; sometimes called Homostelea; only a few
centimeters long, they are composed of interlocking, generally rectangular plates
arranged in an oval shape resembling a small tennis racket, with a short extension at
the narrow end; Europe and North Africa, middle Cambrian, 509–497 Ma.
Cirripedia: the barnacles, a subclass of marine crustacean arthropods traceable to
the Cambrian but uncommon in the fossil record until the middle Cenozoic. The
shell is typically constructed of six triangular plates in a conical shape. They are
generally sessile and cemented to the substrate by a firm base, living in the highly
erosional tidal zone. Since only a few genera live in deep water, they are sometimes
used as indicators of paleodepths, estimated by the degree of disarticulation of the
shells. See also Chesaconcavus and Priscansermarinus. Global distribution, mid-
dle Cambrian to Recent.
Citipati: a genus of theropod saurischian dinosaurs in the family Oviraptoridae;
they are carnivores similar to Troodon and Oviraptor, notable for bird-like repro-
ductive behavior; Mongolia, Late Cretaceous, 84–75 Ma.
clade: a monophyletic group of two or more species with a common ancestor, as
well as all the other descendants of that ancestor.
cladodont dentition: broad-based teeth with a large primary cusp and several sec-
ondary cusps, common in Paleozoic sharks but little known after the Triassic.
cladogenesis: see anagenesis.
Cladoselachidae: an early family of sharks; about 6 feet long and almost scaleless,
they were characterized by cladodont dentition, two dorsal fins, and large and
paddle-like pectoral fins, but they had no anal fin; Devonian to Carboniferous.
Cladoxylon: a genus of early vascular plants in the large group called cladoxylop-
sids; it grew only about a foot high and may have been leafless, but its central stem
marks it as a forerunner of sizeable trees that appeared in the Devonian; global
distribution, Middle Devonian to Early Carboniferous.
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Clarkecaris: a genus of crustaceans in the order Anaspidacea and proposed family

Clarkecarididae; Brazil, early Permian.
Clarno Nut Beds: a botanically rich site near John Day, Oregon; although the Nut
Beds of the Clarno Formation contain few vertebrate fossils, they are one of the
most important paleobotanical sites in the world; they contain hundreds of well-
preserved botanical fossils from a large forest of the Eocene, dated at 44 Ma; North
America, middle Eocene.
Clathropteris meniscioides: a species of reticulate ferns, known worldwide from
the Late Triassic to Middle Jurassic.
Clavatoraceae: a Mesozoic family of charophytic algae; several species represent-
ing gradual evolutionary changes are index fossils from the Late Jurassic to the end
of the Cretaceous.
Cleveland-Lloyd Dinosaur Quarry: an important Late Cretaceous site, espe-
cially for dinosaurs. Located on the site of a boggy marsh dated at 147 Ma, the
quarry’s Morrison Formation strata have produced thousands of disarticulated dino-
saur bones. There are also dinosaur tracks in the vertical cliffs and specimens from
all other vertebrate classes. Of the dinosaurs, many more carnivores than herbivores
are represented. Only a few whole skeletons have been found, evidently because of
the slow turbidity of the clay sediments in the bog. Central Utah, Late Jurassic.
Climacograptus: a genus of scandent graptolites important as index fossils in the
Ordovician; characterized by a single serrated stipe; Middle Ordovician to Lower
Devonian.
Climatiiformes: an order of small, spiny acanthodian fish, only about 3–4 inches
in length; they had several stout spines and a heterocercal tail fin; global distribu-
tion, Silurian to Carboniferous.
Climatius: a primitive genus of acanthodians in the family Climatiidae; although
only about 3 inches long, it had 15 spines on its body and fins; North America and
Europe, late Silurian to Early Devonian.
Cloudina: an Ediacaran microfossil found as a tube with double-layered walls of
calcium carbonate; thought to have been made by a suspension-feeding worm;
named for American paleontologist Preston Cloud; Namibia, late Ediacaran.
Clymeniida: a large early order of ammonoids, notable for an unusual dorsal
siphuncle, as opposed to the ventral siphuncle of other ammonoids; less than 2 inches
across and very diverse in shell type and ornamentation; they diversified rapidly in
the Famennian, the final stage of the Devonian, but then abruptly disappeared at the
end of the period; Europe, North Africa, and possibly Australia; Middle to Late
Devonian.
Clypeasteroida: the modern sand dollars, an order of euechinoids in the superor-
der Gnathostomata; global distribution, late Paleocene to Recent.
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Clypeus ploti: “swimming shield,” a species of irregular echinoids in the family

Clypeidae, called the “pound-stone” because it fossilized as a rounded, fist-sized
stone of about 22 ounces; common in eighteenth-century Oxfordshire, England, it
was among the first echinoids studied; Europe and Africa, Middle Jurassic.
Cnidaria: a phylum of diploblastic animals, mostly marine, usually with stinging
tentacles around a mouth; comprising chiefly the corals, sea anemones, and jelly-
fish, the phylum includes more than 10,000 named species, divided into three
classes: Anthozoa, Scyphozoa, and Hydrozoa; Ediacaran to Recent.
coccolithophore: a one-celled planktic algae that surrounds itself with a many-
layered calcite skeleton (a coccosphere); the skeletons are the major component of
chalk deposits around the world, and coccolithophores are considered the main cal-
cite producer in the ocean; significant in the fossil record since the Triassic, espe-
cially as an indicator of climate change.
Coccosteus: an important Devonian genus of arthrodire placoderms in the extinct
suborder Brachythoraci; similar in some ways to its much larger cousin Dunkleosteus,
Coccosteus was 8 to 16 inches long and is found mainly in freshwater deposits; it
had large pectoral and pelvic fins and a long finned tail; the arthrodires were joint-
necked fish, and Coccosteus had two joints, which allowed it to feed on large prey;
Europe and North America, Middle to Late Devonian.
Cochliodontidae: a large family of elasmobranch fish similar to sharks but with a
bradyodont toothplate dentition; Devonian to Permian.
Coelacanthidae: an extinct family of actinistian lobe-fin fish in the order
Coelacanthiformes; Permian to Jurassic.
Coelacanthiformes: an order of lobe-fin fish, considered extinct since the
Cretaceous until two species of the genus Latimeria were discovered in the Indian
Ocean in 1938. Latimeria (Latimerioidei: Latimeriidae), which appeared in the
Triassic, is very similar to the Cretaceous coelacanths. Even with a poor fossil
record, coelacanths are significant in evolution because they are closely related to
the early lobe-finned fish, which are thought to be the first fish to develop air-
breathing skills that let them begin to live on land about 400 Ma. Devonian to
Recent.
Coelenterata: an obsolete term, formerly considered a phylum that included a
large group, the Cnidaria, and a smaller group, the Ctenophora; these are now con-
sidered separate phyla.
Coelodonta: a genus of large perissodactyl mammals in the extant suborder
Ceratomorpha and subfamily Rhinoceratinae; coelodonts were the woolly rhinocer-
oses of Eurasia, surviving up to about 10,000 years ago; they were similar in size to
the modern rhinoceros, typically about 13 feet long and weighing 3 tons, with some
almost 7 feet high; they bore two forward-pointing horns on the nose; Europe and
Asia, Pliocene to late Pleistocene.
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Coelophysis a genus of early saurischian dinosaurs, theropods in the family

Coelophysidae; bipedal, lightly built, and about 10 feet long, it is a typical coeluro-
saur; there is some inconclusive evidence that it was gregarious, perhaps hunting in
packs; North America, Late Triassic to Early Jurassic.
coelurosaur: “hollow-tailed lizard,” a large group of small, lightly built, theropod
dinosaurs in the order Saurischia; these bipedal predators, some of them feathered,
gave rise to later saurischians such as the Ornithomimidae and Tyrannosauridae,
and to birds; Triassic to Late Cretaceous.
Coenopteridales: a large and varied group of ancestral ferns, usually considered
as belonging to the order Primofilices; Devonian to Triassic.
coevolution: an evolutionary process in which unrelated lineages exert selective
pressure on each other, resulting in evolutionary changes in both. The best-
documented cases are the interactions between insects and entomophilous and ento-
mophagous flowers, and between birds and ornithophilous flowers, but many other
cases have been described of coevolutionary relationships between predatory ani-
mals and prey and between herbivorous animals and plants.
Coleodontidae: a family of conodonts in the order Neurodontiformes, with teeth
characterized by a lamellar structure; Ordovician to Triassic.
Coleoidea: a subclass of soft-bodied mollusks in the class Cephalopoda, including
the extinct belemnites and the still-extant squids, octopuses, and cuttlefish; disputed
specimens are known from as early as the Cambrian, but the earliest definitive cole-
oids are dated as Mississippian.
Coleoptera: the beetles, an order of pterygote insects in the clade Neoptera. They
are named “sheathed wing” for the hard elytra, the front pair of wings that protect
the membranous back wings and the abdomen. The oldest definitive beetle dates
back to the Permian, but a possible specimen has been found at the Late Carboniferous
Mazon Creek site. Following the end-Permian extinction, the Coleoptera diversified
rapidly in the Triassic. Global distribution, Pennsylvanian? to Recent.
Collembola: a still-extant order of primitive apterygote arthropods; up to half a
centimeter long, they are commonly called springtails because of the jumping abil-
ity given them by a furcula, a tail-like forked appendage at the end of the abdomen,
normally folded under the body; known from the Early Devonian Rhyniella prae-
cursor and other fossil specimens beginning in the Cretaceous.
Collenia: a widespread genus of cyanobacteria that produced stromatolites;
Precambrian.
Collignoniceras: a small genus of ammonoids in the order Ammonitida and super-
family Acanthoceratoidea; up to 7 inches across, evolute, and very strongly ribbed,
Collignoniceras is the type genus of the family Collignoniceratidae; global
distribution, Late Cretaceous, 96–88 Ma.
Collinsium ciliosum: see Hallucigenia.
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Collyrites: a genus of small irregular euechinoids in the family Collyritidae,

p ossibly paraphyletic; they were oval, up to an inch long, and did not have a lantern;
Europe and Asia, Middle Jurassic to Early Cretaceous.
Columba omnisanctorum: an extinct species of early Pliocene pigeons in the
order Columbiformes; found on the Gargano Peninsula of Italy, related to a similar
columbiform found nearby, C. pisana; see also Deinogalerix.
Columbia: see Rodinia.
Colymbosathon ecplecticos: a species of ostracods notable for the well-preserved
soft-tissue features in one intact specimen buried by volcanic ash about 425 Ma and
found in a carbonate nodule. Although the fossil is only 0.2 inches long, organs vis-
ible in a digital three-dimensional image include eyes, gills, a circulatory system,
anus, and penis. The presence of the penis makes this specimen the oldest known
definitively male fossil. See Herefordshire Nodules. England, Silurian.
comb jellies: see Ctenophora.
Como Bluff: a Konzentrat-Lagerstätte on a ridge in Wyoming noted for the many
dinosaurs discovered there in strata of the Morrison and Sundance Formations;
United States, Middle and Late Jurassic.
Composita: a genus of brachiopods in the extinct order Athyridida; from 1 to
2 inches in diameter, smooth-shelled, with a distinct fold and sulcus; global distri-
bution, Late Devonian to Early Triassic.
compound eye: an eye formed of a mosaic of smaller eyes, characteristic of many
arthropods.
Compsognathus: “dainty jaw,” a genus of theropod saurischian dinosaurs; bird-
like, small and light in build, its body was as long as 2 feet and it weighed up to 8
pounds; a fast runner, it had long hind legs and a longer tail; Jurassic.
concretions: hard, layered structures, usually rounded, formed by precipitation
and cementation of minerals in the soil or rock, sometimes enclosing a fossil.
condylarths: an informal group of primitive placental mammals ancestral to the
perissodactyl ungulates and to the cetaceans. Previously considered an order, the
condylarths have all been assigned more precisely to different orders. Arising in the
Cretaceous, they underwent an extensive radiation of species in the Paleocene and
Eocene. They were mainly herbivorous, but some had hooves and some had claws.
Global distribution except Australia, Late Cretaceous to Oligocene.
condyle: an articular process on the joint of a bone (or mandible of an insect).
Confuciusornis: a genus of primitive birds in the family Confuciusornithidae, of
which C. sanctus is the type species. Crow-sized, it is the earliest-known toothless
beaked bird and seems to be the earliest that could fly for a considerable distance.
About 11 inches high and bearing true feathers, it was similar to Archaeopteryx
except that it was toothless and probably had more feathers. Three other species
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have been named, all found in the Yixian and Jiufotang Formations of northeastern
China (see Jehol biota) and dated between 130 and 125 Ma, 10–15 million years
after Archaeopteryx. Confuciusornis was probably not in the line of descent to mod-
ern birds. China, Early Cretaceous.
Coniconchia: a proposed class of mollusks to include the hyoliths and tentacu-
lites; 2017 research, however, indicates the hyoliths are lophophorates and closely
related to brachiopods; the tentaculites have generally been considered as an order
in their own right, the Tentaculitida.
conifers: see Pinopsida.
Coniopteris: a genus of Mesozoic ferns in the still-extant family Dicksoniaceae; it
bore lanceolate or lobate leaves 2–4 inches wide; its spores were carried marginally,
in cup-shaped pouches at the end of lateral veins; especially common in the Jurassic;
global distribution, Permian to Cretaceous.
Conocardium: a genus of rostroconch mollusks in the family Conocardiidae; up to
6 inches long, its thick shell surface is marked by fine rays; global distribution,
Conoclypus: a genus of euechinoids in the order Clypeasteroida and family
Conoclypidae, sand dollars with a high test about 1.5 inches across and petaloid
ambulacra; Europe, early Eocene to Miocene.
Conodonta: a class of enigmatic eel-like chordates. Although considered verte-
brates, they are still known mainly by their feeding apparatus, referred to as con-
odont elements; fossil specimens show no soft-tissue detail. Their jaw structure
differs greatly from that of later animals. They are now classified as chordates and
divided into protoconodonts, paraconodonts, and euconodonts. Appearing in the
early Cambrian, they ranged from an inch to more than a foot long. Only a few
whole-body specimens are known, the best-known of which is Promissum pul-
chrum, q.v. Many conodonts became extinct at the end of the Ordovician, but they
recovered and continued to be important stratigraphically through the Paleozoic,
becoming completely extinct at the end of the Triassic.
conodont elements: the earliest tooth fossils, in the form of rudimentary teeth and
very rudimentary jaws; despite the inability until recently to discover whole-body
specimens of the animal which bore them, conodont elements have long been
important in stratigraphic research; see for example Iapetognathus and
Streptognathodus.
Conophyton: one of many plant-like organisms that formed stromatolites in the
Mesoproterozoic Era; North America and Africa, Archaean.
Constellaria: a genus of stenolaemate bryozoans in the extinct order Cystoporata
and family Constellariidae; they formed branching colonies and are distinguished
by generally star-shaped maculae on the surface of branches about half an inch
across; global distribution, Middle Ordovician to early Silurian.
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continental drift: see paleogeography.

Conulariida: a problematic clade of extinct cnidarians, generally considered scy-
phozoans but not well understood. Their superficially shell-like structure consists of
rows of calcium phosphate rods that form a quadrilateral cone, with new rods being
added at the broader end. The surface has a wavy ridged appearance, with the rows
of rods intersecting at a slight angle in the middle of each side. The narrow end was
attached to the substrate, and soft tentacles extended from the broad end. The larger
conulariids were as long as 20 inches, but most were about 4 inches. Also called
Conulata and Conularia. Late Cambrian to Early Triassic.
Conulidea: a superfamily of echinoids in the order Echinoneoida; Europe, Asia,
and North America; Early Cretaceous to Eocene.
Conulus: a small genus of conulidean euechinoids in the order Echinoneoida and
family Conulidae; dorsally high-domed, almost conical, smooth and non-petaloid;
the flat oral side has a faint pentameral pattern; Europe, Asia, North America; Early
to Late Cretaceous.
convergent evolution: the process in which unrelated lineages living in similar
ecological niches independently develop superficially similar adaptations.
Cooksonia: a genus of tracheophytic plants in the family Rhyniaceae; it is the old-
est demonstrably vascular plant and is close to the point in time when the lycopsids
branched off from other plants; cosmopolitan, middle to late Silurian.
Cope’s Rule: the name given to the observation that groups of animals tend to
evolve toward larger size over evolutionary time. The idea is associated with the
nineteenth-century paleontologist Edward Drinker Cope and has been elaborated by
many others. The tendency is obvious in many lineages but absent in others, and of
course must be subject to limitations of sufficient food sources and of the ability to
move efficiently. See also Unspecialized Descent.
Copodus: a little-known genus of primitive holocephalic fish in the extinct family
Copodontidae; bradyodont, with one tooth plate in each jaw; North America and
Europe, Devonian and Carboniferous.
coprolites: fossilized residues from the digestive tracts of animals. Their contents
may provide evidence of the existence of other organisms, and they are important in
investigating eating habits of animals, but it is difficult to identify the producer. The
largest known specimen is about 17 inches long, found in Saskatchewan in 1995,
thought to have come from a large tyrannosaur just before the end of the Mesozoic
era. Usually in the form of dung, coprolites may also be regurgitalites (regurgitated)
or cololites (fossilized inside the body).
corallite: the chamber or skeleton constructed by an individual coral animal.
corals: marine animals that secrete corallite skeletons composed of calcium car-
bonate, often in conjunction with algae; they generally thrive in warm, shallow
water, but some cold-water corals that do not need algae are found as deep as
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3,000 feet; see Rugosa, Tabulata, and Scleractinia; worldwide, Cambrian to

Recent.
Cordaicarpus: the form genus of seeds of the family Cordaitaceae.
Cordaitales: an order of primitive gymnosperm plants in the class Pinopsida; they
were tree-like, growing as high as 100 feet, with long, strap-like leaves; possibly
ancestral to conifers, cycads, and ginkgos; global distribution, Carboniferous to
Late Triassic.
Coronacollina: the earliest-known animal with skeletal body parts, possibly
related to sponges. A genus of benthic organisms 1–2 inches high, with 3 spicules
radiating from a central thimble-shaped body; the long spicules probably supported
the animal on the ocean floor. Its resemblance to the Cambrian sponge Choia indi-
cates a link between the two periods. Described in 2012 from hundreds of speci-
mens found in upper Ediacaran strata in Australia, 560–550 Ma.
Coronodon havensteini: a species of mysticete whales known only from a large,
complete skull with unusual dentition. It has strikingly ridged molar teeth that show
no wear from cutting and shearing prey, leading some to consider Coronodon evi-
dence of a transition from the toothed condition of the early odontocete whales to
the baleen of the mysticetes. The very large molars, 3–4 inches in diameter, might
have sieved food from the water as baleen does in modern mysticetes; however, this
interpretation is in dispute. United States, early Oligocene.
Corynexochida: a diverse order of early trilobites that appeared at about the same
time as the short-lived Redlichiida and the Ptychopariida; generally characterized
by large eyes and large glabellae extending nearly to the front of the cephalon; early
Cambrian to Middle or Late Devonian.
Coryphodontidae: “peaked tooth,” a family of pantodont mammals in the extinct
order Cimolesta; the type genus Coryphodon was 3 feet at the shoulder and about
8 feet long, and some genera were larger; although the canine teeth were relatively
long, it was a forest browser; North America, Europe, and Asia; late Paleocene to
middle Eocene.
Corystospermaceae: a family of seed plants in the order Peltaspermales; similar
to seed ferns, but the relationship is obscure; characterized by pairs of heavily
veined leaflets on a central stalk; widespread in the Triassic throughout Gondwana
and present also at least in China; Early to Late Triassic.
Corythoraptor jacobsi: a species of feathered theropod dinosaurs in the family
Oviraptoridae, known from one almost-complete skeleton. Described in 2017, C.
jacobsi was 5–6 feet long, with small forelimbs and strong hindlimbs, probably
flightless but a fast runner. In shape it resembled the modern cassowary, especially
because of a tall, characteristic crest. It is one of several oviraptoroids found recently
in the Nanxiong Formation; Southern China, Late Cretaceous, 84–66 Ma.
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Corythosaurus: “Corinthian-helmet lizard,” a hadrosaurid dinosaur in the

subfamily Lambeosaurinae; similar to Lambeosaurus, it was about 30 feet long,
with a stocky body and long tail; its most notable feature was a tall crest on the top
of its head that may have made a loud trumpet call possible; North America, Late
cosmopolitan distribution: existing in appropriate but scattered environments
across all or most of the world.
Cotylorhynchus: a genus of synapsids in the family Caseidae; it was 10–12 feet
long, with a large, barrel-shaped body and a very small head; the type species C.
romeri was widespread in the southern part of North America in the early Permian;
North America, Carboniferous to middle Permian.
cotylosaurs: an informal term for the paraphyletic Captorhinida.
craniate: having a specialized and distinctly formed head, distinguishing most
chordates from tunicates and cephalochordates.
Crassatella: a long-lived genus of articulate marine burrowing bivalves; Crassatella
was 1.5 inches long, with a strong shoulder on both shells and strong concentric
ribs; Cretaceous to Recent.
Crato Formation: a Lower Cretaceous Konzentrat-Lagerstätte in the Araripe
Basin of Northern Brazil, dated uncertainly but probably between 115 and 112 Ma,
slightly earlier than the neighboring Santana Formation. The occurrence of the
Crato fossils differs from that of the Santana fossils, which are found in limestone
concretions. Those of the Crato, found in finely bedded limestone similar to that of
Solnhofen, are predominantly arthropods, especially insects and arachnids, with
some specimens of fish such as Dastilbe and crested pterosaurs such as Anhanguera.
There are other vertebrates and a variety of plants including gymnosperms and other
early seed plants, as well as a few angiosperms.
Craugastor: a still-extant genus of tree frogs known from amber fossils found in
the Dominican Republic and Mexico, about half an inch long; late Oligocene to
Recent.
creodonts: an informal term for a polyphyletic group of flesh-eating mammals
formerly considered an order, including the Oxyaenidae and Hyaenodontidae; the
diverse group included carnivores as small as squirrels and as large as bears; Late
Cretaceous to Miocene.
Crepiporinae: see Ceramoporidae.
Cretaceous: the third and final period of the Mesozoic Era, from 146 to 66.0 Ma,
divided into Early (146–100) and Late (100–66.0) Epochs. It was a period of high
levels of carbon dioxide, very little polar ice, and the highest sea levels in Earth’s
history, with shallow, saline seas covering much of the land. The first angiosperms
appeared early in the period and by the end of the Cretaceous were the dominant
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plants worldwide. Insects and spiders diversified rapidly along with the angiosperms.
Dinosaurs continued their reign on land, and marine life was dominated by the
greatly diversified bony fish and large predators such as sharks and mosasaurs.
Cretaceous extinction: the most recent and second-most severe of the five great
extinctions, ending the Mesozoic Era at about 66.0 Ma. The most striking result was
the disappearance of the last of the dinosaurs and pterosaurs. Other large terrestrial
animals were also decimated, so that the largest that survived into the Paleocene was
about the size of a turkey. In the seas, the last of the ammonoids and other groups
became extinct, as well as the plesiosaurs and mosasaurs. As to possible causes, the
preponderance of evidence points to a large asteroid striking the Earth off the
Yucatan Peninsula and causing global environmental disruption, and the volcanic
eruption creating the basalt flows called the Deccan Traps in India may have con-
tributed; but, as with the four earlier mass extinctions, the causes are uncertain.
Although the number of dinosaur species decreased during the last several million
years of the Cretaceous, the evidence for a relatively abrupt extinction event is
stronger than in the other mass extinctions.
Cretaceous Inland Seaway: see Western Interior Seaway.
Cretaceous Terrestrial Revolution: abbreviated as KTR, the period of angio-
sperm radiation from 125 to 90 Ma, when flowering plants, insects, and spiders
radiated dramatically, leading to an accompanying radiation of many other
animals.
Cretoxyrhinidae: an extinct family of very large Cretaceous sharks in the still-
extant order Lamniformes; up to 23 feet long and characterized by very sharp
3-inch-long teeth, they preyed on mosasaurs, bony fish, and turtles; cosmopolitan,
Crinoidea: commonly called “sea lilies” but actually animals, a class of echino-
derms in the subphylum Crinozoa. There are 100 extant genera and 600 extant spe-
cies of crinoids. About 7000 extinct species are known, with the greatest diversity in
the Paleozoic. They may have arisen from the Edrioasteroidea or from the eocri-
noids and their descendants the cystoids. Crinoids were widespread and numerous
throughout the Paleozoic up to the end-Permian extinction, and many Paleozoic
limestones are made up almost entirely of crinoid fragments. There are stalked,
unstalked, and free-swimming genera, but the classic Paleozoic genera were stalked.
Crinoids declined rapidly in the Mesozoic for poorly understood reasons. Early
Ordovician to Recent.
Crocodylia: a still-extant order of predatory, semi-aquatic reptiles; global distribu-
tion, Late Cretaceous to Recent.
Crocodylomorpha: a large superorder of archosaurs that includes the crocodilians
and their extinct relatives, going back to the sphenosuchians of the Late Triassic;
global distribution, Late Triassic to Recent.
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Cro-Magnon: the common name for the hominins who competed with
Neanderthals in Europe after 40,000 years ago; now referred to as “European early
modern humans,” the Cro-Magnons are classified in the same subspecies as modern
humans, H. sapiens sapiens, because the skeletal differences between the two are
insignificant.
Crossopterygii: see Sarcopterygii.
crown group: a monophyletic group comprised of a set of extant species, together
with their ancestors back to their latest common ancestor and all other descendants
of that common ancestor; see also stem group.
Crustacea: a large subphylum of arthropods that includes crabs, lobsters, shrimp,
krill, barnacles, ostracods, branchiopods, and other similar animals; middle
Cambrian to Recent.
Cryogenian Glaciation: A period of global cooling around 650 Ma; see acri-
tarchs and icehouse conditions.
Cryolophosaurus: a genus of early theropod dinosaurs found in Antarctica in
1991, probably tetanuran but with some dilophosaurid features, distinctively a thin
bony crest across the top of its forehead. Weighing half a ton and about 21 feet long,
Cryolophosaurus had a shallow skull and a long, stiff tail. It is known only from an
immature specimen, but it was one of the largest theropods of its time and was prob-
ably an efficient predator. Antarctica, Early Jurassic, 190 Ma.
Cryptoclidus: a genus of slender plesiosaurs in the superorder Sauropterygia;
Europe, Asia, and South America; Middle Jurassic.
Cryptodira: a still-extant suborder of turtles in the order Testudines, including
most of the modern turtles and tortoises. They differ from the Pleurodira in several
ways, notably by retracting the head inside the shell by vertical flexure, directly
back instead of to the side. They evolved along with the pleurodires in the Early
Jurassic, and by the end of the period had largely replaced them. Global distribution
except Australia, Early Jurassic to Recent.
Cryptograptus: a genus of scandent pterobranch-like graptolites in the suborder
Glossograptina; cosmopolitan, Middle Ordovician to late Silurian.
Cryptolithus: a genus of asaphid trilobites in the family Trinucleidae; almost cir-
cular in shape and eyeless, but with a pitted fringe around its head that may have
allowed it to sense the direction of ocean currents as it fed on small animals in ocean
sediments; in addition to this “lace collar,” Cryptolithus was remarkable for its pair
of long genal spines; it became extinct in the Ashgill event, 440 Ma; cosmopolitan,
Late Ordovician.
Cryptostomata: an order of Paleozoic colonial bryozoans in the still-extant class
Stenolaemata; the zoaria are short but erect, and the colony forms bifurcating
branches; Ordovician to Permian.
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Cryptovolans: see Microraptor.

Cryptozoon: an enigmatic genus of stromatolitic reef-forming algae common in
the Cambrian and Ordovician, possibly still-extant in the stromatolites of Shark
Bay, Australia.
Ctenodactylidae: a still-extant family of rodents in the suborder Hystricomorpha,
named for the stiff bristles on their hind feet. The earliest definitive specimen is
from the late Eocene, but DNA evidence indicates that the first Ctenodactylidae
appeared in the early Eocene in Asia. The family is still represented by several pika-
like North African genera, called gundis, which are commonly called “comb-rats.”
Asia, Europe, and Africa; Eocene to Recent.
Ctenodonta: a genus of burrowing bivalve mollusks, about 1 inch long, in the sub-
class Palaeotaxodonta and order Nuculida; Early Ordovician to Silurian.
Ctenodus: “comb-tooth,” a genus of sarcopterygian fish in the order Dipnoi; ances-
tral to the modern lungfishes in a well-documented evolutionary line; North
America, Europe, and Australia; Early Carboniferous.
Ctenophora: the comb-jellies, a phylum of gelatinous marine animals that move
by the beating motion of their cilia; generally bell- or ribbon-shaped, they are her-
maphroditic and predatory, feeding on zooplankton and fish larvae; global distribu-
tion, early Cambrian to Recent.
Ctenothrissiformes: a small order of teleost fish in the subclass Neopterygii, well-
known from the genera Ctenothrissa and Aulolepis; appearing only in the
Cenomanian Stage of the Late Cretaceous, they seem to be close to the ancestry of
most modern bony fish; deep-bodied and about a foot long, characterized by ctenoid
scales and large dorsal and pectoral fins without true spines; Europe and Southwest
Asia, Late Cretaceous, 95–90 Ma.
Ctenurella: “small tail comb,” a genus of benthic ptyctodontid placoderm fish;
about 5 inches long, it was lightly armored, like the other ptyctodontids; it had a
whip-like tail and two dorsal fins, the one near the tail comb-shaped, long and low;
it had small toothplates and was probably viviparous; Germany and Australia, Late
Devonian.
cursorial: of terrestrial animals, adapted to rapid running movement in open
country.
Cuvieronius: a genus of New-World proboscideans in the extinct family
Gomphotheriidae; a small gomphothere, Cuvieronius (also known as Cordillerion)
was 9 feet high, weighed about 4 tons, and was notable for its two spirally twisted
tusks; it lived in North America from the Pliocene up to the end of the Pleistocene,
and in South America until about 9,000 years ago, the latest specimen being found
in Chile; Pliocene to Holocene.
Cyanophyta: a phylum of bacteria, one of the earliest forms of life on Earth, found
in Precambrian deposits as old as 3 billion years. Since they produce oxygen through
C 69
photosynthesis, they may have significantly changed the composition of the

atmosphere when they arose. Also called cyanobacteria and “blue-green algae”
(although not algae at all).
Cyathaspidiformes: an order of jawless heterostracan fish, the second-largest
group of heterostracans; fusiform and heavily armored, they were all benthic;
Middle Ordovician to Middle Devonian.
Cycadales: one of the two orders of the Cycadopsida, the Cycadales include the
living cycads and their immediate ancestors; they are subtropical plants with large,
stiff, fernlike leaves, forming a crown around the top of the stem; cycads are gym-
nosperms and are not related to palms, which are angiosperms; early Permian to
Recent.
Cycadeoidales: the second order of tree-like plants in the Cycadopsida; the
Cycadeoidales include the many extinct cycad-like plants that were widespread,
along with the true cycads, throughout the Mesozoic; Permian to Cretaceous.
Cycadofilicales: see Pteridospermatophyta.
Cycadopsida: the class of early seed plants that includes principally the cycads;
early Permian to Recent.
Cyclacantharia: a genus of productid brachiopods variously placed in the family
Richthofeniidae or the family Cyclacanthariidae; with a large conical valve
cemented to the substrate, it is an aberrant form, characterized by many prop-like
spines; known especially from specimens found in the Glass Mountains of Texas,
with the calcite shell replaced by silicification; United States, Africa, and Asia; early
to late Permian.
Cyclida: an enigmatic order of arthropods, formerly considered crustaceans but
now classified as maxillopod branchiurans, related to modern fish-lice; originally
called Cycloidea, they are crab-shaped with a circular carapace 1–3 inches in diam-
eter; a dozen genera are recognized, including Apionicon, Carcinaspides, Halicyne,
and Hemitrochiscus; Europe and North America, Carboniferous to Late Cretaceous.
Cyclocystoidea: a small class of echinoderms in the subphylum Echinozoa, for-
merly placed in the class Edrioasteroidea; small and disc-shaped, characterized by
a marginal ring of ossicles around a plated disc; North America and Europe, Middle
Ordovician to Early Carboniferous.
Cyclomedusa: a very contentious fossil widespread in the Ediacaran biota, possi-
bly related to Aspidella. Circular in shape and up to 10 inches in diameter, it is
generally flat, with many concentric growth ridges, and some specimens display
fine radial lines and possible stems. It is variously considered a holdfast, possibly
from an octocoral-like organism; a colonial microbial organism; or an anemone-like
benthic polyp. Global distribution, Ediacaran.
Cyclopygidae: a family of small to medium-sized trilobites in the order Asaphida;
characterized by very large eyes and found mainly in deep-ocean sediments; along
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with many other groups of trilobites, it became extinct at the end of the Ordovician;
global distribution, Early to Late Ordovician.
Cyclostomatida: a still-extant order of bryozoans in the class Stenolaemata; domi-
nant in the Mesozoic and declining since then, but still widespread; also called
Tubuliporata; Early Ordovician to Recent.
Cyclothyris: an obscure extinct genus of articulate brachiopods in the still-extant
order Rhynchonellida; subtriangular shape, with an erect beak; cosmopolitan,
Cretaceous.
Cynodesmus: a genus of dog-like mammals in the still-extant family Canidae and
extinct subfamily Hesperocyoninae; it was long considered the immediate ancestor
of Tomarctus, but studies since the 1990s indicate a more distant relationship; North
America, late Oligocene to early Miocene.
Cynodictis: a genus of carnivorous mammals in the family Amphicyonidae, pos-
sibly ancestral to many modern carnivores; Europe and Asia, late Eocene to early
Oligocene.
Cynodontia: “dog tooth,” a large suborder of therapsids that arose in Africa in the
late Permian, following the theriodonts in the line leading to mammals; the cyn-
odonts, one of the few synapsids that survived the end-Permian extinction, diversi-
fied globally in the Middle Triassic and probably gave rise to the chiniquodontids,
possible ancestors of mammals; the cynodonts were then displaced by mammals by
the end of the Jurassic; see also Charassognathus and Procynosuchus; late Permian
to Late Jurassic.
Cynognathus: a genus of large early Mesozoic cynodonts, about the size of a wolf;
South Africa and South America, Early Triassic.
Cyonosaurus: a genus of gorgonopsian therapsids of the late Permian; there is
recent but inconclusive evidence that it may have survived the end-Permian extinc-
tion; South Africa, late Permian.
Cypridea: a genus of freshwater podocopid ostracods, possibly related to the mod-
ern Cyprididae; Middle Jurassic to Cretaceous.
Cystoidea: a class of primitive echinoderms in the subphylum Blastozoa; attached
to hard substrates by a stalk and generally characterized by pentameral symmetry
and triangular pore openings; may have given rise to the crinoids (see also
Edrioasteroidea); Middle Ordovician to Devonian.
Cystoporata: “sac-like passages,” an order of marine bryozoans in the still-extant
class Stenolaemata; characterized by elongated zooecia on supporting blister-like
vesicular structures called cystopores; Ordovician to Permian.
D
Dactylioceras: a genus of planulate ammonoids in the order Ammonitida; only

3 inches wide at the most but very numerous; it was part of the radiation of ammo-
nitids that followed the extinction of most ceratitid ammonoids at the end of the
Triassic; occurring in the Pliensbachian, always in a stratum just above that of
Amaltheus, it is a significant index fossil; global distribution, Early Jurassic,
200–172 Ma.
Daeodon: a genus of artiodactyl mammals in the family Entelodontidae; mainly a
scavenger, it was bison-sized and resembled a giant warthog; the type genus, D.
shoshonensis, was the largest entelodont, 12 feet long, 6 feet high at the shoulder,
and with a skull 3 feet long; widespread across North America, Oligocene to middle
Miocene;
Dakosaurus: “biting lizard,” a genus of large marine crocodylomorphs in the clade
Thalattosuchia and family Metriorhynchidae. It was up to 16 feet long and a good
swimmer, with a finned tail and paddle-shaped limbs. The skull was up to 3 feet
long, with a jaw containing dozens of sharp, serrated teeth. Dakosaurus was mainly
marine, but whether it gave birth at sea or on land is unclear. North and South
America, Europe, and Russia; Late Jurassic to Early Cretaceous, 160–135 Ma.
Dalmanites: a genus of phacopid trilobites in the suborder Phacopina and family
Dalmanitidae; 2 to 3 inches long, Dalmanites is characterized by large and raised
schizochroal compound eyes and a significant tail spine; global distribution, Silurian
to Early Devonian.
Dapedium: a genus of actinopterygian fish in the order Semionotiformes and fam-
ily Dapediidae; a durophagous fish, it had peg-like teeth useful for crushing hard-
shelled invertebrates; it was up to 16 inches long, deep-bodied, protected by thick
dermal head bones and a covering of rhombic ganoid scales; North America and
Europe, Late Triassic to Late Cretaceous.

72 D
Daphoenodon: a genus of large carnivorous canids in the family Amphicyonidae;

widespread in North America, late Oligocene to Miocene.
Darwinius masillae: a contentious species of early cat-sized primates from about
48 Ma, about 2 feet long. A nearly complete skeleton was found in 1983 in the
Messel Shale Pit that has opposable thumbs and fingernails rather than claws; absent
are the toothcomb and grooming claws characteristic of more primitive animals. Its
authenticity and importance, however, are in dispute, partly because of collection
difficulties. Like most specimens from Messel, the specimen of Darwinius is part
and counterpart of a split bedding plane, covered with resin to preserve it, so that the
bones cannot be individually handled and carefully examined. Germany, Middle
Eocene.
Darwinopterus: a genus of pterosaurs in the family Wukongopteridae; dated at
161 Ma, it is notable for possessing characteristics of both large groupings of ptero-
saurs, the “rhamphorhynchoids” and their descendants the pterodactyloids; China,
Late Jurassic.
Daspletosaurus torosus: “horrible muscular lizard,” the only named representative
of a genus of large tyrannosaurs in the subfamily Tyrannosaurinae; closely related
to the later T. rex, Daspletosaurus grew up to 30 feet long; there is some evidence
that it lived in social groups; Alberta and Montana, Late Cretaceous, 77 to 74 Ma.
Dastilbe: a genus of gonorhynchiform fish in the family Chanidae; from 1 to
2 inches long, it has been found in large numbers in the Crato Formation of Brazil;
South America and Africa, Early Cretaceous, 125–110 Ma.
Dasyatidae: a family of rays in the order Myliobatiformes; commonly called
whiptail stingrays, they are characterized by flattened fin discs and a generally oval
outline, and also by venomous stingers at the base of the tail; they may have origi-
nated in the Late Jurassic and were widespread from the early Eocene to the late
Miocene; global distribution, Early Cretaceous to Recent.
Dasyuromorphia: the still-extant order of marsupial mammals that includes the
Dasyuridae and the extinct Thylacinidae; Australia and New Guinea, early Oligocene
to Recent.
Davidsonina septosa: a spire-bearing species of articulate brachiopods in the class
Rhynchonellata, order Spiriferinida, and family Davidsoninidae, significant as an
index fossil; Europe, Early Carboniferous.
Daxatina canadensis: a species of ceratitid ammonoids in the family
Trachyceratidae; its first appearance marks the beginning of the Late Triassic at
237 Ma; Italy.
Dayiidae: a family of brachiopods in the class Rhynchonellata, order Atrypida,
and superfamily Dayioidea; Europe, Asia, and North Africa; early Silurian to Early
Devonian.
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Deccan Traps: an area of large volcanic flows in India, mainly basalt, resulting
from an eruption near the end of the Cretaceous; it is one of the largest geological
features of Earth, covering an area about 200,000 square miles.
Decennatherium: a genus of artiodactyl mammals in the family Giraffidae, similar
to the later Sivatherium, q.v. Decennatherium has been known for several decades
from several specimens found in Spain and is now well-known from an almost com-
plete skeleton described in 2017. The recent specimen, named D. rex, was about
10 feet tall and 7 feet at the shoulder; it has two small ossicones behind its eyes and
two longer ones at the back of its skull. Europe, late Miocene, 9 Ma.
Declinognathodus nodiliferus: a species of conodonts whose first appearance
marks the beginning of the Late Carboniferous (Pennsylvanian) at 323.2 Ma; United
States.
Deinocheirus: a genus of primitive theropod dinosaurs in the family Deinocheiridae;
36 feet long, it was the largest ornithomimid; known mainly from its large forearms,
which were 8 feet long, and its hands tipped with three 10-inch claws; Mongolia,
Deinogalerix: “terrible shrew,” a genus of omnivorous hedgehogs in the family
Erinaceidae. Some species grew to the size of a wolf, an example of island gigan-
tism. Deinogalerix had a long, thin skull and hair, not the spines of modern hedge-
hogs. Restricted to the Gargano Peninsula of Italy, which was the location of several
islands during the Miocene; Miocene to Pliocene.
Deinonychus: a monospecific genus of saurischian dinosaurs in the family
Dromaeosauridae; about 11 feet long, with 3 claws on each manus and pes, D. antir-
rhopus was an efficient predator; like other dromaeosaurs, it had a sickle claw on
the second toe of each pes; North America, Early Cretaceous, 115 to 108 Ma.
Deinopoidea: a still-extant superfamily of orb-weaving spiders in the order
Araneae. They produce a web of cribellate silk that appears to be combed, and in
which insect legs become caught in the same way that the Velcro fastening fabric
works. Fossil specimens of their webs are sometimes found because of insects
trapped in them; the oldest known complete orb web with prey attached (a fly, a
mite, a beetle, and a wasp leg) was found in 2005 in a Lower Cretaceous deposit in
Spain. Inconclusive phylogenomic research has challenged the monophyly of the
Deinopoidea. See also Araneoidea. Global distribution, Middle Jurassic to Recent.
Deinosuchus: “terrible crocodile,” a genus of gigantic crocodilians that lived for
less than 10 million years, from about 81 to 73 Ma; about 30 feet long and weighing
up to 5 tons, Deinosuchus inhabited the shallow sea in the middle of North America,
where its diet included dinosaurs and turtles; a five-foot long skull found in Texas in
1997 bears teeth 5 to 6 inches long; Late Cretaceous.
Deinotheriidae: a family of late Cenozoic elephant-like proboscideans whose
most notable feature was the pair of long, down-curving tusks on the lower jaw;
similar to the modern elephant but probably not ancestral to it; their extinction
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coincided with the worldwide Quaternary extinctions of many large mammals;

Europe, Asia, and Africa; Miocene to late Pleistocene.
Deiphon: a genus of phacopid trilobites in the family Cheiruridae; characterized
by a set of numerous strong spines and by the warty surface of the glabella, which
is similar to that of Phacops rana; Silurian.
Deltadromeus: a genus of ceratosaurian theropod dinosaurs; agile and probably a
fast runner although it weighed up to 2 tons and grew to 26 feet long; its relationship
to the similar Bahariasaurus is still unclear; Kem Kem beds, between Morocco and
Algeria, Late Cretaceous.
Deltatheridium: a genus of basal metatherian mammals in the order Deltatheroida
and family Deltatheridiidae; 6 inches long, scansorial, and carnivorous, it has a
combination of marsupial and placental characteristics; Mongolia and possibly
Deltatheroida: an order of basal metatherian mammals, near the beginning of the
line of descent leading to the marsupials; they were scansorial omnivores and wide-
spread in the Cretaceous; see also Gurbanodelta kara; Central Asia and North
America, Early Cretaceous to late Paleocene.
Delthyridina: see Spiriferidina.
Deltoblastus: a genus of blastoid echinoderms in the order Spiraculata; character-
ized by a conical shape with depressed ambulacra; attached to the substrate, from 1
to 3 inches long; similar but not directly related to the better-known Pentremites, it
was one of the last of the Blastoidea, which disappeared at the end of the Permian.
Demospongiae: the largest and most diverse class of the phylum Porifera, the
sponges; ranging from encrusting forms to large genera up to 3 feet in diameter,
they comprise 80 percent of known modern sponges; their skeletons are composed
of siliceous spicules, spongin fibers, or both; see also Sphaeractinoidea; Cambrian
to Recent.
Dendrerpeton acadianum: an obscure species of temnospondyl amphibians, an
early tetrapod known only from disarticulated fossils; Nova Scotia, Early
Pennsylvanian.
Dendrograptus: a genus of irregularly branching graptolites in the order
Dendroidea; Europe, Asia, and North America; late Cambrian to Middle Devonian.
Dendroidea: an order of sessile branching graptolites that originated in the middle
or late Cambrian, the earliest order of the Graptolithina; probably ancestral to the
other major graptolite order, the Graptoloidea, which appeared in the Early
Ordovician; most genera of the Dendroidea were benthic, attached by holdfasts to
the substrate and often bushy; the order is exemplified by Dendrograptus,
Callograptus, and Dictyonema; early Cambrian to Early Carboniferous.
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Dendropithecus: a genus of early primates in the superfamily Hominoidea and

family Pliopithecidae; 2 feet tall, it may be ancestral to the gibbons; East Africa,
early to middle Miocene.
Denisovans: an extinct species of humans in the genus Homo. The Denisovans
were a relatively small group of hominins who lived in Southern Siberia between
110,000 and 50,000 years ago. They are related to Neanderthals, who are known to
have lived between this area and Western Europe up to 40,000 years ago, and they
may also have interbred with other hominins, such as Homo erectus. This culture
supplies more evidence that many types of slightly differentiated hominins co-
existed with H. erectus for long periods of time.
denticle: pointed, placoid scales of cartilaginous fish; also, serrations on the teeth
of other animals.
dentition: Since teeth fossilize easily and are the most common fossil specimens
of many animals, patterns of dentition are paleontologically important. The type of
tooth often tells a great deal about diet and lifestyle, and the number and types of
teeth are often used in classifying animals.
The manner of replacement of teeth is also sometimes a significant descriptive
characteristic. Monophyodonty refers to having only one lifetime set of teeth.
Diphyodonty refers to having two sets of teeth, one deciduous (“milk teeth”) and
one permanent, and is one of the characters that differentiate mammals from therap-
sids. Polyphyodonty is the condition of having an indefinite number of tooth
replacements. See also brachydont, bradyodont, bunodont, heterodont, hypso-
dont, lophodont, petalodont, sectorial tooth, and selenodont.
Deperetellidae: a family of perissodactyl mammals in the superfamily Tapiroidea,
ancestral to modern tapirs; Asia, middle to late Eocene.
derived character: a novel character that first appears in the most recent common
ancestor of a lineage and is passed on to its descendants.
dermal ossicles: bony plates attached to the skin or cuticle of many animals, some-
times constituting an armored covering; found in dinosaurs, reptiles, turtles, echino-
derms, and several xenarthran mammals, such as the ground sloth Paramylodon and
the glyptodonts.
Desmatochelys: a genus of cryptodiran marine turtles in the extinct family
Protostegidae; the older of its two species, D. padillai, is the oldest known sea turtle,
7 feet long and dated at around 120 Ma; South America, Early Cretaceous.
Desmatosuchus: a genus of aetosaurs in the family Stagonolepididae; about
16 feet long, the crocodile-like Desmatosuchus bore a series of striking spines on its
sides, the farthest forward as long as 18 inches and pointing backward; North
America, Late Triassic, 230–210 Ma.
Desmodonta: a suborder of bivalve burrowing mollusks; characterized by the
absence of lateral teeth and few if any cardinal teeth; Ordovician to Permian.
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Desmostylia: “bundle pillar,” an order of quadrupedal aquatic perissodactyl

u ngulates, the only known extinct order of marine mammals. About 6 feet long and
up to 450 pounds, they were very similar to proboscideans and modern manatees,
but the similarities may be only a result of convergence. Desmostylians are charac-
terized by forward-pointing incisors and tusks, and by a dentition of closely packed
molars resembling a bundle of pillars. Japan and North America, late Oligocene to
early Miocene.
Deuterostomia: a superphylum including the phyla Chordata and Echinodermata,
and the subphylum Hemichordata; the earliest deuterostomes are microscopic
organisms of the early to middle Cambrian, the earliest accepted being Saccorhytus,
dated at 540 Ma; see also Vetulicolia; cosmopolitan, early Cambrian to Recent.
Devonian: the fourth period of the Paleozoic Era, from 416 to 359 Ma, divided
into Early, Middle, and Late Epochs. Known as the age of fish, the Devonian saw a
great radiation of marine and freshwater fish, including the first sharks. The first
ammonoids developed from the bactritid nautiloids in the Early Devonian.
Evolutionarily, the period is most important for its terrestrial developments. Aquatic
fish like Eusthenopteron in the Middle Devonian gave rise to tetrapods like Tiktaalik
and Ventastega that began to move ashore toward the end of the period. The first
insects appeared at the same time as a rapid radiation of terrestrial plant life. At the
beginning of the period, the tallest plant was only about 3 feet high, but as the vas-
cular plants spread, trees and even forests appeared. The progymnosperms and other
ancestors of most living plant groups were present by the end of the period.
Devonian extinction: a series of closely spaced extinction pulses during the last
5 million years of the period (some argue for seven events over the final 25 million
years). The principal extinctions occurred at the beginning and end of the
Famennian Stage, respectively the Kellwasser and Hangenberg Events. Major
groups that disappeared included the placoderms and most agnathans. The coral
and stromatolite reef-builders were so severely affected that there was little reef-
building until the rise of the scleractinian corals in the Mesozoic. The trilobites
were especially affected by the Kellwasser Event, and only those of the order
Proetida survived into the Carboniferous. Some have pointed to global cooling and
widespread anoxia in the oceans as a cause of some of these extinctions, but the
evidence is inconclusive.
Diacodexis: a genus of small Eocene mammals in the family Dichobunidae;
Diacodexis is the oldest known artiodactyl mammal, although its feet still had five
toes, with elongated third and fourth digits; it was less than 2 feet long, but it had a
longer tail; North America, Europe, and Asia; early to middle Eocene.
Diadectomorpha: a group of bulky anthracosaurian amphibians closely related to
the early amniotes, although their mode of reproduction is still obscure; reptile-like
and among the largest early tetrapods, some herbivorous genera (such as Diadectes)
grew up to 10 feet long; North America, Europe, and Asia; Late Mississippian to
late Permian.
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Diadematoida: a still-extant order of euechinoid sea urchins; characterized by

very long and slender spines and ten buccal plates around the mouth; Late Jurassic
to Recent.
Diademopsis: a genus of primitive echinoids in the order Pedinoida and family
Pedinidae, a common echinoid in Europe during the Early Jurassic. Generally
1–2 inches wide, with tesselate plating; long, slender spines; and a lantern with
broad, grooved teeth. Probably ancestral to euechinoids. Europe and South America,
Late Triassic to Early Jurassic.
diagenesis: in geology, a term for the process of changes in a rock over time, such
as a sedimentary rock becoming metamorphosed. In paleontology, diagenesis refers
to the final taphonomic process undergone by a fossil, especially to the process of
mineralization, which varies widely according to the burial site (such as in sedi-
ment, amber, tar, hydrothermal vent, or volcanic material). Successive diagenetic
events may occur if the burial environment is disturbed.
Diania cactiformis: a wormlike animal dated at about 520 Ma, related to an extinct
group of armored lobopods; characterized by robust spiny limbs, possibly jointed;
it may be a forerunner of true arthropods; Chengjiang, China, early Cambrian.
Dianulitidae: a family of stenolaemate bryozoans in the extinct order Cystoporata;
North America, Europe, and Asia; Ordovician and Silurian.
Diapsida: “two arches,” the stem group of early amniotes that gave rise to snakes,
lizards, and crocodiles; identified by two temporal fenestrae behind each eye, except
that modern snakes and lizards have lost one or both of these openings; the diapsids
appeared late in the Pennsylvanian, after the first synapsids.
diatoms: a kind of glassy-shelled algae in the class Bacillariophyceae; generally
flat and round like two pie plates stuck together, they constitute an important area of
study in micropaleontology; they appeared in the fossil record in the early Jurassic
and exist globally in marine and freshwater bodies of water; see also Stephanodiscus.
Diatryma: see Gastornis.
Dibunophyllum: a genus of solitary rugose corals in the family Aulophyllidae;
North America and China, Late Devonian to Late Pennsylvanian.
Dicellograptus: a widespread and varied genus of graptoloids in the family
Dicranograptidae; part of the leptograptid fauna, an important graptolite index fossil
group in the Late Ordovician; global distribution, Middle to Late Ordovician.
Diceratherium: “two-horned beast,” a genus of large perissodactyl mammals in
the family Rhinocerotidae; it weighed about a ton and was characterized by a pair
of horns side by side on the nose, the first rhinocerotoid to develop horns; wide-
spread in North America, Europe, and Asia; Oligocene to Miocene.
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Dichobunidae: a family of primitive artiodactyl mammals, characterized by

b unodont upper teeth and typified by Diacodexis, q.v.; global distribution, Eocene
and Oligocene.
dichograptid fauna: an important fossil group of the Ordovician, including the
many-branched Dichograptus and other graptoloid index fossils; see Graptolithina.
Dichograptus octobrachiata: an unusual species of graptolites, living in a circular
colony 3–4 inches in diameter; its 8 equal-length stipes were slender, with very
small individual animals; North America, Europe, and Australia; Early Ordovician.
Dickinsonia: an enigmatic genus of flat-bodied fossils in the family Dickinsoniidae.
It is an iconic member of the Ediacaran biota and variously considered to belong to
the fungi, the coelenterates, or an otherwise unknown class. The body is a bilaterally
symmetrical ribbed oval, usually a few inches long, but the largest known was about
3 feet long and 2 feet wide. Although hundreds of specimens have been studied,
little is known about its mode of life. Its trace fossils, generally ferruginous impres-
sions on the underside of an overlying stratum, have been dated between 579 and
555 Ma. Australia and Eastern Europe, Ediacaran.
Dicksonia: a still-extant genus of pteridophyte tree ferns in the order Cyatheales
and family Dicksoniaceae; Australia and New Zealand, Jurassic to Recent.
Dicranograptus: “two-headed,” a genus of bilateral graptolites in the order
Graptoloidea and family Dicranograptidae; a planktic member of the leptograptid
fauna, it is characterized by convoluted thecae; cosmopolitan, Early to Late
Ordovician.
Dicranurus: “two-head-tail,” a genus of spiny trilobites in the order Odontopleurida
and family Odontopleuridae; a very spinose trilobite, characterized by two thick,
curling spines behind the head; similar to the spiny trilobites of the order Lichida;
North America, North Africa, and Australia; middle Silurian to Lower Devonian.
Dicroidium: a poorly known genus of woody seed ferns in the family
Corystospermaceae; its leaves have been found attached to branches and some
reports refer to it as a tree dozens of feet high, but its fossils are known mainly as
leaves of a wide variety of types; widespread in Gondwana throughout the Triassic.
Dictyoclostus: a genus of brachiopods in the extinct order Productida and family
Productidae; its concave and convex valves are characterized by a network of fine
lines, grooves, and spines; D. americanus is an index fossil in the Late Pennsylvanian;
global distribution, Carboniferous to Permian.
Dictyonellida: a Paleozoic order of articulate brachiopods in the class Chileata;
characterized by a fine network of rays on the surface of the shell; Ordovician to
Permian.
Dictyonema: “net thread,” an important genus of many-branched dendroid grapto-
lites; sessile, characterized by its net-like structure and many small thecae; late
Cambrian to Early Ordovician.
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Dictyospongia: a genus of hexactinellid reticulate sponges in the family

Dictyospongidae; characterized by a smooth reticulum and an inverted cone shape;
North America, Silurian to Carboniferous.
Dictyothyris: a genus of articulate brachiopods in the still-extant order Terebratulida
and superfamily Terebratuloidea; characterized by an erect beak and large pedicle
aperture; Middle Jurassic to Early Cretaceous.
Dicynodontia: an infraorder of herbivorous therapsids; short and strongly built,
with some genera such as Placerias growing two tusks, they appeared in the middle
of the Permian; along with the theriodonts, they were the sole synapsid survivors of
the end-Permian extinction; they then diversified globally in the Triassic, most dis-
appearing in the end-Triassic extinctions, but some possibly surviving in Australia
into the Cretaceous.
Didelphidae: “two wombs,” the opossums, a still-extant family of arboreal marsu-
pials; didelphids probably originated in North America in the Early Cretaceous;
they were abundant in the Late Cretaceous but became extinct there in the Miocene;
after migrating in the Eocene to Europe and spreading worldwide, they again
appeared in North America near the end of the Pliocene.
Didelphodon: a genus of marsupial mammals in the family Stagodontidae; the
type species D. vorax, although only as large as a small badger, is estimated to have
had a stronger bite than a hyena; North America, Late Cretaceous.
Didolodontidae: an extinct family of middle-sized condylarths in the superorder
Meridiungulata; about 2 feet long, they were scansorial insectivores; South America,
Paleocene to early Miocene.
Didymograptus: “double branch,” one of the larger graptolites, a graptoloid genus
identified by its two long branches; often referred to as the “tuning fork” graptolite;
Early to Middle Ordovician.
Dielasma: a widespread genus of articulate terebratulid brachiopods; it had a
rather smooth shell surface, with the pedicle valve much larger than the brachial
valve; Carboniferous and Permian.
Diictodon: one of the most successful synapsids of the Permian, a genus of her-
bivorous dicynodont therapsids in the family Pylaecephalidae; less than 2 feet long
and armed with a pair of tusks extending down from the upper jaw, Diictodon bur-
rowed helical tunnels as deep as 5 feet where articulated skeletons have been found;
known mainly from Southern Africa but also Asia, 260 to 254 Ma.
Dikelocephalus: a genus of large North American asaphid trilobites in the family
Dikelocephalidae; only slightly longer than wide, it was 16–20 inches long; late
Cambrian.
Dikika baby: found in 2000 and possibly the most complete example of
Australopithecus afarensis, a fossil of a baby of nursing age; it includes a complete
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torso, a face, and some leg bones; 3.3 Ma, Dikika region of Ethiopia, across the
Awash River from Hadar.
Dilong: a monospecific genus of small dinosaurs in the family Proceratosauridae;
in the Jehol biota, it is generally considered an early tyrannosauroid (perhaps the
oldest) although it seems to have had a feather-like body covering; from 5 to 6 feet
long, D. paradoxus is known from almost complete specimens from the Lower
Cretaceous Yixian Formation; China, 125 Ma.
Dilophosaurus: “two ridges,” a monospecific genus of theropod dinosaurs; rela-
tively small but an efficient predator. D. wetherilli was about 23 feet long and weighed
about 1100 pounds, with long teeth; its most distinctive feature is a pair of crests
along the front of its skull; Kayenta Formation, Arizona, Early Jurassic, 193 Ma.
Dimaitherium: a monospecific genus of early paenungulate mammals in the order
Hyracoidea; D. patnaiki, discovered in 2010, is the earliest-known hyracoid; Fayum,
Egypt, late Eocene.
Dimetrodon: a genus of large synapsids in the family Sphenacodontidae, charac-
terized by its very large spinal fin, which gives it the nickname “sailbacked.” Most
species of Dimetrodon ranged from 6 to 15 feet long, and it was an apex predator. It
is named for its unusual heterodont dentition, having several sizes of teeth in a large
jaw. The sail may have regulated body temperature, but recent researchers have
challenged that theory; other theories also have little support. North America and
Europe, early Permian.
Dimorphodon: a genus of rhamphorhynchoid pterosaurs. It had a wingspan of
almost 5 feet, a large head, and three large claws on its wings. Dimorphodon is
named for its heterodont dentition, having sharp, protruding anterior teeth and also
shorter and flatter teeth toward the back of the jaw. The type species is D. macronyx,
and a possible second species (D. weintraubi) was named in 1998. Europe and pos-
sibly Mexico, Early Jurassic.
Dimylidae: an extinct family of insectivorous mammals in the still-extant super-
family Erinaceoidea; 4–8 inches long, it is distantly related to the modern desman
and hedgehog; known only from the Miocene in Europe.
Dinaelurus: a genus of feliform carnivores in the family Nimravidae; known only
from a relatively broad skull and partial skeleton found in the John Day Fossil Beds;
North America, Oligocene to early Miocene.
Dinocephalia: “terrible head,” a clade of large therapsids of the Permian. Ranging
from dog-sized (Pampaphoneus biccai) to more than 15 feet long (Moschops), they
were the least advanced therapsids and were similar to their ancestors the pelyco-
saurs. Among the largest animals of the Permian, some were herbivorous and others
carnivorous. The dinocephalians are known only from the period 272 to 260 Ma,
becoming extinct with no known descendants. See also Titanophoneus potens.
South America, Europe, Asia, and South Africa; middle to late Permian.
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Dinocerata: an order of large, herbivorous ungulate mammals; the niche of the

Dinocerata as large browsing animals was gradually taken over in the late Eocene
by the brontotheres; the best-known dinoceratan is Uintatherium; Laurasia, late
Paleocene to late Eocene.
dinoflagellates: “whirling whips,” an enigmatic group of protists; second only to
diatoms as primary producers in the oceans, dinoflagellates are also present in
brackish and freshwater environments; on the basis of biomarkers, they are thought
to have appeared in the late Proterozoic or early Cambrian, but suggested specimens
from the middle Paleozoic have been rejected (see Arpylorus); known more fully
from the Triassic to Recent.
Dinornithidae: a family of moas, giant flightless ratite birds in the infraclass
Palaeognathae. The largest of the moas grew to about 12 feet high and weighed
more than 500 pounds; moas displayed a wide degree of sexual dimorphism, the
female being much larger than the male. Along with the other family of moas, the
Emeidae, they were extant in New Zealand from the Pleistocene until about two
centuries ago, when the last of the moas were killed off by the Maori inhabitants.
Dinosaur Cove: a location on the coast of Victoria in southeastern Australia that
has produced specimens of several dinosaurs, including the only Leaellynasaura
and the first Allosaurus of the Southern Hemisphere.
Dinosauria: a large and diverse group of animals that dominated terrestrial life in
many parts of the world for almost 160 million years. Several primitive dinosauri-
forms in the Middle and Late Triassic have been suggested as ancestors (see for
example Saltopus), but definitive dinosaurs appeared only in the Late Triassic. The
superorder Dinosauria is generally taken as including two orders, the “lizard-
hipped” Saurischia and the “bird-hipped” Ornithischia. Both probably evolved from
one of the thecodonts in the Late Triassic around 220 Ma and became widespread in
the Jurassic. A contentious large group of dinosaurs is the Theropoda, which
includes the coelurosaurs, the most likely ancestral group of birds. The theropods
have traditionally been classified as belonging to the Saurischia, but 2017 research
argues that they are a separate group that shares more traits with the Ornithischia;
this proposal joins them with the Ornithischia in the revived clade Ornithoscelida.
Ranging from rat-sized to whale-sized, various families radiated into almost all
the terrestrial ecological niches, and the larger dinosaurs are the largest terrestrial
animals known. None became completely marine animals, but most were probably
capable of swimming, as swim tracks in Spain and Utah have shown. It seems
probable that all dinosaurs were oviparous, often burying the eggs. Some genera
have left clear evidence of nest-building, and some seem to have cared for the
hatchlings. On this point, see for example Egg Mountain, Maiasaura, Oviraptor,
Titanosauria, and Troodon. The number of dinosaur species declined toward the
end of the Cretaceous, and none survived the end-Cretaceous extinction. Global
distribution, Triassic to Cretaceous.
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Dinosaur National Monument: a large Konzentrat-Lagerstätte on the border of

Colorado and Utah that contains several hundred important fossil sites from near the
end of the Jurassic; the fossils occur in strata of the Morrison Formation and consist
mainly of dinosaurs and other large vertebrates; see also Abydosaurus; United
States, Late Jurassic.
Dinosauromorpha: a clade of archosaurian reptiles that includes the clade
Dinosauria (including birds) and the Lagerpetidae; Middle Triassic, 245 Ma, to
Recent.
Dinosaur Provincial Park: a World Heritage Site in southern Alberta, Canada;
renowned for its fossil assemblage from the Campanian Stage of the Cretaceous, it
is the richest source of Late Cretaceous dinosaurs in the world.
Diogenornis: a basal genus of rheas in the family Rheidae; the earliest-known rat-
ite, they were about two-thirds the size of the modern rhea; South America,
Paleocene.
diphyodonty: see dentition.
Diplacanthus: a genus of small, edentulous acanthodian fish in the order
Climatiiformes and family Diplacanthidae; named for their two prominent and
deeply embedded dorsal spines; synonym: Rhadinacanthus; North America and
Europe, Middle to Late Devonian.
Diplichnites: the generic name for a type of trace fossil consisting of two parallel
rows of tracks; it differs from Protichnites only in the absence of a medial furrow
and could be made by the same animal.
Diplobathrida: an order of crinoids in the subclass Camerata, characterized by a
conical or dicyclic cup and two circlets of plates; Middle Ordovician to Late
Mississippian.
diploblastic: a term referring to animals whose blastula is composed of two germ
layers, the ectoderm and endoderm; the principal diploblastic animals are the
Cnidaria and Ctenophora.
Diplocaulus: a genus of large, flattened, newt-like amphibians in the extinct sub-
class Lepospondyli and order Nectridea; 3 feet long, it had the nectridean
boomerang-shaped skull; North America, early to late Permian.
Diplodocus: “double-beam,” a genus of large sauropod dinosaurs; the name refers
to the two rows of bones on the underside of the tail vertebrae, which helped to sup-
port an extremely long tail; one of the largest dinosaurs, Diplodocus reached lengths
of more than 90 feet; its head, however, was only 2 feet long; the largest species, D.
hallorum, was formerly called Seismosaurus; North America, Late Jurassic.
diplograptid fauna: one of the important graptolite faunas of the middle Paleozoic;
characterized by the diplograptid pattern of long, feather-like stipes; see
Graptolithina; global distribution, Middle Ordovician to early Silurian.
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Diplograptus: a genus of scandent biserial graptolites in the order Graptoloidea;

characterized by a cap-shaped float to which featherlike stipes containing large
assemblages of organisms were attached; cosmopolitan, Ordovician and Silurian.
diploid: double or twofold.
Diplomystus: “double whiskers,” a genus of primitive herring-like fish in the order
Clupeiformes; generally small but some genera 2 feet long; distantly related to mod-
ern herrings and sardines; well-known from the Green River Formation in Wyoming,
in association with Knightia; global distribution, Early Cretaceous to early Eocene.
Diplopoda: a still-extant class of myriapod insects that includes the millipedes;
generally detritivorous, they are named “double foot” because they typically have
two pairs of legs on each body segment; see also Archipolypoda and Arthropleura;
Silurian to Recent.
Diploporita: an extinct class of blastozoan echinoderms in the class Eocrinoidea,
characterized especially by paired pores in the thecal plates; similar to the
Rhombifera; Early Ordovician to Middle Devonian.
Diplotrypa: “double hole,” a genus of stenolaemate bryozoans in the order
Trepostomata; generally a dome-shaped colony up to 4 inches across, composed of
circular tubes inhabited by autozooids; named for these holes and the larger holes
bored in the surface by other organisms; global distribution, Early Ordovician to
middle Silurian.
Diplura: “twintail,” a still-extant order of eyeless arthropods superficially similar
to springtails (Collembola); characterized by two pairs of appendages, two antennae
and two cerci, in most genera all equally long; divided into three families:
Campodeidae, Japygidae, Parajapygidae, and Testajapygidae; also commonly called
telsontails; Carboniferous to Recent.
Dipneusti: see Dipnoi.
Dipnoi: a long-lived order of bony fish, one of the oldest still-extant groups of
animals; the Dipnoi flourished from the Early Devonian to the Triassic, declining
since then but still represented by the tropical freshwater lungfish.
Dipodidae: the jerboas, a still-extant family of primitive omnivorous rodents in the
infraorder Myomorpha; from 2 to 10 inches long, some with tails three times longer
than their body, they are notable for their bipedal jumping ability; North America,
Europe, Asia, and Africa; Eocene to Recent.
Diprotodon: a genus of marsupial mammals in the family Diprotodontidae; the
largest marsupial known, Diprotodon was 10 feet long and more than 6 feet high at
the shoulder, and weighed 3 tons; its clawed forefeet turned inward like those of its
descendant the wombat; it inhabited open and wooded grasslands across the entire
continent of Australia from the early Pleistocene to about 45,000 years ago; the date
of its extinction is contentious, however, with some reports that it may have occurred
as late as 28,000 years ago.
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Diprotodontia: a still-extant, very diverse order of marsupial mammals, generally

considered restricted to Meganesia; the order includes kangaroos and wallabies (see
Macropodidae), koalas (see Phascolarctidae), wombats (see Vombatidae), and
others; the largest were the extinct Diprotodontidae (see Diprotodon); Australia,
Tasmania, and New Guinea; Oligocene to Recent.
Diprotodontidae: a family of herbivorous Australian marsupials in the still-extant
order Diprotodontia; the family includes genera that can be characterized as marsu-
pial hippopotamuses and rhinoceroses; the last of the Diprotodontidae became
extinct about 11,000 years ago; Australia, late Oligocene to Pleistocene.
Diptera: “two wings,” a large order of insects considered the true flies, estimated
to contain more than a million species, about 125,000 of them named. Dipterans are
distinguished from other insects by having only one pair of wings, which are
attached to the first thoracic segment; the second pair evolved into balancing organs.
None have stings, but they generally have sucking mouth-parts and can bite. They
seem to have evolved much later than the earliest insects, appearing only in the
Triassic; early dipterans are numerous in the insect fauna of the Madygen Formation
of Kyrgyzstan. Global distribution, Middle Triassic to Recent.
Dipteronotus: a genus of ray-finned fish in the family Perleididae; 2–4 inches long,
the freshwater and marginally marine Dipteronotus was abundant in the Grès à
Voltzia and is also known from a few other occurrences in Europe; Middle to Late
Triassic.
Dipterus: a genus of bony sarcopterygian fish in the order Dipnoi; about 14 inches
long and having a lung, it also closely resembles the modern lungfish in other
respects; it is known only from strata of the uppermost Devonian; Europe and
Australia, Late Devonian.
Disasteridae: a family of euechinoids in the superorder Atelostomata; no lantern,
ovoid and some heart-shaped; Europe, Jurassic and Cretaceous.
Discinoidea: a small but long-lived superfamily of inarticulate brachiopods in the
order Acrotretida; characterized by a cap-shaped, discoidal brachial valve; Early
Ordovician to Recent.
Discoidiidae: a family of globular or conical euechinoids in the extinct order
Holectypoida; Jurassic to Cretaceous.
Discosauriscus: “little-disk lizard,” a genus of aquatic stegocephalian seymouria-
morph stem-tetrapods in the family Discosauriscidae; about 16 inches long, flat-
tened, with wide jaws, and similar to its larger cousin Seymouria; characterized by
a small discoidal structure in the vertebrae; known definitively only from several
hundred juvenile specimens found in the lacustrine deposits of the Boskovice
Furrow site in the Czech Republic, where the genus constituted a near monoculture;
restricted to early Permian, 299–290 Ma.
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Discoserra: a genus of actinopterygian fish in the small order Guildayichthyiformes,

proposed as a possible stem neopterygian; its circular body is more than 12 inches
in diameter and is characterized by rhombic ganoid scales and small forward-facing
hooks on the dorsal scute; North America, Early Carboniferous.
disjunct endemism: the ecological occurrence of a species in a unique geographic
range and nowhere else in the world.
Distacodontidae: a family of primitive conodonts characterized by a fang-like
structure resulting from separation of the lamellae of the teeth; formerly called
Distacodidae; Ordovician to Silurian.
distal: see proximal.
Ditomopyge: a genus of small trilobites in the family Phillipsiidae; only an inch
long, it had strong genal spines and a broad and distinctively furrowed glabella; one
of the last of the trilobites; Late Carboniferous to the end of the Permian.
division: a level of classification used in botanical taxonomy, the equivalent of an
animalian phylum.
Docodonta: the shortest-lived and least diverse order of mammals in the Mesozoic.
Slightly larger than mice, in the Middle and Late Jurassic they radiated mainly in
Laurasia but almost globally. Their rectangular, multi-cusped teeth were well-
adapted for crushing and grinding; some genera may have been aquatic. Middle
Jurassic to Early Cretaceous.
Docofossor brachydactylus: a species of early mammaliaforms in the order
Docodonta; found in the Tiaojishan Formation and dated at about 160 Ma, the
6-inch-long Docofossor is the earliest-known burrowing mammal; its species name
refers to its having a reduced number of phalanges and short but wide fingers
adapted to digging; China, Late Jurassic.
dodo: see Raphinae.
Doedicurus: “pestle tail, a genus of xenarthran mammals in the subfamily
Glyptodontinae; the largest glyptodont at 13 feet long and 5 feet high, Doedicurus
is known only from southern South America, becoming extinct about 11,000 years
ago; late Pleistocene to Holocene.
Dokophyllum: a genus of solitary rugose corals in the family Ketophyllidae; a rhi-
zosessile horn coral, Dokophyllum produced root-like holdfasts that anchored it to
the sediment; North America, Europe, and Asia; middle to late Silurian.
Dolichothoraci: a group of primitive arctolepid joint-necked fish (arthrodires),
bottom fish characterized by long pectoral spines; see also Brachythoraci.
Doliocaris ingens: an enigmatic species of small arthropods, tentatively placed in
the proposed group Thylacocephala; it was probably 5 to 8 inches long and is espe-
cially notable for its very large and complex eyes, each of which was about one-
fourth of the body length; La Voulte-sur-Rhône, France, Middle Jurassic.
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Doliodus problematicus: a species of bottom-dwelling shark; a specimen found in

New Brunswick and dated at 409 Ma is the oldest intact shark fossil, earlier sharks
being known mainly from their teeth; similar to the modern angel shark, but notable
for a pair of inch-long bony spines on the front edge of the fins behind the head,
reminiscent of the acanthodian fish of the Silurian; Canada, Early Devonian.
Domeykosaurus chilensis: see Atacamatitan.
Dorudon: a genus of archaeocete whales in the family Basilosauridae; like other
basilosaurids, it was notable for a blowhole halfway to its eyes, an evolutionary step
toward the crown cetacean blowhole on the top of the head; it was completely
aquatic and had a rudimentary tail that was almost a caudal fluke; Egypt, late
Eocene.
Dorygnathus: a genus of small, basal pterosaurs in the family Rhamphorhynchidae;
it had heterodont teeth of different types and a wingspan of about 4 feet; known only
from several sites in France and Germany, especially from complete skeletons of
Holzmaden; Europe, Early Jurassic.
Dorypterus: “spear fin,” a genus of advanced palaeoniscid fish in the family
Dorypteridae; 8 inches long and characterized by a high dorsal fin almost as long as
its body; late Permian.
Doushantuo Formation: an important Ediacaran Konservat-Lagerstätte in
Guizhou Province, Southern China; covering the period 600 to 550 Ma, the forma-
tion is notable for its exceptional preservation of Ediacaran phosphatic fossils and
earlier Cryogenian acritarch fossils.
Dracorex: a proposed genus of ornithischian dinosaurs in the family
Pachycephalosauridae; known from only one possibly juvenile specimen, a skull
and several vertebrae; its flat, heavy skull and snout are covered with small spikes,
and some have argued that these spikes are simply juvenile forms of a
Pachycephalosaurus skull; South Dakota, Late Cretaceous, 71–66 Ma.
Dreadnoughtus schrani: a species of titanosaurian sauropod dinosaurs known
from two partially articulated specimens found in Patagonia in 2005 and described
in 2014; up to 85 feet long and 30 feet tall, it may be the largest land animal known,
but its weight is disputed, variously estimated at between 40 and 60 tons; Argentina,
Drepanaspis: “sickle-shield,” a genus of heterostracan fish in the family
Psammosteidae; with wide-spaced eyes and a row of spines all along its spine from
neck to tail, it was named for the sickle-shaped anterior outline of its flattened,
semicircular head; well-known from the Hunsrück Slate of Germany, Early
Devonian, 410 Ma.
Drepanellidae: a family of possibly paleocopid ostracods in the suborder
Beyrichicopina; one of several index fossils in the family is Drepanella crassinoda
of the Ordovician; late Cambrian to Late Ordovician.
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Drepanosauridae: a family of unusual reptiles in the order Protorosauria; up to

2 feet long, drepanosaurs are characterized by a sickle-shaped structure at the tip of
a prehensile tail, elongated limbs adapted to climbing, and a dorsal neural hump;
although generally found in moist environments, specimens were discovered in
2013 in a site in Utah that in the Triassic was a desert oasis; North America, Late
Triassic, 225–215 Ma.
Dromaeosauridae: “running lizard,” a family of feathered, carnivorous ornithis-
chian dinosaurs of small to medium size but well-provided with weapons, including
a long sickle claw on the hind feet that may have been used to disembowel or to
simply grasp prey; global distribution, Middle Jurassic to Late Cretaceous.
Dromaeosaurus: a genus of theropod ornithischian dinosaurs in the family
Dromaeosauridae; only 7 feet long and weighing just over 30 pounds, it was a small
carnivore; known only from fragments, it has been reconstructed in museum casts
from what is known of other dromaeosaurids; North America, Late Cretaceous,
77–75 Ma.
Dromiceiomimus: a possible genus of bird-like dinosaurs of the Late Cretaceous,
generally considered synonymous with Ornithomimus, q.v.
Dromomeron: a genus of dinosauromorph archosaurs in the family Lagerpetidae;
known from several fragmentary specimens from New Mexico and Argentina, it
was lightly built, about 3 feet long and undoubtedly a fast runner; North and South
America, Late Triassic, 220–210 Ma.
Dryolestidae: a family of small pantothere eutherian mammals; known in North
America and Europe, Middle Jurassic to Late Cretaceous.
Dryopithecus: “oak-ape,” a genus of monkey-like primates in the family Hominidae;
about 4 feet long, it closely resembled modern orangutans in its long arms, short
legs, and curved fingers; Asia, Africa, and Europe; late Miocene, 12 Ma.
Dryptosaurus: a famous genus of primitive tyrannosaurid dinosaurs; known defin-
itively from a partial skeleton of one individual, it is estimated to have been as much
as 25 feet long; named “tearing lizard” by E.D. Cope because of the 8-inch claws on
at least one of the three fingers on its forelimbs; the only known large carnivorous
dinosaur of Eastern North America; Late Cretaceous, 67 Ma.
Dunbarella: a thin-shelled pectinoid bivalve characterized by numerous fine rays
on the shell exterior; only 1.5 inches wide, it was similar to the modern scallop; its
fossils sometimes occur in association with ammonoids, which undoubtedly preyed
on it; North and South America, Europe, and Asia; Carboniferous.
Dunbaria: a genus of pterygote insects in the order Palaeodictyoptera; it had suck-
ing mouth-parts and dragonfly-like wings with a wingspan of 1.25 inches; a speci-
men from Kansas shows veins and color bands in the wings; North America and
Russia, early to middle Permian.
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Dunkleosteus: a genus of very large arthrodire placoderms in the order

Brachythoraci, one of many armored fish that became extinct at the end of the
Devonian Period. Originally named Dinichthys terrelli, Dunkleosteus grew at least
20 feet long and weighed more than a ton, with some estimates much higher. Its
posterior skeleton has not been preserved, probably because it was cartilaginous.
Armed with very powerful jaws and bony plates instead of teeth, it preyed upon
many marine animals. Cosmopolitan, Late Devonian.
durophagy: see Mesozoic Marine Revolution and Globidens.
Dusignathus seftoni deméré: a subspecies of Pleistocene walrus intermediate
between its terrestrial ancestors and later walruses; characterized by a large bone
crest on its head and short, tusk-like canine teeth; one of 24 species of walruses that
evolved from bear-like land animals since the late Oligocene, with Odobenus being
the only surviving species; see also Valenictus; Southern California, 3 Ma.
Dyticonastis: a monospecific genus of amphisbaenian reptiles; a burrowing limb-
less lizard, D. rensbergeri is known only from sites in central Oregon and is the only
known amphisbaenian west of the Rocky Mountains; United States, late Oligocene
to early Miocene.
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Echinerpeton: a poorly known monospecific genus of early synapsids; E.

intermedium is known only from fragmentary remains found in Nova Scotia and its
relationship to other synapsids is unclear; nevertheless, it may be the oldest known
synapsid, dated between 311 and 308 Ma; one specimen may have borne neural
spines; see also Archaeothyris; Canada, Late Pennsylvanian.
Echinocorys: a genus of irregular burrowing euechinoids in the family Holasteridae;
subconical with a flattened base; North America, Europe, and Asia; Late Cretaceous
to late Paleocene.
Echinocystitidae: a family of early echinoids in the subclass Perischoechinoidea;
characterized by four columns of plates in each ambulacrum; Europe, late Silurian
to Devonian.
Echinodermata: “spine skin,” a phylum of marine, mostly benthic animals known
definitively from the late Precambrian to the present, with about 7,000 extant and
13,000 extinct named species; the largest phylum with no terrestrial or freshwater
representatives; characterized by five-point radial symmetry and an internal skele-
ton of calcareous plates; includes starfish, sea urchins, sand dollars, sea cucumbers,
and sea lilies.
Echinoidea: the sea urchins and sand-dollars, a class of echinoderms in the sub-
phylum Echinozoa. Echinoids are characterized by a generally subspherical or
spherical shape, moveable appendages, and a skeleton of interlocking calcareous
plates. Most have a feeding apparatus called the lantern, which is composed of five
pyramidal plates, each with a single calcitic tooth; those without the feeding lantern
are referred to as atelostomate. Echinoids typically have an anal opening (periproct)
in the upper surface; in regular genera the periproct is in the center, while in irregu-
lar genera it is located away from the center. Global distribution, Middle Ordovician
to Recent.

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Echinoneoida: an order of irregular echinoids established in 2010 to include the

Conulidea, Echinoneidae, Cluniaster, and Mattsechinus; North and South America,
Europe, and Asia; Early Cretaceous to Pliocene.
Echioceras: a genus of evolute ammonoids in the order Ammonitida; generally
2 inches wide and strongly ribbed; E. raricostatum is an index fossil in the Early
Jurassic; North America, Europe, and Asia; Jurassic.
echolocation: the ability to locate objects by biological sonar; notable terrestrially
in bats and used to a lesser extent in some birds and small mammals; in marine
environments, it was first developed in the Eocene by odontocete whales and became
widespread in dolphins and porpoises of the Miocene.
Ecphora: a large genus of predatory snails in the family Muricidae; the name
“bearing out” is an architectural term going back to Vitruvius and refers to the dis-
tinctive T-shaped ribs protruding from the spiral shell; Eastern North America, late
Eocene to Pliocene.
Edaphosaurus: “pavement reptile,” a genus of pelycosaurian synapsid reptiles in
the order Pelycosauria and family Edaphosauridae. Perhaps the first large terrestrial
herbivore, it is named for thick clusters of teeth in both the lower and the upper jaws.
It is also characterized by the showy sail that more than doubled its skin surface,
extending upward along its backbone on almost two dozen long, curving neural
spines more than two feet long. Edaphosaurus was smaller than some of its spena-
codontid cousins, such as the even more spectacular Dimetrodon. North America
and Europe, late Pennsylvanian to early Permian.
edentates: see Xenarthra.
edentulous: lacking teeth.
Ediacaran: the final period of the Precambrian Eon and Neoproterozoic Era, 635–
541 Ma; formerly called Vendian.
Ediacaran biota: a general term for late Precambrian assemblages of primitive
organisms first found in the Ediacara Hills of Australia and later worldwide. Very
difficult to identify and classify, they are mostly impressions in overlying strata. The
Ediacarans include the first true multicellular animals: jellyfish, soft corals, worms,
and other animals that have not been identified. Many seem likely to be evolutionary
dead-ends that are unrelated to later taxa, but some of the world’s current animal
body plans originated in the Ediacaran. Almost all of the other current body plans
originated in the great radiation of species at the beginning of the Cambrian.
Edmontosaurus: a genus of late Maastrichtian hadrosaurs, duck-bill dinosaurs in
the subfamily Saurolophinae; synonym: Anatosaurus. With a short neck but almost
40 feet long, some were among the largest hadrosaurids. Well-preserved skeletons
of one species, E. annectens, show skeletal articulation and skin impressions. One
was originally named Trachodon annectens and referred to as the “Trachodon
mummy” because of its skin impressions and complete skeleton; and another called
E 91
“Dakota,” found in 1999, shows possible stripes on its fossilized skin. Although
widespread across Western North America, Edmontosaurus was first discovered in
the Horseshoe Canyon Formation of Southern Alberta. North America, Late
Edrioasteroidea: a long-lived class of stemless encrusting echinozoan echino-
derms in the paraphyletic subphylum Pelmatozoa; they may have given rise to the
crinoids in the Ordovician. The edrioasteroids are characterized by curved and
sometimes spiral ambulacra. Although never numerous, they survived from the
early Cambrian (possibly the Ediacaran) to the Permian. Global distribution.
Effigia: a famous example of convergent evolution, a genus of crocodylomorph
archosaurs in the family Shuvosauridae. It was 6–8 feet long, edentulous but omniv-
orous, and primarily bipedal, closely resembling the theropod ornithomimid dino-
saurs of the Early Cretaceous. But close examination showed that, like its close
relative Shuvosaurus, it was more closely related to crocodilians. New Mexico, Late
Egg Mountain: a well-known site of a purportedly colonial dinosaur nesting
ground in the Two Medicine Formation of Western Montana, where the first dino-
saur eggs in North America were found. The site is notable especially for the
Maiasaura nests with numerous eggs and possible indications of dinosaur parenting
behavior. There are also specimens of Orodromeus and Troodon, as well as Troodon
eggs. Late Cretaceous, 84–75 Ma.
Eileanchelys: a genus of early turtles known only from partial specimens found in
Scotland; about a foot long and low-domed, it is similar to the Early Jurassic
Kayentachelys; Middle Jurassic, 164 Ma.
Einiosaurus: a medium-sized horned and plated herbivorous ceratopsian dinosaur,
known only from Montana; about 20 feet long and weighing about 2 tons,
Einiosaurus looked like a small Triceratops except that it had a downward-curving
horn extending forward from its nose; United States, Late Cretaceous, 75–73 Ma.
Elasmobranchii: “plated gills,” a subclass of chondrichthyan fish that includes the
Batoidea (skates, rays, and sawfish) and the Selachii (sharks); characterized by
small scales on the skin surface and by 5 to 7 pairs of gills with exposed slits; middle
Silurian to Recent.
Elasmosauridae: a later, highly specialized group of plesiosaurs, notable espe-
cially for a very long neck; the type species Elasmosaurus at 46 feet was the longest
plesiosaur; global distribution, Early to Late Cretaceous.
Elephantidae the elephants and mammoths, a still-extant family of large probos-
cidean mammals that split off from mastodons and the other proboscideans in the
Pliocene and spread throughout the Northern Hemisphere; represented today by two
monospecific genera, which are slightly smaller than some of the Pleistocene gen-
era; see also mammoths and Mammutoidea; North America, Europe, Asia, and
Africa; late Eocene to Recent.
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Elginerpeton: a monospecific genus of early tetrapods, intermediate between fish

and amphibians; E. pancheni was found in Upper Frasnian strata and dated at
375 Ma, 10 million years earlier than Ichthyostega; known from humerus, tibia, and
jaw bones, it was about 5 feet long; Scotland, Late Devonian.
Ellesmeroceras: a genus of small nautiloid cephalopods in the order
Ellesmerocerida; the shell is usually orthoconic but may also be curved; North
America and Asia, late Cambrian to Early Ordovician.
Elliot Formation: a rock formation in South Africa that has produced many sau-
ropodomorph specimens, such as Massospondylus and Melanorosaurus; Late
Triassic to Early Jurassic.
El Montsec Lagerstätte: a site in Catalonia that has produced many fossils from
the beginning of the Cretaceous, about 130 Ma, slightly earlier than the better-
known Las Hoyas site of Eastern Spain. El Montsec is known for the only specimen
of Noguerornis, several important specimens of early orb-weaving spiders, and
many beetles and other insects. Spain, Early Cretaceous.
Elrathia kingi: a genus of medium-sized trilobites in the order Ptychopariida; one
of the most common trilobites of North America, commercially exploited especially
from the Wheeler Shale Formation of Utah; North America, Europe, and Asia; mid-
dle Cambrian.
Embolomeri: a contentious suborder of large, primitive fish-eating anthracosau-
rian amphibians. Although they have some reptilian features, they are probably not
ancestral to reptiles. The embolomeres evolved from reptiliomorph amphibians in
the Early Carboniferous, developing vertebral adaptations that facilitated undula-
tory swimming. They survived the end-Permian extinction and may have partially
adapted to terrestrial life before becoming extinct. North America and Europe,
Carboniferous to Early Triassic.
Embrithopoda: “heavy-footed,” an extinct order of large subungulate eutherian
mammals, represented mainly by genera of the Paleocene and Eocene but known
best by the latest embrithopod, Arsinoitherium, q.v.; despite their resemblance to
the rhinoceros, the Embrithopoda are more closely related to elephants; Africa,
Asia, and Europe; late Paleocene to early Oligocene.
Emeidae: a family of moas; see Dinornithidae.
Emuellidae: a family of early trilobites in the order Redlichiida; characterized
generally by an elongated body with numerous thoracic segments, and by long
genal and pleural spines; Australia and Antarctica, early Cambrian, 520–516 Ma.
Enaliarctos: a genus of early pinnipeds in the superfamily Otarioidea, probably
ancestral to the desmatophocids; it had a short tail and retained the carnassial teeth
of its terrestrial ancestors (possibly related to the ursids) but had flipper-like limbs;
North America, late Oligocene to early Miocene.
E 93
Enaliornis: “sea bird,” a genus of hesperornithine birds in the small family

Enaliornithidae; the type genus Enaliornis is the oldest-known hesperornithine,
dated between 100 and 94 Ma, near the beginning of the Late Cretaceous (Albian);
about the size of a large pigeon, Enaliornis was a foot-propelled diver and probably
nested near water; it is known from three partial skulls and many bone fragments;
originally named Pelagornis but not closely related to that bird of the Miocene;
England, Late Cretaceous.
Enantiornithiformes: an order of birds in the subclass Enantiornithes, known
only from the Cretaceous. These “opposite birds” seem to have originated in the
Early Cretaceous and to have achieved global distribution toward the end of the
period; they may have been the predominant type of bird during the Cretaceous, but
they were all extinguished in the crisis that marked the end of the Mesozoic era. The
enantiornithines were more advanced than Jurassic birds in having enlarged breast
bones, shortened tails, and a pygostyle; but they were less similar to modern birds
than later genera such as Ichthyornis and Hesperornis. The sparrow-sized
Iberomesornis (Spain, Early Cretaceous, 125 Ma) had a pygostyle but with a claw
on each wing was one of the most primitive of the group. The type genus Enantiornis
(Argentina, Late Cretaceous) was more advanced and seems to have been a good
flyer. See also Eoenantiornis and amber. South America, Spain, and China; Early
to Late Cretaceous.
Enchodus: a genus of salmoniform neopterygian fish in the family Enchodontidae;
up to 5 feet long and weighing 25 pounds, E. petrosus was a predatory fish with
fangs more than 2 inches long; seven other species were smaller; global distribution,
Late Cretaceous to early Eocene.
Encrinus: an important genus of pelmatozoan inadunate crinoids in the order
Encrinida; encrinites are characterized by their unopened tulip shape, with a large
cup from 1 to 2 inches long, at the top of a long stalk covered with ossicles; some
Triassic formations are dominated by accumulations of encrinite stalks; Europe,
Asia, New Zealand, and North America; Early to Middle Triassic.
Encrinurus: a widespread genus of trilobites in the order Phacopida and family
Encrinuridae; characterized by the large tubercles on its headshield which give it the
appellation of “strawberry-headed”; global distribution, Middle Ordovician to Early
Devonian.
Endoceratidae: a family of large, straight-shelled nautiloid cephalopods in the
order Endocerida; the family includes possibly the largest known Paleozoic fossils,
specimens of the giant Cameroceras reaching a length of 20–25 feet; global distri-
bution, Early Ordovician to Early Devonian.
Endothyroidea: an extinct superfamily of large benthic foraminiferans in the
order Fusulinida; index fossils in the Late Devonian and Early Carboniferous;
global distribution, Devonian to Late Triassic.
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Ensifera: the crickets and katydids, a suborder of stridulatory chewing insects in

the order Orthoptera, among the earliest flying insects; global distribution, Late
Enteletoidea: a superfamily of punctate articulate brachiopods in the order
Orthida; global distribution, Middle Ordovician to Early Triassic.
Entelodontidae: a family of large, primitive artiodactyl mammals in the super-
family Entelodontoidea; although long assigned to the Suina, they are now consid-
ered closer to whales and hippopotamuses than to pigs; they are characterized by
bony flanges or bumps on each side of the jaw, so that they superficially resemble
warthogs; North America, Europe, and Asia; middle Eocene to early Miocene.
entognathous: of insects, having the mouth parts retracted below the surface of the
head; a group of proto-insects in the subphylum Hexapoda are sometimes referred
to as the Entognatha.
Entomoconchacea: a superfamily of large ostracods in the suborder Myodocopa;
characterized by the absence of a rostrum above the permanent aperture; Europe,
Eoabelisaurus: a genus of ceratosaurian theropod dinosaurs in the family
Abelisauridae; described in 2012 and dated at about 170 Ma, it is now the earliest
(by 40 million years) and most complete skeleton of an abelisaurid; Patagonia,
Middle Jurassic.
Eoalulavis: a monospecific enantiornithine genus, the only species being
Eoalulavis hoyasi; found at Las Hoyas, it is the earliest-known bird with an alula, a
tuft of small feathers attached to the base of the thumb, improving maneuverability
in flight; Spain, Early Cretaceous, 125 Ma.
Eoandromeda: an Ediacaran genus of possible animals of an uncertain nature; cir-
cular and up to 1.5 inches wide, they have 8 spiral arms that seem to be tubular;
known only from slightly different specimens from Australia and China, they may
be stem-ctenophores, but widely different interpretations have been advanced;
580–550 Ma.
Eobasileus: a genus of dinoceratan mammals in the family Uintatheriidae; the
largest of the uintatheres, it was the size of a modern rhinoceros at 4 tons, 13 feet
long, and 7 feet high at the shoulder; armed with two short tusks and three pairs of
blunt horns, Eobasileus closely resembled its cousin, the more widespread
Uintatherium; North America, middle to late Eocene.
Eocarcharia dinops: a species of large dinosaurs in the family
Carcharodontosauridae; characterized by a massive brow ridge above the eyes,
hence its species name, “fierce-eyed”; may have been as long as 20–26 feet; Niger,
Early Cretaceous, 112 Ma.
Eocene: the second epoch of the Cenozoic Era, from 56 to 34 Ma. The earliest
orders of modern mammals were present by the end of the epoch, and modern birds
E 95
diversified throughout. At the end of the Eocene, modern grassland ecosystems had
spread, along with modern moths and butterflies as well as the larger grazing mam-
mals. Lower world temperatures occurred toward the end of the period, introducing
minor glaciations.
Eocrinoidea: a paraphyletic class of pelmatozoan echinoderms in the subphylum
Blastozoa, ancestral to several other classes; the earliest-known stalked, arm-bearing
echinoderms; Europe, North America, and China; early Cambrian to late Silurian.
Eodalmanitina: a genus of phacopid trilobites in the family Dalmanitidae; from 1
to 2 inches long, it is notable for its raised, crescent-shaped, schizochroal eyes; it
had more than a dozen thoracic segments and the small pygidium ended in a short
spine; Europe, Middle Ordovician, 461–458 Ma.
Eoderoceratidae: a family of ammonoids in the order Ammonitida; characterized
by an evolute shell and small spines; South and North America, Europe, and Africa;
Early Jurassic, Sinemurian to Pliensbachian.
Eodiscina: a suborder of trilobites in the order Agnostida; they are isopygous and
very small, less than an inch long; early Eodiscina genera are notable for their
abathochroal eyes, but later genera are eyeless; global distribution, Cambrian to
Silurian.
Eodiscoidea: a superfamily of trilobites generally considered part of the order
Agnostida; see also Pagetia; global distribution, early to late Cambrian.
Eodromaeus: “dawn runner,” a basal genus of theropod saurischian dinosaurs
about 4 feet long but weighing only about 11 pounds; a possible common ancestor
of all other dinosaurs; Argentina, Late Triassic, 230 Ma.
Eoenantiornis: a monospecific genus of enantiornithine birds known from one
specimen found in the Yixian Formation; dated at the beginning of the Aptian Stage,
125 Ma, E. buhleri was about 5 inches long; it is one of the most primitive and old-
est of the known enantiornithine birds; Jehol biota, China, Early Cretaceous.
Eognathacantha ercainella: a possible early species of chaetognaths, known only
from a whole-body specimen found in the Maotianshan Shales; China, early
Cambrian, 520 Ma.
Eohippus: a genus of perissodactyl ungulates in the family Equidae; formerly syn-
onymous with Hyracotherium, it is now considered a separate monospecific genus;
E. angustidens was the size of a medium-sized dog; North America, early Eocene.
Eolympia: See Ceriantharia.
Eomaia: an enigmatic, monospecific genus of early mammals; dated at 125 Ma
and known from one complete skeleton, E. scansoria was only about 6 inches long
nose to tail; although it had several features common in placental mammals, includ-
ing fur, it lacked several features common to all placentals; it also had an epipubic
bone, a marsupial character; Jehol biota, China, Early Cretaceous, 125 Ma.
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Eomanis: a genus of pangolins in the suborder Eupholidota; the earliest-known

pangolin from Europe, it was only 20 inches long and had scales only on its body;
Messel Shale Pit, Germany; early Eocene.
Eomoropidae: a primitive family of perissodactyl ungulate mammals in the sub-
order Ancylopoda, related and possibly ancestral to the chalicotheres; although they
bore claws on their feet, they were sheep-sized grazers or browsers; Asia and North
America, early Eocene.
Eomyidae: a family of rodents in the suborder Castorimorpha and superfamily
Geomyoidea; the family includes the earliest gliding squirrel (Eomys quercyi) and a
variety of small and middle-sized genera; North America, Europe, and Asia; middle
Eocene to Pleistocene.
Eophalangium: see Opiliones.
Eophrynus: a genus of spider-like arachnids in the order Trigonotarbida; charac-
terized by numerous dorsal knobs and half a dozen ventral ridges, unable to spin
silk; Europe, Late Carboniferous.
Eoraptor: a 1992 discovery in Argentina of one of the earliest dinosaurs, dated at
231.4 Ma. A primitive saurischian close to the ancestry of both sauropodomorhs and
theropods, Eoraptor was the size of a greyhound, about 3 feet long. It was probably
mainly carnivorous but more primitive than Herrerasaurus and other early thero-
pods in having no jaw-hinge, and its degree of bipedality is unclear. It had curved
and serrated teeth and three-fingered hands adapted to tearing flesh. Argentina, Late
Triassic.
Eosimias: a genus of extremely primitive anthropoid primates, significant because
it seems to represent a transitional figure between the higher and lower primates.
Eosimias is characterized especially by a deep chin and dagger-like canine teeth
similar to those of many modern monkeys. Smaller than expected for an intermedi-
ate form between the upper and lower primates, it is the size of a shrew; the largest
of the four species so far described weighed only 10 ounces. China, middle Eocene,
40–45 Ma.
Eosuchia: a polyphyletic group formerly considered an order of diapsid reptiles,
now discarded except as an informal term; once considered numerous, the eosuchi-
ans are now restricted to the small family Younginidae, q.v.
Eothele: see Palaeothele.
Eotitanops: a genus of brontotheres, perissodactyl grazing mammals; the earliest
accepted brontothere, Eotitanops was less than 2 feet high and resembled a small
modern tapir; it had 5 digits on the manus and 3 on the pedes; North America and
Asia, early to middle Eocene.
Eotitanosuchus: a genus of basal therapsids, placed tentatively in the suborder
Biarmosuchia although significantly different from other biarmosuchians; known
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from a single partial skull, it is estimated to have been about 9 feet long; Russia,
middle Permian.
Eotragus: a genus of early ruminants in the family Bovidae; the size of a gazelle,
about 40 pounds, and characterized by two vertical horns, Eotragus was a browser
in open and wooded grasslands; the earliest-known bovid, it evolved in Eurasia
around 20 Ma and appeared in Africa soon after; Asia, Europe, and Africa; early to
late Miocene.
Eozostrodon: a contentious genus of early cynodont mammals tentatively placed
in the family Morganucodontidae; known only from disarticulated fragments,
mainly teeth, it is considered by some to be synonymous with Morganucodon;
found in England near the boundary of the Triassic and Jurassic.
Ephemeroptera: the mayflies, an extant order of palaeopteran insects; they are
aquatic and are characterized by the primitive characters of long tails and wings that
cannot be folded over their abdomen; global distribution, Late Carboniferous to
Recent.
Epidendrosaurus: a genus of sparrow-sized saurischian dinosaurs in the suborder
Theropoda; notable for extreme elongation of the forelimb digits and other adapta-
tions to a tree-dwelling habit; synonym: Scansoriopteryx; China, Late Jurassic,
160 Ma.
epifaunal: referring to benthic organisms that live on the surface of a sea or lake
bed, either attached or free-moving, as opposed to infaunal organisms, which bur-
row into the sea bed or construct burrows or tubes.
Epigaulus: see Ceratogaulus.
epi-planktic: of marine organisms attached permanently to a floating substrate.
Epithyris: a genus of articulate terebratulid brachiopods in the family
Postepithyrididae; punctate with a short loop; Europe and Africa, Middle to Late
Jurassic.
Eporeodon: a genus of oreodonts in the family Merycoidodontidae and subfamily
Eporeodontinae; herbivorous artiodactyls with four digits and weighing 200–300
pounds, they are related to camels; not numerous and restricted to Western North
America, early Oligocene to early Miocene.
Equidae: the horses, a family of perissodactyl mammals in the suborder
Hippomorpha and the superfamily Equoidea. The palaeothere Hyracotherium, from
the middle Paleocene, had long been considered the first horse but is no longer con-
sidered ancestral to the Equidae. The earliest unquestioned Equidae are Eohippus
and Protorohippus, fox-sized animals of the early Eocene. Early horses increased in
size very slowly from the late Paleocene to the late Oligocene, but then diversified
rapidly in the early and middle Miocene. From the late Miocene to the Pleistocene,
their size continued to increase, with smaller species becoming extinct. The line of
descent of the later Equidae is well-documented, from Miohippus (Oligocene),
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Hypohippus (Miocene), Merychippus (Miocene), and Dinohippus (Miocene to

Pliocene) to the earliest-known species of the genus Equus, E. simplicidens (late
Pliocene to Pleistocene). Equus quickly migrated to Eurasia and Africa but became
extinct in North America about 12,000 years ago.
Equisetum: a still-extant genus of horsetails in the class Equisetopsida; they
appeared in the Devonian and became widespread in the Carboniferous; there are
about 25 extant species of Equisetum; global distribution.
Equisetopsida: a still-extant class of plants now considered a relative of the ferns
and assigned to the Pteridophyta; formerly called Sphenopsida or Sphenophyta;
global distribution, Late Devonian to Recent.
Eramosa Lagerstätte: a fossil locality in Ontario, Canada, that has produced
well-preserved autochthonous specimens of eurypterids and other arthropods, anne-
lid worms, conodonts, and fish from the middle to late Silurian.
Erbenochile: an unusual genus of phacopid trilobites in the family Acastidae;
notable for its two straight-sided eyes extending perpendicularly above the body;
each eye had up to 500 lenses, and the largest species had 360-degree vision; North
Africa, 403–393 Ma.
Eremotherium: a genus of large ground sloths in the subfamily Megatheriinae;
from 18 to 20 feet long and weighing 3 tons, it could sit at least partially erect;
almost toothless, it was unusually sexually dimorphic, males being almost 50 per-
cent larger than females; it appeared about 5 Ma and became extinct about
11,000 years ago; North and South America, Pleistocene.
Eritherium: a proposed genus of proboscideans described in 2009, considered a
basal proboscidean, much earlier than Moeritherium (late Eocene) and other ances-
tral proboscidean genera; known only from a partial skull, jaw, and teeth; Morocco,
Paleocene, 60 Ma.
Eryops: “drawn-out face,” a monospecific genus of semi-aquatic temnospondyl
amphibians in the family Eryopidae; perhaps the largest land animal of its time at
5–10 feet long and 200 pounds, Eryops is one of the best-known temnospondyls; its
muzzle is almost as long as that of an alligator, and its ribs are unusually short and
straight; North America, Late Pennsylvanian to early Permian.
Erythrosuchus: a genus of thecodont reptiles, close to the ancestry of the last com-
mon ancestor of the archosaurs; Erythrosuchus was 16 feet long and one of the
largest predators of the Middle Triassic, competing with therapsids in Southern
Africa in the period of time before they disappeared; restricted to Southern Africa,
250–230 Ma.
Essexella: see Scyphozoa.
Euarchonta: a proposed grouping of primates, tree shrews, colugos, and the
extinct Plesiadapiformes; early Paleocene to Recent.
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eucynodonts: the clade of cynodont therapsids that includes the ancestors of

ammals; global distribution, Early Triassic to Recent.
m
Euechinoidea: a subclass of echinoids that includes all post-Paleozoic echinoids;
global distribution, Late Triassic to Recent.
Eugeneodontida: an order of holocephalic fish; distinguished by a tooth-whorl of
sharp teeth in the lower jaw; poorly known although numerous; global distribution,
Early Carboniferous to Early Jurassic.
Euhoplites: “well-armored,” a genus of small ammonitid ammonoids; character-
ized by strong ribbing of the shell, which was covered with tubercles and other
bumps; North and South America and Europe, Early Cretaceous, 110–100 Ma.
Eukaryota: the domain of organisms whose complex cells have a distinct nucleus
surrounded by a membrane; they range from microscopic amoebae to the largest
plants and animals; although the oldest eukaryote fossils are about 1.7 billion years
old, genetic evidence points to an age of about 3 billion years; see also Archaea and
Bacteria.
Eulipotyphla: an order of small insectivorous mammals that became widespread
toward the end of the Cretaceous and is still represented by the shrews, hedgehogs,
and moles; see for example Zalambdalestidae and Asioryctitheria; global distri-
bution, Late Cretaceous to Recent.
Eumalacostraca: the subclass of crustaceans that includes most of the modern
crustaceans, such as crabs, lobsters, shrimp, crayfish, and sow bugs; global distribu-
tion, Cambrian to Recent.
Eunotosaurus: a genus of broad-bodied reptiles that may be an intermediate form
between turtles and their Paleozoic ancestors; known from relatively rare but well-
described specimens, they have a broad body with several flattened ribs and turtle-
like vertebrae; see also Pappochelys and Kayentachelys; South Africa, middle to
late Permian.
Euomphalus: a widespread genus of marine gastropods in the family Euomphalidae,
generally about 2 inches wide; the closely coiled shells are usually planispiral but
occasionally have a slightly raised spire; Silurian to middle Permian.
Eupantotheria: an order now considered obsolete; its members, generally small
therian mammals between the Middle Jurassic and the Early Cretaceous, probably
include the ancestors of the later marsupials and placentals; they are being reclassi-
fied into separate groups.
Euparkeria: a genus of primitive reptiles in the possibly monogeneric family
Euparkeriidae; known primarily from the Karoo Basin in South Africa, it is close to
the ancestry of the archosaurs; about 3 feet long but only 10 pounds, characterized
by recurved teeth, forelimbs slightly shorter than the hindlimbs, and a row of
osteoderms down the spine and tail; Africa, Early to Middle Triassic.
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Europasaurus: a genus of small sauropod dinosaurs in the family Brachiosauridae;

estimated at between 5 and 20 feet long, Europasaurus seems to represent an exam-
ple of island dwarfism; Germany, Late Jurassic.
Europejara olcadesorum: a species of pterodactyloid pterosaurs in the family
Tapejaridae; the oldest tapejarid known and the first found in Europe, it is a rela-
tively small species, with a wingspan of 7 feet; dated between 130 and 125 Ma, it
was discovered in 2012 in a Lower Cretaceous formation in Spain.
Euryalida: the basket stars, a still-extant order of asterozoan echinoderms in the
class Ophiuroidea; their numerous arms grow up to 2 feet long and can branch in
any direction, so that they often resemble a small, bushy plant; they have a poor fos-
sil record; global distribution, Carboniferous to Recent.
euryapsids: a polyphyletic group of early Mesozoic marine reptiles including the
ichthyosaurs; noted for a single temporal fenestra near the top of each side of the
skull, higher than in the synapsids.
euryhaline: tolerant of a wide range of salinity; bivalves are more euryhaline than
trilobites, which became extinct at the end of the Permian, and brachiopods, which
have declined since the Paleozoic; dinoflagellates are also generally euryhaline.
Eurymylidae: a family of basal rabbit-like mammals, possibly ancestral to all later
rodents; Mongolia, late Paleocene to middle Eocene.
Eurypterida: an extinct order of sea scorpions, arthropods in the class Chelicerata;
they had gills, and most were less than 12 inches long, but some were larger than a
man (see Jaekelopterus and Pentecopterus); they generally had more than one pair
of walking legs, the last of which was larger and used for swimming; they probably
gave rise to the Arachnida, but the horseshoe crab is their closest modern relative;
global distribution, Ordovician to late Permian.
Eurypterus remipes: “wide wing oar-foot,” the first eurypterid to be discovered
and the most common, a species of chelicerates in the family Eurypteridae; gener-
ally less than a foot long, but individuals as long as 4 feet are known; E. remipes had
the spiny chelicerae and walking legs, distinctive broad paddles, and spiked telson
that are characteristic of the order; North America and Europe, late Silurian.
Eusthenopteron: a genus of large rhipidistian fish, possibly stem-tetrapods; differ-
ences in its teeth, spine, skull, and especially the bones of its swimming fins may
have led to the lobe-finned sarcopterygians and the labyrinthodont amphibians such
as Ichthyostega; see also Acanthostega and Tetrapoda; Canada, Middle Devonian.
Euthacanthus: a genus of edentulous acanthodian fish in the family Climatiidae;
England and Scotland, Early Devonian.
Eutheria: the subclass of mammals that includes all living and extinct placental
mammals. The earliest eutherians arose in the Early Cretaceous or perhaps Late
Jurassic, and diversified slowly until the beginning of the Paleocene, when they
began a very rapid radiation of species into the many ecological niches that became
E 101
available after the end-Cretaceous extinction. There are fourteen orders in the sub-
class, in some classifications grouped as four large clades: Afrotheria,
Euarchontoglires, Laurasiatheria, and Xenarthra. Early Cretaceous to Recent.
Euthycarcinoidea: an obscure group of arthropods with affinities to both insects
and crustaceans; about an inch long, they had a cephalon with small antennae, an
abdomen with many legs, and a terminal tail spine; their fossil record is sparse but
widespread: Australia, North America, Scotland, Germany, and France; Silurian to
Middle Triassic.
Euthycarcinus “straight crab,” a genus of obscure arthropods in the subclass
Euthycarcinoidea, q.v.; known from Germany and France, Carboniferous to
Triassic.
Eutriconodonta: an enigmatic order of primitive mammals of the Mesozoic; for-
merly called Triconodonta, a polyphyletic grouping, but now restricted to a mono-
phyletic assemblage of a dozen carnivorous or omnivorous genera; they include the
largest Mesozoic mammals: the badger-size Repenomamus (3 feet long and weigh-
ing 30 pounds) and Jugulator amplissimus, somewhat larger; Middle Jurassic to
Late Cretaceous.
evolute: “unrolled,” especially of ammonoids, loosely coiled, with successive
whorls barely overlapping earlier ones, so that all whorls are visible.
evolutionary adaptation: the process by which a taxon, in the course of many
generations, accommodates itself to its environment, eventually giving rise to new
species as a result of the changes it undergoes.
Exogyra: a large genus of marine bivalves, oysters, in the still-extant family
Gryphaeidae and extinct subfamily Exogyrinae; characterized by a very thick shell
and a spirally twisted left valve; global distribution but declining after the Cretaceous;
Jurassic to Miocene.
extinction: the global disappearance of a taxon resulting from its failure to pro-
duce surviving direct descendants. A marked decrease in the amount of life on Earth
is referred to as a mass extinction. Background extinctions are those that occur
continuously between the periods of mass extinction. The extinctions at the end of
the Ordovician, Devonian, Triassic, Permian and Cretaceous periods are considered
mass extinctions, and it is likely there were other mass extinctions before the
Phanerozoic.
F
Fabaceae: the principal family of plants in the order Fabales; the most prominent
subfamilies are the Papilionoideae and Mimosoideae, important for their role in fix-
ing atmospheric nitrogen in concert with symbiotic bacteria such as the Rhizobiales;
the earliest-known fossil specimens have been found in Paleocene strata, but the
family’s roots go back to the radiation of angiosperms in the Early Cretaceous;
global distribution, Paleocene to Recent.
Falcarius utahensis: a primitive therizinosaurid saurischian dinosaur, known only
from a find near the Green River in Utah. About 13 feet long and 4 feet tall, it is
named for its large, sickle-shaped claws. Falcarius is notable for its herbivorous
characteristics — small teeth adapted for shredding leaves, large gut, stout legs, and
a long neck — all of which contrast with its smaller and faster carnivorous ancestors
and seem to connect Falcarius with the later therizinosaurs, which were large her-
bivores. Early Cretaceous, 126 Ma.
Falcatus: a genus of very small early sharks; characterized by an unusual dorsal fin
that hung over the head of the male; also by a tall and strongly forked tail, and by
very small pectoral and pelvic fins; North America, Middle Mississippian to Early
Pennsylvanian.
faunal succession: the principle that groups of fauna and flora change over time,
each succeeding group above the previous group in sedimentary rock strata that are
often characteristic of certain intervals of time; established by William Smith, it is
the basic concept of biostratigraphy.
faunal zone: a period of time or a stratigraphic unit that is characterized by the
presence of a unique assemblage of fossils known as a guide fauna.
faunule: a small fauna, especially a small group of animal fossils found only in a
single stratum.
Favositidae: a family of colonial corals in the extinct order Tabulata; the tightly
packed corallites are long, thin tubes with punctate walls pierced by mural pores;

104 F
the type genus Favosites, called honeycomb coral, is especially important in petro-
leum geology; global distribution, Late Ordovician to Late Triassic.
Fenestella: a large genus of stenolaemate bryozoans in the family Fenestellidae;
fan-shaped colonies with a net-like surface; especially widespread from the Middle
Devonian to the late Permian; global distribution, Middle Ordovician to Late
Triassic.
Fenestellidae: a large family of stenolaemate bryozoans in the order Fenestrida;
the colony’s skeleton consists of many stiff branches connected by crossing arms,
resulting in a fenestrate, fan-shaped net; the shape, position, and orientation of the
zooecia, used to identify different species, vary greatly; global distribution,
Ordovician to Late Triassic.
fenestra: in biology, a hole or small opening in an organism’s body; in paleontol-
ogy, fenestrae are important usually as slits or holes in the skull, often behind the
orbits (as in diapsids and synapsids), or in an individual bone (as in a neck frill); the
root fenestr- also appears in many names of organisms with net-like structures.
Fermeuse Formation: one of several formations on the Avalon Peninsula in
Newfoundland containing fossils of the Ediacaran Mistaken Point assemblage, q.v.
Fezouata Formation: a Tremadocian Konservat-Lagerstätte in Morocco, discov-
ered in the 1990s, that has produced very well-preserved fossils from a period
between the Lagerstätten of the Cambrian and the Soom Shale of the Late
Ordovician; among the described species are the sponge Pirania auraeum and a
possible marrellomorph tentatively named Furca mauretanica; North Africa, Early
Ordovician, 480–477 Ma.
Filicopsida: the ferns, a class of plants in the division Tracheophyta; formerly
called Pteridophyta and Polypodiopsida; the origins and the classification of ferns
are unclear; global distribution, Devonian to Recent.
First Family Site: the location in Hadar, Ethiopia, of a cluster of finds of
Australopithecus afarensis, including some nearly complete skeletons, the most
famous called “Lucy”; most of these specimens are dated between 3.2 and 3.4 Ma.
Fistuliporidae: a large and diverse family of stemless marine bryozoans in the
class Stenolaemata and extinct order Cystoporata; the colonies were initially
encrusting, and later genera became dendroid; Silurian to Late Triassic.
flatfish: fish with the unusual pleuronectiform character of having both eyes on the
top side of the body; this trait has made flatfish important in the history of evolution,
research on some transitional species showing gradual movement of one eye toward
the upper side; see Amphistium, Gerrothorax, and Heteronectes.
Flexibilia: a small subclass of stalked or creeping crinoids; characterized by a flex-
ible tegmen with open ambulacral grooves, the calycal plates being loosely united;
global distribution, Middle Ordovician to late Permian.
F 105
Florissant Fossil Beds National Monument: an important site in Central

Colorado for insects and plants in lacustrine deposits of volcanic ash laid down at
the boundary of the Eocene and Oligocene; it contains large petrified stumps of
sequoia-like trees, the finest fossil insects of North America, and many angiosperms;
North America, late Eocene.
Florissantia: a genus of sterculioid angiosperms in the family Malvaceae; its
1-inch-wide five-petaled flowers have been found throughout North America, espe-
cially in Oregon and the Florissant Fossil Beds of Colorado; North America, Eocene
to Oligocene.
Fontechevade Man: name given to two hominin specimens found at Fontechevade,
France; bearing characteristics of both H. sapiens and H. neanderthalensis, they
were first dated as Pliocene and interpreted as a surprisingly early appearance of
modern humans in Europe, antedating Neanderthals; but research in 2007 dated
them as Pleistocene, when H. sapiens and H. neanderthalensis co-existed.
Foraminifera: a class of one-celled organisms in the phylum Retaria, almost all
aquatic and mostly marine. They are usually less than 2 mm but range up to several
centimeters long. These one-celled organisms are mostly benthic, living on or under
the sea floor, while a few are planktic. Most foraminiferans develop a calcitic test,
often in several concentric layers, so that many are important index fossils.
Planispiral taxa are coiled, in either an evolute or an involute manner. Almost 9000
extant and almost 2000 extinct species are known. Cambrian to Recent.
Fordilla: the earliest-known bivalve, a widespread genus of small, primitive
bivalves in the order Fordillida; suboval, about one-tenth of an inch long; only three
species of Fordilla are currently recognized, all from the early Cambrian, about
520 Ma; see also Pojetaia.
form genus: a biological classification of a part of an organism, used when the
relationship of the part to the whole organism is unclear, and mainly for plants
because they seldom fossilize whole; also referred to as a parataxon.
Formicidae: the ants, a family of hymenopteran insects in the superfamily
Vespoidea; ants evolved from primitive wasps between 130 and 100 Ma, along with
the flowering plants; they became the ecologically dominant insect by the end of the
Paleocene, and 10 percent of the species known from the Eocene are still present;
Early Cretaceous to Recent.
form taxonomy: a taxonomic system based primarily on morphology rather than
on phylogeny (biological relationship).
Fort Union Formation: a formation in the Powder River Valley in Wyoming and
Montana that straddles the period between the end of the Cretaceous and the begin-
ning of the Paleocene; it contains many important fossils, such as a still-unnamed
ornithurine bird that may have survived the extinction, the taeniodont mammal
Alveugena, and plant fossils such as Cercidiphyllum; 66–64 Ma.
106 F
fossil: There are two primary types of fossils. Body fossils are parts of an organism
preserved by carbonization or permineralization, or as casts or molds. Trace fossils,
such as worm and mammal tunnels or tracks, are records of some action of an
organism. Fossils are typically found in sedimentary strata but may be found
deformed in metamorphic deposits. Mud flows, lahars, and even igneous lava flows
also often preserve many types of fossils.
fossiliferous: fossil-bearing, referring to any deposit or stratum of rock that con-
tains the fossilized remains of an organism.
fossorial: adapted to digging and living in burrows, typical of many rodents but
also badgers and some bees and wasps.
fossoriality: the habit of living in burrows underground; known in many instances
since the Paleozoic but increasingly important as a lifestyle for small animals as
open grassland increased globally during the Miocene.
Fractofusus misrai: an Ediacaran rangeomorph fossil known only from the
Mistaken Point assemblage; Newfoundland, Canada, 575–560 Ma.
Frondicularia: a genus of foraminifera in the family Lagenidae; generally leaf-
shaped; global distribution, Triassic to Recent.
Fruitadens haagarorum: a species of ornithischian dinosaurs in the family
Heterodontosauridae; only about 4 inches high, between 2 and 3 feet long, and
weighing less than 2 pounds, it is the smallest known ornithischian; like modern
birds, it ate fruit and insects; closely related to horned, duck-billed, and armored
dinosaurs; United States, Late Jurassic, 150 Ma.
Fruitafossor: a monospecific genus of small burrowing mammals; known only
from a complete skeleton found in 2005 in Fruita, Colorado, F. windscheffeli was
only 4 inches long but has several adaptations for fossoriality and probably fed
mainly on termites; it is a striking example of convergent evolution, being unrelated
to any other known animals with similar habits; United States, Late Jurassic.
fullerenes: the term for a molecule of carbon in a hollow sphere or other shape.
They form closed carbon lattices found in rocks, and because they may trap atmo-
spheric gases at the time the rocks are formed, they have been used since the 1990s
to investigate atmospheric changes in different geological time periods. For exam-
ple, some investigators claim that fullerenes from some end-Permian rocks display
noble-gas isotope ratios that are typical of meteorites, not Earth; these ratios are
possibly evidence of a meteorite impact that may have contributed to the end-
Permian extinction.
furcula: generally, the wishbone of birds and some dinosaurs, but see also
Collembola.
Fusulinida: an extinct order of foraminiferans, important as index fossils between
the Devonian and the end of the Permian; the order contains three superfamilies —
F 107
the Endothyroidea, Fusulinoidea, and Parathuramminoidea; global distribution,

early Silurian to late Permian.
Fusulinidae: a family of large, spindle-shaped foraminiferans in the superfamily
Fusulinoidea; mostly about the size of a grain of wheat but some as long as 2 inches;
global distribution, Early Pennsylvanian to late Permian.
Fusulinoidea: a superfamily of foraminiferans in the extinct order Fusulinida; it
comprises the families Fusulinidae, Ozawainellidae, Schwagerinidae, Staffellidae,
and Verbeekinidae; global distribution, generally considered extinct at the end of the
Permian.
Futalognkosaurus: a genus of herbivorous sauropod dinosaurs; known from three
partial specimens, it was found near Lake Barreales in Argentina in 2000; one of the
largest dinosaurs known, it is estimated to have been at least 90 feet long; Late
G
Galeritidae: a family of irregular echinoids in the superfamily Conulidea; charac-

terized by simple ambulacral plating, they appeared about 70 Ma and became
extinct at the end of the Cretaceous; Europe and Asia.
Galliformes: the order of birds that includes chickens, pheasants, turkeys, Guinea
fowl, and grouse. The galliforms probably arose in the late Cretaceous, but few
Cretaceous candidates have been proposed; one is Vegavis iaai, a possible anseri-
form bird found in Antarctica. The galliform family Phasianidae is represented by
several genera that appeared in the Eocene. Global distribution, Cretaceous to
Recent.
Gampsonychidae: a family of primitive crustaceans in the extinct order
Palaeocaridacea, possibly related to the eurypterids; Carboniferous.
ganoid scales: fish scales with a bony, enamel-like surface, common in sturgeon
and gar; generally diamond-shaped and non-overlapping but linked together at
joints.
Gansus yumenensis: a species of Cretaceous ornithuromorphan birds found in
Gansu and Liaoning Provinces; known from several partial specimens, it seems to
be close to the ancestry of modern birds; similar in various ways to Hesperornis,
loons, and ducks; China, Early Cretaceous, about 120 Ma.
Garstang’s Hypothesis (of vertebrate origins): an early twentieth-century the-
ory of chordate evolution that proposed an alternative route for chordate evolution
from echinoderms; Garstang opposed Ernst Haeckel’s theory, based on the principle
that ontogeny recapitulates phylogeny, which is now discredited; most aspects of
Garstang’s theory are now accepted.
Gasparinisaura cincosaltensis: a species of herbivorous ornithopod dinosaurs;
about 6 feet long, it weighed about 40 pounds; Argentina, Late Cretaceous,
85–82 Ma.

110 G
Gastonia: a genus of basal ankylosaurian dinosaurs, generally classified in the

family Nodosauridae; from 15 to 20 feet long and about 1 ton, Gastonia had a
strong set of spikes, scutes, and plates; Utah, United States, Early Cretaceous,
126–123 Ma.
Gastornis: a genus of large flightless birds in the order Anseriformes; they had
massive skulls and legs and were probably good runners; although they are quite
similar to the South American phorusrhacids of the Miocene, the two groups are not
related; the largest species, G. giganteus and Diatryma, grew to almost 7 feet high
and were about the size of the largest moas; researchers still dispute whether they
were herbivorous, carnivorous, or both; North America and Europe, late Paleocene
to Eocene.
Gastrioceras: a genus of goniatitid ammonoids in the family Gastrioceratidae;
characterized by a wide umbilicus and a globose shell with a goniatitic (simple and
relatively smooth) suture pattern; widespread in the Middle to Late Pennsylvanian,
several species are index fossils; North America and Europe, Late Carboniferous,
possibly to Permian.
Gastropoda: “stomach foot,” the class of mollusks that includes land, marine, and
freshwater snails and slugs; the most speciose class of mollusks, originating in the
late Cambrian; see Chippewaella and Strepsodiscus.
Geiseltaliellus: a genus of lizards in the family Iguanidae; see Palaeopython;
Europe, Eocene to Oligocene.
Gelocidae: a family of hornless ungulate mammals, possibly polyphyletically
related to the Pecora; among the earliest ruminants but more advanced than the
Hypertragulidae; North America, Europe, Asia, and Africa; Eocene and Miocene.
Geminiraptor suarezarum: a species of raptor-like dinosaurs in the family
Troodontidae, found in Utah in 2010. Its head was large, as in raptors; but since it is
known only from a partial jaw bone, its relationship is uncertain. It is dated between
130 and 125 Ma, almost 50 million years earlier than the numerous North American
raptors of the Late Cretaceous. United States, Early Cretaceous.
Gemündina: a monospecific genus of rhenanid placoderms in the family
Asterosteidae; its flattened body and very large, wing-like pectoral fins gave it a
shape similar to that of rays (see Batoidea), which evolved 200 million years later
in the Triassic; G. stuertzi is known from several exceptional specimens from the
Hunsrück Lagerstätte; notable for the absence of the characteristic placoderm tooth-
plates; Germany, Early Devonian.
generalized: of an organism: having few specific adaptations to its ecological niche.
Geniohyus: a genus of primitive hyraxes in the extinct family Pliohyracidae and
subfamily Geniohyinae; Egypt, known only from the early Oligocene.
Geologic Time Scale (simplified for this book’s purposes): Precambrian Eon, up
to 541 Ma, ending in the Neoproterozoic Era, which consists of the Tonian,
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Cryogenian, and Ediacaran Periods. Phanerozoic Eon (541 Ma to the present),

consisting of three eras: Paleozoic Era, Cambrian, Ordovician, Silurian, Devonian,
Carboniferous (for North America, divided into Mississippian and Pennsylvanian
Epochs), and Permian; Mesozoic Era, Triassic, Jurassic, and Cretaceous; and
Cenozoic Era, divided into the Paleogene Period (Paleocene, Eocene, and Oligocene
Epochs), Neogene Period (Miocene and Pliocene Epochs) and Quaternary Period
(Pleistocene and Holocene Epochs). The Phanerozoic periods are divided into
epochs, and the epochs into shorter spans called stages. For a more detailed list, see
the ICZN’s International Chronostratigraphic Chart.
Georgiacetus: a genus of protocetid whales in the suborder Archaeoceti; its pelvis
was not attached to its spine, representing the loss of functional hindlimbs in whales’
evolution from terrestrial ancestors; Southeastern United States, middle Eocene,
40–37 Ma.
Geosaurus: a genus of marine crocodylomorphs in the clade Thalattosuchia, fam-
ily Metriorhynchidae, and subfamily Geosaurinae; up to 10 feet long, its classifica-
tion is contentious because of small differences in specimens; probably related to
Dakosaurus, Cricosaurus, and Rhacheosaurus; South America and Europe, Late
Jurassic to Early Cretaceous.
Gerrothorax: a probably monospecific genus of stereospondyl amphibians in the
family Plagiosauridae; about 3 feet long and extremely flattened, G. pulcherrimus
was pleuronectiform, with both eyes on the top of its broad head; it retained its lar-
val gills, so that it could remain underwater indefinitely, probably lying on the bot-
tom awaiting prey; Europe, Middle to Late Triassic.
Ghost Ranch: a Konzentrat-Lagerstätte in New Mexico that is known for thou-
sands of specimens of dinosaurs from the end of the Triassic; the best-known dis-
covery was the first Coelophysis; United States, Late Triassic, 220–202 Ma.
Giganotosaurus: one of the largest known carnivores, a genus of carcharodonto-
saurid dinosaurs; up to 43 feet long and weighing up to 14 tons, Giganotosaurus
was almost as large as Spinosaurus; known only from a skull, femur, lower jaw, and
other fragments; the type species is G. carolinii; Argentina, Late Cretaceous,
99–96 Ma.
Gigantopithecus: a genus of hominoid primates in the family Pongidae, known
from partial skeletons and teeth of the Pliocene in India and the Pleistocene in
China, as recent as 100,000 years ago; they stood as high as 10 feet and weighed up
to 1200 pounds; the largest known ape, Gigantopithecus was primarily
herbivorous.
Gigantoproductus: a genus of large brachiopods in the order Productida and the
family Monticuliferidae; up to 12 inches wide, G. giganteus is the largest brachio-
pod known; global distribution, Carboniferous.
Gigantopteridales: a diverse group of large spermatophyte trees at the end of the
Paleozoic. Among the most advanced plants of the Paleozoic, they are possible
112 G
ancestors of later plant forms, including angiosperms. They had many characteris-
tics of flowering plants but are not known to have flowered themselves; in fact, their
mode of reproduction is unclear. They appeared near the end of the Permian and
flourished in Asia and North America for 10 million years, briefly surviving the
great extinction before disappearing in the Early Triassic.
Gigantoraptor xu: a species of theropod dinosaurs in the bird-like family
Caenagnathidae; dated at 85 Ma but possibly younger, it had no teeth but had large
claws and probably a large beak; not well-known, it is estimated to have been about
25 feet long and 2 tons in weight; see also Anzu; Mongolia, Late Cretaceous,
Campanian.
Gigatitan: a genus of large predatory insects in the order Titanoptera; notable for
fluting on the wings that could produce stridulation when rubbed by the hindlegs, in
the manner of modern crickets; it had a 16-inch wingspan and stout spines on the
forelegs; Madygen Formation, Kyrgyzstan; Late Triassic, 230–225 Ma.
Ginkgoites: a form genus of the leaves of the Ginkgoales, with indented but not
deeply divided fan-shaped leaves; global distribution in the late Mesozoic.
Ginkgophyta: a division of gymnospermous plants that first appeared in the late
Permian but is now represented by only one genus, Ginkgo biloba, in the order
Ginkgoales; the ginkgos are known in the fossil record primarily by their distinctive
lobed and veined leaves.
Giraffatitan: a genus of titanosaurian sauropodomorph dinosaurs in the family
Brachiosauridae; known from partial skeletons found in Tanzania, Giraffatitan is
closely related to Brachiosaurus; it is estimated to have been more than 70 feet long
and to have weighed up to 80 tons; Africa, Late Jurassic.
Girtyocoelia: a genus of Permian calcareous sponges in the subclass
Ceractinomorpha. An important reef builder late in the Permian, especially in the
economically important Permian reef complex of Texas, it seemed to be extinct in
the Early Triassic but then reappeared in the Middle Triassic. Close relatives appear
earlier, in Ordovician strata, but it is unclear whether they are Girtyocoelia itself.
North America, Middle Pennsylvanian to Permian or Triassic.
Girvanopyge: a genus of deep-ocean trilobites in the order Asaphida and family
Cyclopygidae; characterized by an unusual cylindrical thorax and a very broad gla-
bella; Europe, Middle to Late Ordovician.
glabella: the convex axial segment of the cephalon in trilobites, usually bulbous on
the anterior end; the animal’s major organs lie beneath the glabella, which is cov-
ered by a solid cephalic shield, the carapace.
Glabrocingulum: a genus of low-spired marine snails in the family Gosseletinidae;
United States, Carboniferous.
G 113
Glacialisaurus: a genus of relatively small massospondylid dinosaurs of Antarctica,

one of several sauropodomorphs found in the Hanson Formation; known only from
fragments of two specimens; Early Jurassic.
Gliridae: a still-extant family of sciuromorph rodents in the superfamily Gliroidea,
descended from the Ischyromyidae; the family includes the modern dormice;
Europe, Asia, and Africa; early Eocene to Recent.
Global Boundary Stratotype Sections and Points: Markers selected by the
International Commission on Stratigraphy on which the International
Chronostratigraphic Chart is based. The choices are based principally on strati-
graphic first appearances of particular fossils, with a few extinctions also noted. The
list also includes some markers based on paleomagnetic and climatic data. See for
example Cardioceras, Iapetognathus, Marsupites, and Monograptus.
Globidens: a genus of small mosasaurs known for its rounded teeth, specialized for
durophagy — crushing turtles, ammonoids, and other hard-shelled animals; North
America, Africa, and Asia; Late Cretaceous, 85–66 Ma.
Globigerinida: still-extant order of globose planktic foraminiferans, important as
an index fossil in the Cretaceous. Simple but multi-chambered foramiferans, the
globigerines seem to be unusually resistant to mass extinctions. The genus
Globigerina, which appeared in the Eocene, now dominates, and large areas of
ocean floor worldwide are covered by the bodies of various species in the order,
constituting what is called a Globigerina ooze. In reference to the end-Cretaceous
extinction, see also Guembelitria. Global distribution, Jurassic to Recent.
glossopterid flora: an important group of pteridospermatophyte plants found in
Permian and Early Triassic strata laid down in Gondwana; the absence of glossop-
terids in northern land masses was important early evidence of the existence of the
southern supercontinent; Antarctica, Australia, New Zealand, Southern Africa,
India, and South America.
Glossopteris: a genus of seed ferns in the extinct order Glossopteridales; character-
ized by a tongue-shaped leaf with a prominent midrib and veins, arranged spirally
around the ends of the branches; it was a dominant plant in the Permian glossopterid
flora but is rare in Triassic and Jurassic deposits; Australia, Antarctica, India, Asia,
Southern Africa, and South America; Permian to Jurassic.
Glyphocyphidae: a family of epifaunal sea urchins, irregular echinoids in the still-
extant superorder Echinacea and order Camarodonta; South America, Africa,
Europe, and Asia; Early Cretaceous to Eocene.
Glyptocrinina: a suborder of camerate crinoids in the order Monobathrida; large,
one-inch thecae in some families; Middle Ordovician to Permian.
Glyptodontinae: a subfamily of large, heavily armored xenarthran mammals in
the still-extant order Cingulata and family Chlamyphoridae. Their striking armor
consisted of a dome-shaped covering of scutes over the whole body and skull.
114 G
The type genus, Glyptodon, was about 11 feet long and almost 5 feet high, and
weighed up to 2 tons; the southern genus Doedicurus was slightly larger. Living in
South America for 20 million years, they entered North America in the Pliocene by
migrating across the newly formed Panama land bridge. They were probably hunted
to extinction by the first humans to arrive in the Americas. South and North America,
early Miocene to Pleistocene.
Glyptops: a genus of cryptodire turtles in the family Pleurosternidae; the type spe-
cies, G. plicatulus, is about a foot long and is well-represented in the Morrison
Formation; restricted to North America, Late Jurassic to late Cretaceous.
Glyptostrobus: a still-extant genus of trees in the family Cupressaceae; related to
the modern bald cypress and still represented by the Chinese swamp cypress,
Glyptostrobus bears a cone at the tip of a branch covered with scale-like leaves and
grew as high as 115 feet; North America, Europe, and Asia; Cretaceous to Recent.
Gnathobelodon: a genus of proboscideans in the family Gomphotheriidae, charac-
terized by a projecting jawbone that resembled a shoe horn; similar to Amebelodon
and Platybelodon except for having no lower tusks but simply the long jawbone;
North America, middle to late Miocene.
Gnathodontidae: a family of conodonts in the order Ozarkodinida; widespread in
the early Carboniferous, they declined rapidly and became extinct in the early
Permian.
Gnathostomata (echinoids): a superorder of irregular sea urchins that possess a
feeding lantern; lower Jurassic to Recent.
Gnathostomata (vertebrates): “jaw-mouthed,” a superclass or infraphylum of
jawed vertebrates. The gnathostomes appeared in the Middle Ordovician, derived
from agnathans that had developed pectoral and pelvic fins as well as enlarged ante-
rior gill arches, which folded over forward in some species, gradually becoming
upper and lower jaws. Jaws gave such an advantage in predation that the gnathos-
tomes became the dominant vertebrates in the oceans from the Silurian through the
Early Devonian. By the end of the Devonian, almost all agnathans had become
extinct. The gnathostomes were the ancestors of almost all modern vertebrates,
descending through placoderms or osteichthyans. Middle Ordovician to Recent.
Gnetales: an unusual, still-extant order of Mesozoic gymnosperms in the class
Gnetophyta, probably related to the Bennettitales; the Gnetales had a vascular sys-
tem similar to that of angiosperms and have been considered possible ancestors of
the flowering plants; still-extant genera include Welwitschia, Gnetum, and Ephedra;
widespread in the Cretaceous, but possible specimens are known from the Permian,
Triassic, and Jurassic; Australia, Antarctica, Africa, and North and South America.
GOBE: the Great Ordovician Biodiversification Event; see Ordovician.
G 115
Gobiatherium: a genus of large ungulates in the extinct suborder Dinocerata;

unlike its uintathere relatives, Gobiatherium had no tusks, horns, or knobs on its low
skull; China, middle Eocene.
Gobipteryx: a genus of early toothless enantiornithine birds; until 1995, when
Confuciusornis was discovered, Gobipteryx was the earliest-known beaked bird;
Mongolia, Late Cretaceous, 75–70 Ma.
Gogia: a genus of mainly North American echinoderms in the class Eocrinoidea; a
common middle Cambrian eocrinoid, the plate-covered Gogia was one of the earli-
est stalked echinoderms, dating back to the late early Cambrian and disappearing
toward the end of the period; North America and Spain.
Gogo Formation: an Upper Devonian (Frasnian) site in Western Australia, a
Konservat-Lagerstätte containing exceptionally well-preserved soft-tissue fossils.
Among its notable specimens are three species of armored placoderm fish that seem
to have been viviparous: Austroptyctodus gardineri, Materpiscis attenboroughi, and
Incisoscutum ritchiei. Two specimens of the latter species show evidence of fertil-
ization of eggs inside a female fish occurring as much as 30 million years earlier
than previously thought, around 380 Ma. Since the fish evidently gave birth to live
young, the two fossils represent the earliest-known vertebrate mothers. The speci-
men of Materpiscis contains a fetus and an umbilical cord.
Gomphos elkema: the oldest complete skeleton of a rabbit-like mammal, found in
the Nemegt Basin of Mongolia; closely related to both lagomorphs and rodents, it
supports the idea that the two groups shared a common ancestor that lived near the
end of the Cretaceous; it also indicates that lagomorphs appeared no earlier than
65 Ma; Mongolia, 55–50 Ma.
Gomphotheriidae: a diverse family of proboscideans in the suborder
Gomphotherioidea; the largest species weighed up to 7 tons and were about 10 feet
high at the shoulder; most species had four tusks and may have had elephant-like
trunks; the last of the gomphotheres became extinct about 9000 years ago; global
distribution except Australia and Antarctica, late Oligocene to Holocene.
Gomphotherium: a genus of proboscideans in the family Gomphotheriidae; 8 feet
high at the shoulder and weighing 5 tons, it had four tusks; North America, Europe,
Asia, and Africa; early Miocene to early Pleistocene.
Gondwana: the Paleozoic southern supercontinent, named for the Gondwana geo-
logical system (dated from the Carboniferous to the Jurassic) in the region of India
where early Glossopteris investigations occurred. Located in the lower part of the
Southern Hemisphere, Gondwana included what is now South America, Africa,
Madagascar, India, Antarctica, Australia, and New Zealand. See also
paleogeography.
Gondwanatheria: a group of primitive mammals known from fragmentary speci-
mens from the Late Cretaceous through the Miocene, related to haramiyidans and
116 G
multituberculates; a discovery in 2010 of Vintana sertichi, an almost complete skull

of a 20-pound gondwanatherian, greatly expanded knowledge of the group.
Goniatitida: a large order of Paleozoic ammonoids; characterized by goniatitic
sutures, with a simple pattern of smooth lobes and saddles; widespread from the
Devonian to the end of the Permian.
Goniopholis: a genus of Late Jurassic medium-sized crocodylomorphs in the clade
Thalattosuchia and family Goniopholididae; 6–9 feet long, Goniopholis was very
similar to modern crocodiles and is considered a neosuchian; possibly the earliest
goniopholidid is Calsoyasuchus, known from a partial skull found in Arizona and
dated at 196.5 Ma; cosmopolitan, Late Jurassic to Early Cretaceous.
Goniopora: a still-extant genus of scleractinian corals in the family Poritidae; a
large and diverse group, generally massive but some species branching; global dis-
tribution, Early Cretaceous to Recent.
Gorgonopsidae: a family of theriodont therapsids; early gorgonopsians were dog-
sized and some later species were the size of a large bear; they had some mamma-
lian characters but are not considered in the direct line of descent, which goes
through their cousins the cynodonts; Africa and Europe, middle to late Permian.
Gorgosaurus: a genus of tyrannosaurid dinosaurs in the subfamily Albertosaurinae;
bipedal, as long as 30 feet and weighing almost 3 tons, it was smaller than
Tyrannosaurus or Tarbosaurus; Canada, Late Cretaceous, 77–75 Ma.
Graeophonus: a genus of whip spiders in the order Amblypygi and family
Paracharontidae; about half an inch long, three species of Graeophonus are known
from Carboniferous sites in Nova Scotia, Illinois, and England.
Grande Coupure: a term used for the great turnover of mammalian faunas in
Europe at the boundary of the Eocene and Oligocene, marked especially by the
migration into Europe of many new Asian taxa.
Graptolithina: the graptolites, a class of colonial marine invertebrates in the sub-
phylum Hemichordata, probably related to the still-extant pterobranchs. A grapto-
lite colony is called a rhabdosome and has a varying number of stipes (branches);
the individual zooids were housed in thecae on the stipes. Very little is known about
the structure, morphology, or feeding habits of the zooids (see Psigraptus). The
largest zooids were about an eighth of an inch long, and they probably could not
leave the theca. Graptolites evolved from bottom-encrusting to sessile forms in the
Cambrian, with the earliest sessile order, the Dendroidea, persisting to the
Carboniferous. It gave rise to the planktic Graptoloidea, which diversified rapidly
worldwide during the Ordovician. Graptolites are important as index fossils of the
early and middle Paleozoic Era because of their global occurrence and their rapid
evolution and radiation into many different species; the strata of the Ordovician and
Silurian have been divided into graptolite biozones of about one million years each.
The anisograptid, dichograptid, diplograptid, leptograptid, and monograptid faunas
G 117
are the most important. See also Chaunograptus. Middle Cambrian to Carboniferous,
global distribution in the Ordovician and Silurian.
Graptoloidea: an order of graptolites and the only completely planktic group;
generally characterized by a smaller number of stipes than the Dendroidea; they
appeared early in the Ordovician and became the most prominent planktic animals
worldwide until their extinction in the Devonian; Early Ordovician to Early
Devonian.
grasses: the Poaceae (or Gramineae). The earliest-known grass fossils go back to
the time of the dinosaurs, with phytoliths from grasses known in dinosaur coprolites
toward the end of the Early Cretaceous. Changing environmental conditions in the
Paleocene may have encouraged the spread of grasses, but the first grasslands seem
to have appeared only much later, in the middle Eocene, and then to have evolved in
conjunction with the rise of ungulate mammals in the Eocene. Since the Eocene,
grasses have spread globally and are the most common of all plants today. See also
fossoriality and ungulates.
Great American Biotic Interchange: the mingling of animal populations between
South and North America that was made possible by the rise of the Isthmus of
Panama about 3 million years ago, joining the two previously separated continents.
Only a few terrestrial animals had moved from one continent to the other until the
late Pliocene. The interchange resulted in many regional changes, notably the
extinction of several South American groups because of the introduction of large
new predators from the North.
Great Oxygenation Event: term referring to the production of oxygen by organ-
isms, mainly cyanobacteria; although the timing and causes are debatable, signifi-
cant oxygen production began around 2400 Ma and resulted in levels of atmospheric
oxygen sufficient for complex oxygen-breathing and sexually reproducing organ-
isms sometime in the Neoproterozoic Era, between 1000 and 600 Ma.
Green River Formation: a lower Eocene formation of the Western United States
with significant outcroppings in Colorado, Utah, and Wyoming, including Fossil
Butte National Monument; the formation has produced large numbers of fish, espe-
cially in finely varved lacustrine strata; other important fossils are early bats, the
North American crocodilian Borealosuchus, and many stingrays, turtles, snakes,
plants and insects; Eocene, 54–49 Ma.
Greererpeton burkemorani: a genus of stem-tetrapod amphibians in the family
Colosteidae. Almost 5 feet long, its body was eel-like, with a flattened skull 7 inches
long. The short limbs had five digits but were used mainly for steering in the water.
It was almost completely aquatic, remaining in rivers and lakes while other stem-
tetrapods were moving ashore in the Carboniferous. North America, Late
Mississippian.
Grès à Voltzia: a Middle Triassic Konservat-Lagerstätte in northeastern France,
dated about 243 Ma. Its brackish-water deposits were laid down in a deltaic environ-
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ment similar to that of the Carboniferous Mazon Creek site. The finely laminated
clay rock of the Grès à Meules stratum contains very well-preserved specimens of
terrestrial and aquatic fossils, including plants, millipedes and other arthropods,
brachiopods, crustaceans, and insects. Fish (see Dipteronotus) and amphibians,
mainly capitosaurid temnospondyls, are also represented.
Groeberiidae: a small family of South American marsupials, rodent-like but with
a skull architecture and dentition different from any other mammals; late Eocene to
early Oligocene.
Groenlandaspis: a genus of arthrodiran placoderm fish; ranging from 3 to 19
inches long, it had small toothplates and strong, pyramid-shaped thoracic armor;
global distribution, Late Devonian.
Gryphaea: a genus of bivalves in the order Ostreoida and family Gryphaeidae. The
distinctively convoluted Gryphaea is characterized by prominent growth bands in
the thick, twisted upper valve. It was considered in the first half of the twentieth
century to be a possible example of orthogenesis because of an erroneous interpreta-
tion of its coiling growth pattern, disproved by Anthony Hallam and Stephen Gould.
Global distribution, Late Triassic to Eocene.
Gryphaeidae: a still-extant family of bivalves, sometimes called “foam oysters,”
in the order Ostreoida; global distribution, Middle Triassic to Recent.
Gryposaurus monumentensis: a genus of herbivorous dinosaurs in the family
Hadrosauridae; a duckbill, Gryposaurus probably ate many kinds of vegetation,
having as many as 800 teeth set in a powerful jaw; North America, Late Cretaceous,
83–74 Ma.
GSSP: see Global Boundary Stratotype.
Guaibasauridae: an early family of saurischian dinosaurs; first proposed in 1999,
its makeup is still in flux; see Panphagia, Chromogisaurus, Agnosphitys, and
Saurischia; Argentina and Brazil, Late Triassic, 231–208 Ma.
Gualicho shinyae: a species of allosaurid theropod dinosaurs similar but not
closely related to Tyrannosaurus rex; also described as similar to Deltadromeus but
with much shorter forelimbs. Known from a partial skeleton found in the Huincul
Formation of Argentina and described in 2016, it was about 16 feet long and weighed
half a ton. Like T. rex, Gualicho had a two-digit manus on forelimbs the size of
those of a human child, a result of convergent evolution. Argentina, Late Cretaceous,
95 Ma.
Guanlong wucaii: a species of theropod saurischian dinosaurs in the superfamily
Tyrannosauroidea, perhaps the earliest basal tyrannosauroid, dated at 160 Ma.
Known from two specimens described in 2006, G. wucaii was only about 4 feet high
and 10 feet long, with three fingers on its strong forelimbs. It bore a large but fragile
crest on its forehead, stretching from behind the nostrils to just behind the eyes and
about 3 inches high. The function of the crest is not known, but it would appear to
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be a liability in combat, since it is only as thick as a tortilla. Western China, Late

Jurassic.
Guembelitria cretacea: a species of planktic foraminiferans in the order
Globigerinida and family Guembelitriidae. Known as an opportunistic “disaster
species” because, although rare at the end of the Cretaceous, it is the most common
of several foraminiferans that survived into the Paleocene. Abundant in the earliest
Paleocene strata, it evidently thrived in the environmental conditions following the
extinction of many other species. See also Globigerinida. Europe, Australia, Asia,
and; South America; Cretaceous to Oligocene.
guide fossil: see index fossil.
Guildayichthys: a genus of early marine actinopterygian fish in the order
Guildayichthyiformes; only about 2 inches long, it had short, conical teeth; its disc-
shaped body and robust pectoral, dorsal, and anal fins gave it good maneuverability;
see also Discoserra; North America, Early Carboniferous.
Guiyu oneiros: a species of sarcopterygian fish, the earliest-known articulated
specimen of an osteichthyan; about a foot long, with ganoid scales, it was found in
Yunnan Province near the site where the similar but slightly later Psarolepis was
found; China, Early Devonian, 419–418 Ma.
gundi: see Ctenodactylidae.
Gunflintia: a genus of cyanobacteria that is among the earliest-known bacteria;
around 2 billion years ago, it and similar photosynthetic organisms began to increase
the oxygen content of the atmosphere to levels that supported more complex life
forms; first found in the 1950s in the Gunflint Chert of Canada and later in Australia,
Proterozoic to Cambrian.
Gurbanodelta kara: a species of basal metatherian mammals described in 2016,
probably in the order Deltatheroida, the latest-known deltatheroidan; known only
from a jaw fragment, it was a scansorial omnivore; dated 10 million years later than
the previously known deltatheroidans, it may be a Lazarus taxon if confirmed;
Xinjiang, China, late Paleocene.
Gymnarthridae: a family of tuditanomorph lepospondyl amphibians in the order
Microsauria; distinguished from other microsaurs by its single row of conical teeth;
North America and Europe, Late Carboniferous to early Permian.
Gymnocodiaceae: an enigmatic family of small algal fossils characterized by
irregular tubes with perforate walls; sometimes classified as rhodophytes and some-
times as chlorophytes; Europe, North and South America, Asia, and Africa; Permian
to Cretaceous.
Gymnolaemata: a class of bryozoans that includes most living bryozoans; global
distribution, Middle Ordovician to Recent.
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gymnosperms: a group of seed-bearing plants that includes cycads, conifers, and

ginkgos; characterized by “naked” seeds not enclosed in an ovary or seed-case as
with angiosperms; they appeared in the Late Devonian and rapidly spread
worldwide.
Gyracanthus: a genus of acanthodian fish in the family Gyracanthidae; 2–3 feet
long, Gyracanthus is notable for its fin spines, which were 18 inches long; global
distribution, Early Carboniferous.
Gyrogonites: the fossil cast of the nucule, the female reproductive structure, of
charophytes. The worldwide distribution and spiral ornamentations of these small,
nut-like casts make them important index fossils in some freshwater deposits.
Gyrogonites appear in the fossil record as far back as the Silurian.
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Haasiophis: a monospecific genus of marine snakes in the family Simoliophiidae;

about 3 feet long, H. terrasanctus is especially notable for its hindlimbs; known
from an almost complete skeleton, it figures in the ongoing debate about the origins
of snakes; Israel, Late Cretaceous, 94 Ma.
Haast’s eagle: see Harpagornis moorei.
Habrohagla: a genus of katydid-like orthopteran insects in the family
Prophalangopsidae, which was formerly called Haglidae; H. curtivenata was
4–5 inches long and was very similar to the larger species of modern katydids; also
called Parahagla; China and Russia, Early Cretaceous.
Hadrocodium wui: a species of mouse-like primitive mammal that has the earliest-
known example of several mammalian ear and brain features; known only from a
single skull, from which the body is estimated to have been about 1 inch long;
China, Early Jurassic, 195 Ma.
Hadrosauridae: a family of ornithischian duck-billed dinosaurs; one of the most
successful dinosaur families, persisting from 86 to 66 Ma; descendants of the iguan-
odontids, the hadrosaurs are divided into five subfamilies, the largest of which are
the Lambeosaurinae and the Saurolophinae; middle-sized herbivores, hadrosaurs
ranged in length from 18 to 54 feet; North and South America, Europe, Asia, and
Antarctica; Late Cretaceous, 90–66 Ma.
Hagerman Fossil Beds: an important late Pliocene site in South-Central Idaho, in
cliffs on the southwest bank of the Snake River. The site contains many mammals
and other remains from the period about 3.2 to 3.7 Ma, notably the Hagerman horse,
Equus simplicidens, which resembled a small zebra, about 4 feet high at the shoul-
der and weighing about 900 pounds. United States, Pliocene.
Haikouella: a genus of primitive animals, found in the Chengjiang biota; consid-
ered a probable chordate, it is similar to Yunnanozoon and to Pikaia of the Burgess
Shale; China, early Cambrian.

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Haikouichthys: a monospecific genus of primitive proto-vertebrate animals from

the early Cambrian; about an inch long, it resembles a narrow Myllokunmingia;
China, early Cambrian.
Halitheriinae: a subfamily of sirenian mammals in the family Dugongidae; the
type genus is Halitherium, which is very similar to modern dugongs; North and
South America, Africa, and Europe; early Oligocene to Miocene.
Halkieria: a genus of enigmatic Cambrian organisms tentatively considered mol-
lusks; in appearance like an armored slug, the halkieriids somewhat resemble
Wiwaxia; they are generally 2–3 inches long and flattened, and have the shape of an
elongated oval; known primarily from H. evangelista of the Sirius Passet site in
Greenland; North America, Europe, Asia, and Australia; early to middle Cambrian.
Hallopora: a large genus of “branching twig” stenolaemate bryozoans in the
extinct order Trepostomata and family Halloporidae. The Hallopora are character-
ized by their dendroid habit, large pores, and many diaphragms in some branches.
Bushy zoaria up to a foot across are seldom preserved intact, but the branches are
important constituents of some reefs, especially Ordovician. Several species are
index fossils. Global distribution, Ordovician to Devonian.
Hallucigenia: one of the famous genera of the Burgess Shale, now known also
from cousins in China and elsewhere; possibly related to modern velvet worms
(Onychophora) and to Collinsium ciliosum, a late Cambrian worm reported in 2015
(517 Ma), but considered by some a basal arthropod; Hallucigenia is notable for 14
spikes on its back, 14 limbs, and a bulbous head with two eyes and a mouth encir-
cled by a ring of small stiffening spikes; there are also teeth farther down its throat;
North America, Europe, and Asia; early to middle Cambrian.
Halysitidae: a family of chain corals in the order Tabulata; colonies ranged from 2
inches to several feet wide, with the corallites connecting to each other in a strik-
ingly chain-like pattern; global distribution, Early Ordovician to Early Devonian,
472–412 Ma.
Hamites: a paraphyletic genus of heteromorph ammonoids in the order Ammonitida
and suborder Ancyloceratina; named for the hook shape of most species, with a very
evolute, spirally shaped shell; a relatively weak swimmer because of the drag cre-
ated by the open shape; global distribution, Early to Late Cretaceous.
Hamulina: a genus of heteromorph ammonitid ammonoids in the family
Hamulinidae; it has an unusual crook-shaped shell; Caribbean and Europe, Early to
Late Cretaceous.
Hangenberg Event: an extinction event dated at about 359 Ma, at the end of the
Famennian, the final stage of the Devonian, characterized by widespread anoxia in
the oceans.
Haootia quadriformes: a species of early cnidarians, the earliest-known animal
with muscles; described in 2014 and dated at 560 Ma, its body has a four-sided
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s ymmetry and is composed of bundles of muscles stacked horizontally, with short

extensions from the corners; Newfoundland, Canada, Ediacaran.
Haplocheirus sollers: a species of avetheropod dinosaurs that seems to be the
earliest-known member of the lineage of dinosaurs that led to birds. Known from
only one specimen, it is the earliest and largest known of the superfamily
Alvarezsauroidea. About 7 feet long in the body, it had a long tail, powerful biceps
and forelimbs, and three long digits on the manus. Xinjiang, China; Late Jurassic,
160 Ma.
Haplohippus: a genus of horses in the still-extant family Equidae and the subfam-
ily Hyracotheriinae; a perissodactyl ungulate, Haplohippus was a forest browser;
North America, known from Oregon and Texas, late Eocene.
haploid: single or simple. Haploid cells are gamete (reproductive) cells that con-
tain one set of chromosomes rather than the two sets of diploid cells. In fertilization,
haploid gametes combine to make a diploid, producing a zygote, a fertilized ovum.
Haplolepis: a genus of small palaeoniscid actinopterygian fish in the family
Haplolepidae; like its relative Canobius, a smaller ray-finned fish of the Early
Carboniferous, it had vertical jaw suspension and a large gape; North America and
Europe, Late Carboniferous.
Haplorrhini: “simple nose,” a suborder of primates that includes humans, apes,
monkeys, and tarsiers, as well as their extinct primate ancestors; the Haplorrhini are
actually called “dry-nosed” in reference to their lacking a rhinarium, the moist area
around the nostrils of the Strepsirrhini and most other mammals; Paleocene to
Recent.
Haramyidae: an enigmatic family of proto-mammals of the Late Triassic.
Haramiyavia is a possible species based on jawbone and teeth fragments in a speci-
men of limestone from about 210 Ma, 30 to 40 million years earlier than the earliest
proven mammals. Research in 2015 seems to show that the haramyids were not true
mammals but part of a side-branch of mammaliaform animals that became extinct.
North America, Europe, Asia, and Africa; Triassic to Cretaceous.
hard parts: mineralized or otherwise hardened structures of an organism that fos-
silize easily, such as shells, bones, horns, bills, teeth, cuticles, claws, and scales.
Harpagornis moorei: Haast’s eagle, a large predator of New Zealand that became
extinct only 500 to 700 years ago, after the arrival of humans on the two islands.
Weighing 22 to 40 pounds, it had a 9-foot wingspan and was at least 30 percent
larger than the largest extant bird of prey. Genetic research indicates that it is most
closely related to a very small bird of prey, the two-pound Hieraaetus eagle of
Australia and New Guinea. The probable common ancestor of Haast’s eagle and
Hieraaetus is a two-pound species that lived between 1.8 and 0.7 Ma, so that Haast’s
eagle represents one of the most rapid instances of evolution from a very small to a
very large species. Before the arrival of humans, the only mammals in New Zealand
were three species of bats, but there were 250 species of birds. Harpagornis
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d ominated the food chain, hunting the 400-pound moas (Dinornithidae); both went
extinct within a century of the arrival of humans and accompanying rats, which
multiplied in the absence of natural predators and ate the eggs of many birds.
Harpetida: an order of trilobites in the subclass Librostoma; characterized by an
unusual brim around the cephalon; probably a feeding apparatus, the brim kept the
animal from enrolling tightly; also characterized by very small eyes and flat genal
prolongations; global distribution, late Cambrian to Late Devonian.
Harpoceras: a genus of ammonoids in the family Hildoceratidae; generally thin-
shelled and flattened laterally, the various species range widely in size, from 5
inches up to 18 inches wide; global distribution, Early to Late Jurassic.
harvestman: see Opiliones.
Hatzegopteryx: a genus of very large azhdarchid pterosaurs in the suborder
Pterodactyloidea, known from fragments found in Romania in 2002; estimated to be
as large as Quetzalcoatlus, to which it is similar, both of them having a wingspan of
about 40 feet; Late Cretaceous, 66 Ma.
Heidelberg man: see Homo heidelbergensis.
Helaletes: a genus of lophiodont tapirs in the superfamily Tapiroidea and extinct
family Helaletidae; derived from Homogalax and close to the ancestry of the mod-
ern tapir, Helaletes was small and agile, and looked like a small horse; North
America and Asia, early Eocene, 50–46 Ma.
Helcionelloida: an early Paleozoic class of small marine mollusks; characterized
by rugose, cap-shaped shells, conical and sometimes coiled; global distribution,
early Cambrian to Early Ordovician.
helens: a pair of long, curved, calcareous extensions often found connected to the
operculum of hyoliths; the function is unclear, perhaps as supporting structures;
named for C.D. Walcott’s wife and daughter.
Heliconiinae: a still-extant subfamily of butterflies in the superfamily Papilionoidea
and family Nymphalidae; famous for examples of Batesian and Müllerian mimicry;
Miocene to Recent.
Helicoplacus: a genus of early Cambrian echinoderms in the class Helicoplacoidea.
It is one of the earliest echinoderms but is represented by only two genera and very
few specimens. Helicoplacus was cigar-shaped and 2–3 inches high, with spirally
arranged plating on three ambulacra, different from other echinoderms in lacking
any radial symmetry. The only complete specimen was found in the White Mountains
of California, but fossil plates have been found across North America.
Helicoprion: a genus of eugeneodontiform holocephalian fish in the family
Agassizodontidae. Shark-like and estimated at 12–30 feet long, it is known only
from heterodont teeth and partial skulls since the cartilaginous body disintegrated
soon after death. Notable for its dentition, with a tooth-whorl as a cutting plate in
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the lower jaw and simple crushing teeth in the upper jaw; the tooth whorl contains
as many as 150 teeth. Global distribution, middle Permian, 285–270 Ma.
Heliobatis radians: “sun ray,” a genus of rays in the order Myliobatiformes and
family Dasyatidae; its generic name refers to the circular outline of its pectoral fin
discs; known from many specimens found in the Green River Formation, it was
generally 1–2 feet long; it had a long tail with numerous spines and up to three
stingers, and its diet was primarily small fish, crayfish, and mollusks; Wyoming,
United States, early Eocene, 54–48 Ma.
Heliolites: a genus of tabulate colonial corals in the family Heliolitidae; circular
corallites, smooth or surrounded by 12 oval segments; cosmopolitan, Middle
Ordovician to Middle Devonian.
Heliomedusa: a monospecific genus of early brachiopods in the family Disciniidae;
circular and less than an inch in diameter, H. orienta was first thought a jellyfish, but
it is now known from dozens of specimens in the Chengjiang biota that show the
lophophore and the growth lines of the shell; China, early Cambrian.
Hell Creek Formation: a very important North American formation that com-
prises strata from the end of the Cretaceous and beginning of the Paleocene, dated
roughly 70–66 Ma. With the Frenchman Formation of Saskatchewan, it extends
over a circular area 500 miles in diameter, centered in northeastern Montana. It is
best-known for dinosaur fossils, including the most complete hadrosaur skeleton
ever found. But it has also produced fossils of invertebrates, plants, fish, reptiles,
amphibians, and birds. Near its boundary with the overlying Fort Union Formation,
there are exposures of the iridium-enriched stratum that marks the end of the
Mesozoic Era.
Helodontidae: an enigmatic family of holocephalic fish, grouped with the
Chimaeriformes, possibly related to sharks; the type genus is Helodus; they are
characterized by a heterocercal tail fin and the absence of toothplates; instead they
have a series of flattened tooth elements in each jaw; Australia, North America and
Europe; Late Devonian to Permian.
Helohyidae: “marsh pig,” a family of small palaeodont artiodactyl mammals, sim-
ilar to the Raoellidae and possibly close to the ancestry of whales; Asia and North
America, late Eocene to late Oligocene.
Hemicidaris: a genus of echinoids in the order Hemicidaroida and family
Hemicidaridae; Europe, Africa, and North America; Middle Jurassic to Early
Cretaceous.
Hemicidaroida: a paraphyletic order of sea urchins, regular echinoids in the sub-
class Euechinoidea; cosmospolitan, Late Triassic to Late Cretaceous.
Hemicyclaspis: a genus of osteostracan fish in the order Atelaspidiformes, closely
related to Cephalaspis; 4–5 inches long, it is pleuronectiform and is characterized
by paired pectoral fins, a low dorsal fin, a row of low dorsal spines, a single nostril,
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and a bony headshield that was almost circular in cross-section; North America and
Europe, Early Devonian.
Hemicyoninae: a subfamily of carnivorous ursid mammals that probably includes
the ancestors of modern bears; the later species, such as Hemicyon, ranged from 4
to 7 feet long and were generally digitigrade; North America, Europe, and Asia; late
Oligocene to late Miocene.
Hemipneustes: a sea urchin, a genus of large irregular euechinoids in the order
Holasteroida and family Holasteridae; characterized by a high-domed test and a
deeply sunken anterior ambulacrum; the largest echinoid of the Cretaceous; Europe
and Asia, Early to Late Cretaceous.
Hemipsalodon: “semi-shearing teeth,” a genus of creodonts in the family
Hyaenodontidae; a scavenging carnivore, H. grandis was 6–7 feet long and weighed
about 250 pounds; the first and second molars were modified for slicing meat;
Western North America, late Eocene.
Hemizonida: an order of echinoderms in the class Asteroidea, primitive sea-stars;
North America and Europe, early Silurian to Late Carboniferous.
Heptodon: a genus of perissodactyl placental mammals in the superfamily
Tapiroidea and family Helaletidae; known in North America soon after the begin-
ning of the Eocene, it was about 3 feet long and closely resembled the modern tapir;
British Columbia, early Eocene, 50–49 Ma.
Hercynian orogeny: see Variscan orogeny.
Herefordshire Nodules: a late Silurian assemblage of carbonate nodules from a
volcanic ash stratum dated at 425 Ma, found in Western England in 1996. The nod-
ules contain exceptionally preserved calcitic fossils of organisms of a shallow epi-
continental sea, including brachiopods, polychaete worms, mollusks, crustaceans
(see Colymbosathon), and echinoderms. Since the fossils cannot be separated from
their matrix, many have been digitally recorded by a complex process of serial thin
sectioning that preserves fine detail.
Herpetotherium: a genus of marsupials in the family Herpetotheriidae, widespread
in North America from the early Eocene to early Miocene; Herpetotherium was a
small ground-dwelling omnivore, closely related to modern marsupials; North
America, Eocene to Miocene.
Herrerasauridae: a family of early saurischian dinosaurs; they are traditionally
considered theropods, but 2017 research argues that their similarities may be simply
the result of convergent evolution rather than phylogenetic relationship; the carnivo-
rous herrerasaurids are relatively small dinosaurs, generally less than 15 feet long;
South and North America, Late Triassic, 231–215 Ma.
Herrerasaurus: a genus of early saurischian dinosaurs in the carnivorous family
Herrerasauridae; the type species, H. ischigualastensis, is dated at 231 Ma; lightly
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built, its length is estimated at between 10 and 15 feet and its weight up to 700
pounds; Argentina, Late Triassic.
Hesperornis: the best-known member of the hesperornithine birds, a more
advanced early group than the enantiornithines; Hesperornis was a large, flightless,
diving sea-bird, almost 6 feet long; North America and Asia, Late Cretaceous,
85–75 Ma.
Hesslerides: a genus of trilobites in the family Proetidae; characterized by a large
glabella comprising most of the cephalon, with crescentic eyes set near the rear of
the cephalon; North America, Early Carboniferous.
Heteractinida: a probably paraphyletic Paleozoic order of sponges in the still-
extant class Calcarea with six- or eight-pronged calcareous spicules; global distri-
bution, early Cambrian to Carboniferous.
heterocercal: referring to an asymmetrical tail fin, generally with the upper part
extending farther than the lower part, seen especially in sharks; on the other hand,
the Anaspida and mosasaurs are characterized by a reversed heterocercal tail fin.
heterochrony: a genetic shift in the evolutionary development of tissue or an ana-
tomical part, causing it to develop at an abnormal time relative to an ancestor; the
change can then be inherited by the offspring; for example, the adults of some later
species of the Jurassic bivalve genus Gryphaea exhibit traits that were juvenile in an
earlier species.
Heterocorallia: an order of late Paleozoic corals in the subclass Zoantharia; it has
one family, the Heterophyllidae; solitary and elongate, possibly related to the
Rugosa but with a different arrangement of septa; global distribution, Middle
Devonian to Early Carboniferous.
Heterocrania rhyniensis: a species of arthropods in the subclass Euthycarcinoidea,
q.v.; Scotland, Early Devonian.
heterodont dentition: having more than one type of teeth, evidence of specializa-
tion in feeding.
Heterodontosauridae: a basal family of ornithischian dinosaurs; a long-lived and
widespread group known from sparse specimens on all continents except Australia
and Antarctica; they were as long as 6 feet, including a long tail, and are character-
ized by long, grasping forelimbs; also characterized by their heterodont dentition
(the trait they were named for), having three different types of teeth; probably her-
bivorous but may also have preyed on small animals; only Heterodontosaurus is
known from a complete skeleton, but see also Abrictosaurus; Late Triassic to Early
Cretaceous.
Heterodontosaurus: a monospecific genus of basal ornithopod dinosaurs; known
from several specimens but primarily from an articulated skull and skeleton dated
between 210 and 190 Ma; it was up to 6 feet long, had three different types of teeth,
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and had five digits on the manus and four on the pedes; South Africa and North and
South America, Late Triassic to Early Jurassic.
Heteromyidae: a still-extant family of rodents in the superfamily Geomyoidea;
widespread in North America in the Oligocene, the family’s range has shrunk, but
the kangaroo mouse and kangaroo rat are two of many surviving genera; North and
Central America, Oligocene to Recent.
Heteronectes: a genus of flatfish showing a transitional stage in the development
of flatfish asymmetry; known only from Italy, Eocene; see also flatfish.
Heterosoricinae: a proposed subfamily of insectivores in the family Soricidae (the
shrews); North America, Europe, Africa, and Asia; Eocene to Pliocene.
heterospory: the production of two types of spores differing in size and sex;
known in several types of terrestrial plants such as Archaeopteris and the club-
mosses in the Devonian.
Heterostraci: “different shield,” a diverse subclass of jawless, armored fish-like
vertebrates; generally small (5–9 inches) and benthic, they evolved during the time
when the mineralized skeleton first appears in the fossil record; the group includes
principally the cyathaspids and their descendants the pteraspids; the family
Psammosteidae was the last of the heterostracans, extinguished at the end of the
Devonian; global distribution, late Cambrian to Late Devonian.
Hexactinellida: a class of glass sponges, characterized by 4-, 5-, and 6-rayed sili-
ceous spicules; formerly called Hyalospongiae; more common as fossils than demo-
sponges because their siliceous spicules fossilize easily; global distribution,
Cambrian to Recent.
Hexanchiformes: a still-extant order of primitive sharks in the superorder
Squalomorphii; characterized by 6 or 7 gills, one dorsal fin, and a heterodont denti-
tion; well-known from the Middle Jurassic through the Cenozoic, but teeth resem-
bling those of hexanchiforms are known as far back as the Devonian; the extant
families are the Hexanchidae and the Chlamydoselachidae, and extinct families are
the Crassonotidae, Mcmurdodontidae, and Orthacodontidae; Devonian? to Recent.
Hexapoda: a subphylum of arthropods that includes true insects and the entogna-
thous proto-insects: the Collembola (springtails), Protura, and Diplura; the group
has also traditionally included myriapods, but genetic evidence may exclude them;
global distribution, Early Devonian to Recent.
Hexathelidae: see Rosamygale.
Heymonsicambria scandica: one of several important discoveries of fossil pen-
tastomes showing that the parasitic Pentastomida (q.v.), tongue-shaped worms,
date back to the Cambrian; Canada, late Cambrian.
Hiemolora: a poorly known genus of Ediacaran fossils, known at least from the
Mistaken Point fauna and possibly elsewhere; it is a tentacular medusoid, with 12–20
arms radiating from a 2-inch circular body; Canada, Ediacaran, 575–560 Ma.
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Hildoceras: a genus of ammonoids in the order Ammonitida and family

Hildoceratidae; characterized by a narrow, discoidal, and evolute shell; Hildoceras
is named for St. Hilda, the founding abbess of Whitby Abbey in Yorkshire, England,
who legendarily turned all the snakes into stone, creating the ammonoids; Europe
and Japan, Middle Jurassic.
Hindeodus parvus: a species of conodonts in the family Anchignathodontidae; its
first appearance marks the beginning of the Mesozoic Era and the Triassic Period at
252.17 Ma; China.
Hippuritoida: the rudists, an order of euryhaline bivalve mollusks in the class
Heterodonta; flourishing in the warm, saline water of the Cretaceous, they became
the basis of the most prominent reefs of the period, cementing themselves together
in tropical reefs several hundred feet high, running for hundreds of miles along
some Cretaceous coastlines; the individuals took various tube and conical shapes,
the latter sometimes as long as 40 inches; see for example Radiolites; cosmopolitan,
Late Jurassic to the end of the Cretaceous.
holarctic: referring to the Holarctic Region, the temperate areas of Europe, Asia,
North America, and to North Africa, from the Jurassic to the present.
holaspis: the final ontogenetic phase of trilobites, followed by several molts that
merely increased body size and added segments; see also meraspis and protaspis.
Holasteroida: an order of Mesozoic sea urchins, heart-shaped atelostomate echi-
noderms; several species are important in biostratigraphy, with a few genera still
extant; Cretaceous to Recent.
Holectypidae: a family of irregular echinoids in the order Holectypoida; the group
includes some of the earliest and most primitive irregular echinoids, similar to the
Pygasteridae; global distribution, Early Jurassic to Late Cretaceous.
Hollinoidea: a superfamily of dimorphic paleocopid ostracods in the extinct order
Beyrichicopina; some forms with sulci and lobation; North America and Europe,
Holocene: the most recent epoch of the Cenozoic Era, the last 11,700 years.
Holocephali: a still-extant subclass of cartilaginous fish in the class Chondrichthyes;
known as far back as the Devonian but only from scattered teeth; the only surviving
order is the Chimaeriformes, the ghost sharks; Devonian to Recent.
holochroal: the type of compound eye in most trilobites, with thousands of tightly
packed lenses covered by a single cornea.
Holoptychiidae: a family of sarcopterygian fish in the order Porolepiformes; these
lobe-finned fish included at least one genus that may have been amphibian,
Laccognathus; North America and Europe, Middle Devonian to Early Pennsylvanian.
Holothuroidea: the sea cucumbers, a still-extant class of echinoderms in the sub-
phylum Echinozoa. Their body is generally cylindrical, with five ambulacra and a
leathery skin which contains ossicles of different types and sizes. They are generally
130 H
6–12 inches long, but the longest known (Synapta maculata) grows to 10 feet. The
fossil record is poor since there is no integrated skeleton, but ossicles and elements
of the calcareous ring around the mouth are known from as early as the Silurian, and
body specimens are known from the Devonian and later. Global distribution,
Silurian to Recent.
holotype: a single physical specimen of the type species of a taxon, designated as
the defining type of the taxon at the time when it is formally described and estab-
lished as a formal group; the holotype anchors specific physical characters to the
name of the taxon.
Holuridae: a family of palaeonisciform chondrostean fish; characterized by an
ossified exoskeleton and robust rhombic scales; Europe, Carboniferous to Permian.
Holzmaden Shale: a Lower Jurassic Konservat-Lagerstätte in Southern Germany,
where several limestone and shale quarries have produced exceptionally preserved
specimens of shallow-sea taxa from the Lower Jurassic Posidonia Shale, dated at
185–183 Ma. Of the vertebrate fossils, the most common are the ichthyosaurs,
including the first to clearly show Stenopterygius with a dorsal fin and an upper lobe
of the tail. Many of these specimens show stomach contents, and there is a high
percentage of juveniles and females with unborn fetuses, indicating that the area
was probably a spawning ground. Other vertebrates include teleosaurid crocodiles
(see Steneosaurus), pterosaurs, and fish. Cephalopods such as the ammonoid
Harpoceras are common, and other invertebrates include crinoids and bivalves. One
of four Lagerstätten dated near 183 Ma; see also Strawberry Bank and Ya Ha
Tinda.
Homacodontidae: a paraphyletic family of palaeodont mammals in the superfam-
ily Dichobunoidea; among the earliest and most generalized artiodactyls; North
America, Asia, and Europe; Eocene.
Homalozoa: a controversial group of early Paleozoic animals traditionally consid-
ered a subphylum of echinoderms; some consider them a separate class called
Carpoidea; also called calcichordates or stylophorans by some, they may even have
an evolutionary link with chordates; characterized by a calyx with a short stem,
projecting arm and tail-like structures, and a lack of any symmetry; global distribu-
tion, middle Cambrian to Early Devonian.
hominid: term referring to the family Hominidae, which includes all modern great
apes (i.e., modern humans, gorillas, chimpanzees, and orangutans) and their imme-
diate ancestors; early Miocene to Recent.
hominin: term referring only to modern humans, extinct members of the genus
Homo, and their immediate ancestors, such as Australopithecus, Paranthropus, and
Ardipithecus; late Miocene to Recent.
Hominoidea: the apes, a superfamily of haplorrhine primates consisting of the
families Hominidae and Hylobatidae; Oligocene to Recent.
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Homo erectus: a species of primates more advanced in several ways than H. habi-
lis and probably ancestral to H. sapiens. Remains of H. erectus have been found in
Java, China, Africa, Europe, and Western Asia, some as old as 1.75 million years
and some possibly as young as 70,000 years ago. Homo erectus was on average
about 5 feet tall and had a cranial capacity slightly less than H. sapiens. Although it
was long thought to be the first hominin to walk upright, other species, notably Au.
afarensis, now seem to have done so as early as 3.4 Ma. A find of a jawbone and two
skullcaps found in the Republic of Georgia in the late 1990s may be H. erectus, and
its age is thought to be about 1.7 Ma; it is similar to a jawbone from Kenya from
1.6 Ma and is possible evidence of early migration of the species from Africa. Some
authorities now divide H. erectus into three separate species in Africa, Europe, and
Asia. See also Denisovans, Neanderthals, and Ngandong fauna.
Homo erectus pekinensis: a form of H. erectus characterized by a massive brow
ridge and a cranial capacity of 900–1200 cc; tools and the remains of modern
humans have also been found at the site; Northern China, Pleistocene, about
750,000 years ago.
Homo ergaster: the group of hominins considered by some to be the branch of H.
erectus that gave rise to H. sapiens in Africa; see also Turkana Boy; southeastern
Africa, 1.9 to 1.3 Ma.
Homo floresiensis: a species of hominins known from partial skeletons of nine
individuals found on the Indonesian island of Flores; between 2 and 4 feet tall, the
specimens have generated intense debate, some claiming that a disease caused their
diminutive size; the emerging consensus, however, is that they do represent a sepa-
rate species; dated between 200,000 and 50,000 years ago.
Homo habilis: possibly the earliest-known representative of its genus, H. habilis
lived from about 2 million years ago to about 1.75 million years ago, with most
specimens found in Africa and Asia; possibly ancestral to H. erectus, with a cranial
capacity (around 650–700 cc), slightly more than that of the australopithecines but
less than that of H. erectus.
Homo heidelbergensis: a disputed species of hominins; considered by some to
represent the branch of H. erectus ancestral to Neanderthals and modern humans;
Europe, Africa, and possibly Asia; 600,000 to 120,000 years ago.
Homoiostelea: a controversial class of early biradiate echinoderms, generally
placed in the subphylum Homalozoa; characterized by a feeding arm and a stem
consisting of a series of plates; cosmopolitan, Cambrian to Devonian.
homologous structures: body parts of different organisms that have anatomical
features in common but that perform different functions. For example, bats’ wings
and human hands have homologous bone structures, but only bats can fly. The
hindlimbs of whales may seem homologous with those of hippopotamuses; but
since they are no longer functional, the whales’ hindlimbs are considered vestigial.
See also analogous structures and vestigial structures.
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Homo naledi: the name given to a species of hominins known only from a deposit
of bones in a cave north of Johannesburg, South Africa, probably a burial site. Found
in loose soil and unaccompanied by other fauna, they are difficult to date, with esti-
mates ranging from 2.5 to 0.9 Ma. The bones represent parts of at least 15 individu-
als, and there are multiple examples of almost every bone in the body, so that the
skeleton constructed from them is more complete than any previously known hom-
inin specimen. This discovery seems to be an early stage of hominin development,
having a brain the size of an orange but also having other more advanced
characteristics.
Homo neanderthalensis: see Neanderthals.
Homo rhodesiensis: the name given to several pre-modern hominin specimens
found in the twentieth century in East and North Africa; dated between 400,000 and
120,000 years ago, the group is thought by some to be an African version of H.
heidelbergensis and probably in the direct line of descent to H. sapiens; whether it
should be called H. sapiens rhodesiensis is in question.
Homo sapiens sapiens: modern humans, the only surviving member of the genus
Homo; see also Cro-Magnon, Denisovans, and Neanderthal; middle Pleistocene
to Recent.
Homotherium: a genus of large felids in the subfamily Machairodontinae. Its
canines were longer than those of Machairodus but shorter than some other saber-
toothed cats. Homotherium was about the size of a modern male lion, but its elon-
gated forelimbs and short tail give it a hyena-like appearance. Its last representatives
seem to have been those in Europe, which disappeared about 28,000 years ago.
Global distribution except Australia and Antarctica, Pliocene to Pleistocene.
honeycomb coral: see Tabulata.
Hoplitoplacenticeras: “heavily armed Placenticeras,” a Late Cretaceous ammo-
noid in the order Ammonitida; its shell can be 3 feet across and so strongly ribbed
that it has a spiky appearance; important stratigraphically in the Campanian, it is the
latest representative of the Placenteratidae, found worldwide into the Maastrichtian.
Hoplopteryx: a genus of teleost actinopterygian fish in the still-extant family
Trachichthyidae; almost a foot long, it had relatively small fins, but the dorsal fin
was supported by a row of strong spines; in an evolutionary advance, a series of
uroneural bones in the tail supported the fin rays and made Hoplopteryx a strong
swimmer; related to the modern roughy; global distribution, Late Cretaceous,
90–70 Ma.
horizon: in paleopedology, a layer of soil; the upper layers lie generally within
1.5 feet of the surface (labeled horizons A and E) and the subsurface horizon lies
within 7 feet of the surface (labeled horizon B).
horned dinosaur: see Ceratopsia.
horseshoe crab: see Limulus.
H 133
House Range: see Wheeler Shale.

Hunsrück Slate: Several quarries in this Konservat-Lagerstätte of the Early
Devonian (about 407 Ma) contain well-preserved soft-tissue pyritization fossils.
Important examples include fish (especially Drepanaspis and Gemündina), echino-
derms (crinoids, asteroids, and ophiuroids), mollusks, and several arthropods (espe-
cially the malacostracan crustacean Nahecaris and the pycnogonid sea-spider
Palaeoisopus). Germany, Early Devonian, Pragian to Emsian Stages.
Hurdia: a genus of stem-arthropods in the order Radiodonta. Known first from the
Burgess Shale assemblage and at first misidentified, it was correctly described in
2009 and seen to be related to Anomalocaris. Between 8 and 20 inches long, it is
characterized especially by large gills attached to its sides and by a thin, pointed
carapace extending forward. North America, Europe, and Asia; middle Cambrian.
Hyaenodon: a short-lived but diverse genus of creodonts in the extinct family
Hyaenodontidae; the largest species, H. gigas, was about 10 feet long and weighed
half a ton; Hyaenodon was cat-like and walked on its toes; not a direct ancestor of
the modern hyenas; North America, Europe, Africa, and Asia; late Eocene to early
Miocene.
Hyalospongiae: see Hexactinellida.
Hybocrinida: an order of early crinoids in the polyphyletic class Inadunata; char-
acterized by a monocyclic cup and circular stem; North America and Europe, Late
Ordovician and possibly early Silurian.
Hybodontiformes: “hump tooth,” a long-lived suborder of euselachian sharks,
named for their compressed conical teeth; dominant Mesozoic predators, hybodonts
were 6–7 feet long; prominent genera include Hybodon and Ptychodus, the type
genera of the large families Hybodontidae and Ptychodontidae; global distribution,
Early Carboniferous to Late Cretaceous.
Hydnoceras: a genus of glass sponges in the class Hexactinellida; horn-shaped and
about 8 inches long, it is an index fossil in the Late Devonian; North America and
Europe, Devonian to Mississippian.
Hydrodamalis gigas: Steller’s sea cow, a genus of sirenians in the family
Dugongidae; as long as 30 feet, it was toothless and fed entirely on seaweed; with a
very thick layer of blubber, it may have been unable to dive; hunted to extinction in
the middle of the eighteenth century; North Pacific Ocean, Pliocene to Holocene.
Hydrozoa: a class of cnidarians in the subphylum Medusozoa. Mostly soft-bodied,
predatory marine animals, solitary or colonial, they include the Portuguese man-of-
war but are separate from the jellyfish (see Scyphozoa). They probably date back to
the Precambrian, but fossils are rare. Three groups — fire corals, milleporines, and
stylasterines — have a stony growth that fossilizes; they appeared in the Cretaceous
and are widespread through the Cenozoic.
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Hyeniales: an order of primitive shrubs with short, forked leaves in whorls; incer-
tae sedis between sphenophytes and pteridophytes; global distribution, Devonian.
Hylobatidae: the gibbons, classified with humans and the Cercopithecidae in the
superfamily Hominoidea and infraorder Catarrhini; Southeast Asia, late Miocene to
Recent.
Hylonomus: a monospecific genus of anapsid reptiles, placed tentatively in the
Romeriida; the earliest-known unquestioned reptile, known from Nova Scotia about
312 Ma, H. lyelli was 9 inches long and had a long tail; Canada, Middle
Pennsylvanian.
Hymenoptera: an order of insects that includes bees, wasps, and ants; global dis-
tribution, Early Triassic to Recent.
Hyolitha: “u-bone,” a group of enigmatic Paleozoic animals, long considered
probable mollusks but in 2017 shown to be lophophorates, related to brachiopods.
Up to 1.5 inches long, they are similar in some ways to annelids, the segmented
worms. They have conical calcareous shells with an operculum closure at the large
end and two curved extensions called helens protruding, along with short feeding
tentacles. The shell generally appears triangular or u-shaped when crushed flat.
There are two orders, the Hyolithida and the problematic smaller group Orthothecida.
Widespread in the Cambrian, their numbers declined through the Devonian, but a
few persisted to the end of the Permian.
Hyopsodontidae: a family of small, primitive ungulate mammals, possibly related
to horses; dachshund-like in appearance, they were generally insectivorous and her-
bivorous; they were unspecialized, having clawed toes and a generalized dentition;
North and South America, Europe, Asia, and Africa; early Paleocene to Eocene.
Hypacrosaurus: a duospecific genus of large hadrosaurid dinosaurs in the subfam-
ily Lambeosaurinae. Similar to Corythosaurus but with a higher and wider crest and
with very high neural spines, for which one species is named H. altispinus. The
other species, H. stebingeri, is especially important for the discovery of a great
number of hatchlings and embryos in eggs the size of a small soccer ball. Research
on the embryos has shown, among other things, that their relatively long incubation
time was almost six months, much longer than in birds. Two Medicine Formation,
North America, Late Cretaceous, 86–71 Ma.
Hypertragulidae: a family of small artiodactyl ungulate mammals, among the
earliest ruminants; weighing only 5–9 pounds, they resemble very small deer;
Europe, Asia, and North America; Eocene to Miocene.
Hyphalosaurus: a genus of choristoderan reptiles in the family Hyphalosauridae.
Very common in the Jehol biota, it reached a length of almost 3 feet, with a very
long neck and tail. Some specimens show skin impressions, and Hyphalosaurus
may have been viviparous. China, Early Cretaceous, about 125 Ma.
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Hyphantoceras: a genus of heteromorph ammonitid ammonoids in the superfam-

ily Turrilitaceae; it has a loose, helically coiled shell that stretches out to 4–5 inches
long; cosmopolitan, Late Cretaceous.
Hypochilidae: a still-extant family of araneomorph spiders in the superfamily
Paleocribellatae; there are no definitive known fossil specimens, but the body plan
of the modern lamp-shade spider Hypochilus seems to be a mosaic of primitive and
derived characters; found only in China and North America.
Hypsilophodon foxii: a species of herbivorous ornithischian dinosaurs in the sub-
order Ornithopoda, the only generally accepted hypsilophodont. Known only from
several specimens found on the Isle of Wight in the nineteenth century, H. foxii was
5–6 feet long and lightly built, weighing less than 50 pounds; it had a stiff tail, five
long fingers on weak forelimbs, and four toes on its hindfeet. Its name refers to a
frill along its spine, not to its dentition. England, Late Jurassic to Early Cretaceous.
Hypsilophodontidae: a large, probably paraphyletic family of small ornithischian
dinosaurs in the suborder Ornithopoda; from 3 to 6 feet long and lightly built, gener-
ally bipedal; global distribution, Middle Jurassic to Late Cretaceous.
hypsodont dentition: high-crowned teeth with a flattened surface for chewing
tough vegetation, with enamel extending the length of the tooth and past the gum
line, and with cementum between the lophs of the tooth.
Hyrachyus: a genus of ceratomorph mammals in the family Hyrachyidae; 4–5 feet
long and related to the Lophiodontidae, it is known from hundreds of specimens; it
is generally considered to be related to the tapirs but has rhinoceros-like dentition;
holarctic distribution, Eocene.
Hyracodontidae: a family of perissodactyl ungulates in the superfamily
Rhinocerotoidea; initially small and tapir-like, the later hyracodontids include the
largest known terrestrial mammals, the Indricotherinae (see Paraceratherium);
North America, Europe, and Asia; early Eocene to early Miocene.
Hyracoidea: a still-extant order of superficially rabbit-like mammals; in the Eocene
the hyracoids ranged from cow-sized to mouse-sized and were widespread in Africa;
related to sirenians and elephants; Africa, Europe, and Asia, early Eocene to Recent.
Hyracotherium: a genus of palaeotheres in the family Palaeotheriidae, close to the
ancestry of Equidae and brontotheres. It had four toes on its forelimbs and three toes
on its hindlimbs but is considered the earliest-known perissodactyl. Its teeth were low-
crowned and it weighed up to 50 pounds. Hyracotherium had a global distribution in
the early Eocene, as evidenced by whole fossil skeletons found in Europe and North
America and partial ones elsewhere. See also Eohippus. Early to middle Eocene.
Hystricospherida: an enigmatic group of spiky and spherical protist microfossils,
one-celled marine algae; not well-known but possibly related to the zygotes of
freshwater chlorophytes or to dinoflagellates; global distribution, Paleozoic to
Recent.
I
Iapetognathus fluctivagus: a species of denticulate conodont elements in the fam-

ily Cordylodontidae; its first appearance (currently considered to be 485.4 Ma)
marks the GSSP for the beginning of the Early Ordovician in Newfoundland,
Canada; the species is also known from the United States, Asia, and Australia.
Iberomesornis: see Enantiornithiformes.
Ibyka: a poorly known genus of early plants tentatively placed in the order
Iridopteridales; Middle to Late Devonian.
Icaronycteris: a genus of early microchiropteran bats in the family Icaronycteridae;
the earliest-known bat, it was about 6 inches long, had a wingspan of 15 inches, and
probably used echolocation; North America, Asia, and possibly Europe; early
Eocene.
ICBN: International Code of Botanical Nomenclature; in 2011 the Eighteenth
International Botanical Congress changed the name to International Code of
Nomenclature for algae, fungi, and plants.
icehouse conditions: periods of extensive cooling on Earth, referring to periods of
lower global temperatures and glaciation. They are associated with low levels of
carbon dioxide, relatively calm volcanic eruptions, and low sea levels. The most
recent, the Pleistocene glaciations, are well-documented, but the earlier cool peri-
ods are not. The Karoo Ice Age was a period of cooling toward the end of the
Carboniferous, extending into the early Permian; evidence for it has been found
especially in Southern Africa and South America, but its extent and effects are not
well-known. A cool period at the end of the Ordovician, roughly 450–420 Ma, is
sometimes referred to as the Andean-Saharan Glaciation. Two longer periods of
more severe cooling occurred in the Cryogenian Period, before the Ediacaran. Dated
at 650–635 Ma, the Marinoan Glaciation ended the Cryogenian after a warm span
of about 25 million years; and the long Sturtian Glaciation, from 717 to 660 Ma,
marked the first part of the Cryogenian. Another long stretch of global cooling,

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probably the most severe but least understood, is called the Huronian Glaciation,
occurring between 2400 and 2100 Ma.
ichnology: see paleoichnology.
Ichthyodectidae: “fish biters,” a family of Mesozoic teleost actinopterygian fish in
the still-extant superorder Osteoglossomorpha and extinct order Ichthyodectiformes;
most were between 3 and 16 feet long, and they preyed on smaller fish; Late Jurassic
to Late Cretaceous.
Ichthyolestes: a genus of archaeocete whales in the family Pakicetidae, fish-eating
but mainly terrestrial; Pakistan, early to middle Eocene.
Ichthyopterygia: a superorder or subclass of diapsid marine reptiles including
ichthyosaurs and their smaller ancestors; predatory and strong swimmers, they had
a relatively short neck and porpoise-like body; Europe and Asia, Early Triassic to
Late Cretaceous.
Ichthyornis: a genus of early toothed birds in the order Ichthyornithiformes, dated
between 95 and 84 Ma; notable for its rows of sharp teeth, similar to those of mosa-
saurs, and for its vertebrae, which are fish-like, concave on both sides; strongly
keeled, it was probably a good flyer; similar to a small tern, it is closer to modern
birds than are more archaic groups like the enantiornithines; North America, Late
Cretaceous.
Ichthyosauria: an order of specialized diapsid marine reptiles in the superorder
Ichthyopterygia, characterized by an elongated snout and a fish-shaped body; their
ancestors may have been diapsids that returned to the sea; ichthyosaurs ranged in
length from 3 feet to more than 50 feet, and they were viviparous; Early Triassic to
Late Cretaceous.
Ichthyostega: a genus of primitive labyrinthodont amphibians in the clade
Tetrapodomorpha; close to the early ancestry of all terrestrial vertebrates and among
the first to adopt an amphibian habit; its pedes had 7 digits, but a fossil manus has
not been found; the skull was broad and flattened, and some species attained lengths
of 3 feet; see Tetrapoda; North America and Europe, Late Devonian.
ICSN: International Code of Stratigraphic Nomenclature, the generally accepted
authority for the names assigned to rock strata.
Ictidosauria: see Tritheledontidae.
ICZN: International Code of Zoological Nomenclature, the generally accepted
authority for animal nomenclature; the current 4th edition was issued in 1999, and
a 5th edition is planned for 2018.
Iguanodon: a genus of bulky ornithischian dinosaurs in the suborder Ornithopoda;
one of the largest animals of its time, the herbivorous Iguanodon ranged from 30 to
40 feet long and weighed about 3 tons; with long arms, it could shift easily from
quadrupedality to bipedality; the genus has seen many revisions and reassignments
I 139
of species, but I. bernissartensis, one of the earliest dinosaur discoveries, is the type
species; its manus is distinctive, with a prominent thumb spike and a prehensile fifth
finger; it was named for its set of iguana-like teeth; Europe, North America, and
probably Asia and Africa; Early Cretaceous, 126–125 Ma.
Illaenus: a genus of medium-sized isopygous trilobites in the family Illaenidae
and order Corynexochida; Illaenus grew as large as 2 inches, its cephalon had a high
profile, and it had two short and recurved genal spines; the Illaenidae persisted from
the Early Ordovician to the late Silurian; Europe and North Africa, Middle
Ordovician.
impunctate: see punctate.
Inadunata: a class or subclass of stalked crinoids that recent research identifies as
polyphyletic; the members of its three orders (Cladida, Disparida, and Hybocrinida)
are being reassigned; global distribution, Early Ordovician to Middle Triassic.
inarticulate: see articulate.
incertae sedis: of uncertain classification.
Incisoscutum ritchiei: see Gogo Formation.
index fossil: a morphologically distinctive fossil taxon that characterizes a particu-
lar time period or biozone, usually found in a limited number of strata; it is abundant
and widespread but short-lived, so that its presence in a stratum makes it possible to
correlate widespread occurrences of the stratum; also referred to as guide fossil or
zone fossil.
Indocetus: a genus of archaeocete cetaceans in the family Protocetidae, one of the
earliest ancestral whales. Its four sacral vertebrae are fused to each other and to its
pelvis, so that it may have been able to support its weight on land. It probably used
its hindlimbs rather than its tail in swimming. India, early Eocene.
Indohyus: a genus of artiodactyl mammals in the family Raoellidae; Indohyus was
deer-like in shape and about the size of a raccoon; several distinctive features show
that it may be closely related to whales; India, middle Eocene.
Indricotherium: see Paraceratherium.
infaunal: see epifaunal.
Iniopterygiformes: an unusual order of holocephalic fish in the class
Chondrichthyes, distantly related to the modern Chimaeriformes, the ghost sharks;
they ranged from 6 to 18 inches long and had a cartilaginous skeleton; they are
named “nape wing” for the distinctive pair of wing-like fins high on their back; a
specimen from Kansas was found in 2009 to have a fossilized brain inside its intact
skull; North America, Devonian to Carboniferous.
Inkayacu paracasensis: a species of large penguins that lived in Peru in the late
Eocene; very similar to modern penguins except that it was about five feet tall,
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weighed about 120 pounds, and had a long beak; the specimen found in the Paracas
National Reserve in Peru in 2009 is also notable for having well-preserved feathers
that may have been brown and gray.
Inoceramus: a very large genus of deeply ribbed bivalves in the order Praecardioida
and subclass Cryptodonta; some species were over 3 feet in diameter; the inocera-
mids were declining gradually for at least 10 million years before disappearing
about 1.5 million years before the end of the Cretaceous; global distribution, Early
Jurassic to Late Cretaceous.
Insecta: a class of arthropods in the subphylum Hexapoda; the largest class of
animals, with almost a million named species. Insects’ most basic apomorphies are
a chitinous exoskeleton, six jointed legs, and compound eyes. The only inverte-
brates that have achieved flight, most groups of insects have four wings presently or
had them at an earlier stage of their evolution (see also Pterygota, Diptera and
Coleoptera). Insects first appeared in the Early Devonian, but their fossil record is
very sparse through the Devonian and Carboniferous. The oldest known insect is a
springtail from the Devonian Rhynie Chert of Scotland (see Rhyniognatha), and
the first definitive flying insects appeared in the Late Carboniferous (see Ensifera).
Insectivora: an abandoned name for a grouping of the many small mammals that
became widespread in the Cretaceous, preying mainly on insects; see Eulipotyphla.
invertebrate: an informal term for all non-vertebrate metazoan animals.
involute: especially of ammonoids, tightly coiled, with successive whorls overlap-
ping and often covering most of the surface of earlier ones.
Iridopteridales: one of the widespread but enigmatic orders of early vascular
plants called cladoxylopsids; the whole-plant concept is unclear because its trunk
and branch connections are not known; global distribution, Middle to Late Devonian.
Isanosaurus: a monospecific genus of sauropod saurischian dinosaurs; I. attavipa-
chi is important as one of the earliest sauropods, but it is poorly known from frag-
ments of one skeleton, dated at about 210 Ma; Thailand, Early Triassic.
Isastraea: a genus of scleractinian corals in the subclass Hexacorallia; it lived in
warm, clear, shallow water; North America, Europe, and Africa; Jurassic to Cretaceous.
Ischadites: a contentious genus of benthic marine organisms currently classified as
algae in the obscure family Receptaculitidae; its globose body resembles a sponge,
with spiral rows of interlocking plates; North America and Europe, Middle
Ordovician to Early Devonian.
Ischnacanthiformes: an order of acanthodian fish; they were slender, lightly
armored nektic predators, characterized by two dorsal fins and stout spines; larger
than other acanthodians, the larger species grew as long as 6 feet; North America
and Europe, late Silurian to Carboniferous.
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Ischyromyidae: a basal family of sciuromorph rodents; known from France to

North America through the Eocene, they gave rise to the Gliridae and other modern
rodents; probably scansorial and probably competitive with early scansorial p rimates
(see also Paramyidae); North America, Europe, and Asia; Paleocene to Oligocene.
Isectolophidae: a family of perissodactyl ceratomorph mammals in the superfam-
ily Tapiroidea; probably descended, along with the Helaletidae, from Homogalax,
an early Eocene tapiroid of Asia and North America; also related to the ancestral
horse Hyracotherium; North America, Europe, and Asia; early to late Eocene.
Isisaurus: a genus of titanosaurian sauropodomorph dinosaurs; about 60 feet long
and 20 feet high, it is characterized by unusually long forelimbs and a thick, verti-
cally directed neck; probably a high browser; India, Late Cretaceous, 71–69 Ma.
island dwarfism: a process resulting in reduced size in a population of animals
isolated on islands from other populations of the same species; see Bubalus,
Europasaurus, and Homo floresiensis.
Isoptera: the termites, a suborder of insects in the order Blattodea, q.v.
isopygous: of trilobites, having a cephalon and pygidium of approximately the
same size.
Isotelus rex: the largest known trilobite, a species in the order Asaphida and family
Asaphidae; characterized by its isopygous shape and its size: 28 inches long, 3
inches high, and 16 inches wide; North America, Middle to Late Ordovician.
Itagnostus: a genus of agnostid trilobites in the family Peronopsidae; isopygous
and in some species with a thorax narrower than the cephalon and pygidium; known
only from Marble Canyon in British Columbia; Canada, middle Cambrian.
Ixodida: the ticks, an order of arachnids, divided into two main families, the
Ixodidae and the Argasidae; poorly known but in the fossil record at least since
99 Ma; they have thrived along with browsing mammals since the spread of grass-
lands in the Eocene; also called Metastigmata; North America and Asia, Cretaceous
to Recent.
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Jamoytius kerwoodi: a lamprey-like jawless fish with long paired fins running the
length of its body; formerly considered an anaspid ostracoderm, its classification is
still in dispute; it is now tentatively placed in the disputed class Hyperoartia;
Scotland, known only from the Silurian.
Janassa bituminosa: the type species of the family Janassidae, poorly known car-
tilaginous fish in the family Petalodontidae; known primarily from teeth; North
America and Europe, Mississippian to Permian.
Janjucetus: a stem mysticete whale in the family Mammalodontidae; about 11 feet
long, it is notable for its short skull, its large eyes, and especially its large teeth; it
had no baleen, a principal character of modern mysticetes; Australia, known only
from one well-preserved specimen of the late Oligocene.
Java Man: a set of fossil bones discovered at Trinil, Java, about 60 miles from
Ngandong, where the Solo Man remains were found; first classified as
Pithecanthropus or a sub-genus Javanthropus, it is now considered Homo erectus;
Pleistocene, about 1.5 million years ago.
Jehol Biota: “hot river,” a fossil community seen in several Konservat-Lagerstätten
of Liaoning Province in China, found in the Lower Cretaceous Jiufotang and Yixian
Formations, an area known for numerous hot springs. The species are dated between
133 and 110 Ma, the dates sometimes being uncertain because of problems dating
the hundreds of fine-grained volcanic lake deposits that have preserved very detailed
structures in the fossils; most of the important fossils are dated between 130 and
120 Ma, in the Yixian Formation. The most striking are a range of feathered dino-
saurs such as Jeholornis and Sinosauropteryx and protobirds such as Confuciusornis,
but there are also significant mammals (see Repenomamus and Eomaia), reptiles
(see Hyphalosaurus), insects, and some of the earliest angiosperms. See also
Caudipteryx, Protarchaeopteryx, Beipiaosaurus, and Sinornithosaurus.

144 J
Jeholornis: one of the many bird-like feathered dinosaurs found in China since the
1970s. Like Archaeopteryx and other proto-birds, it is notable for its long, bony tail
with feathers at the end rather than the pygostyle tail of modern birds. Omnivorous,
eating insects and seeds of various plants, Jeholornis was about the size of a turkey,
almost 3 feet long from beak to tail. While the type species is J. prima, found in
2002, other species with varying dentition, such as J. palmapenis, have been found
since. Hebei province, China; Early Cretaceous, about 120 Ma.
Jeholosaurus: a monospecific genus of ornithischian dinosaurs in the family
Thescelosauridae, J. shanyuangensis is known from two partial juvenile specimens
from the Yixian Formation, dated. about 125 Ma; it was bipedal and probably her-
bivorous; China, Early Cretaceous.
Jianfengia multisegmentalis: a species of early arthropods known only from the
Chengjiang biota; about half an inch long with many short legs and at least two
anterior appendages, it is morphologically similar to Opabinia; China, late Cambrian,
530–515 Ma.
Jinzhousaurus: a genus of ornithischian dinosaurs in the Jehol biota; about 25 feet
long, it is known from a complete skeleton from the Yixian Formation and is tenta-
tively considered a hadrosaurid on the basis of a spike-like thumb and a small crest
at the top of the head; China, Early Cretaceous, 122 Ma.
Jiufotang Formation: a Lower Cretaceous (Aptian) Lagerstätte in Liaoning
Province; important for many feathered dinosaur specimens as well as pterosaurs,
birds, insects, and arachnids; preservation of fine detail was possible especially
because of fine volcanic ash deposits; taxa from the formation are an important part
of the Jehol biota; China, Early Cretaceous.
Joggins Formation: a Lower to Middle Pennsylvanian site in Nova Scotia whose
strata record an area of swamps and periodically flooded forests; important espe-
cially for an early terrestrial vertebrate fauna, including the temnospondyl
Dendrerpeton; Canada, Late Carboniferous, 315 Ma.
John Day Fossil Beds: a series of Eocene, Oligocene, and Miocene Konzentrat-
Lagerstätten in Central Oregon near the towns of Fossil, Mitchell, and Dayville.
Several formations represent one of the most complete sequences of Cenozoic strata
in the world, covering a period of more than 40 million years and including a great
variety of animal and plant specimens. Some of the notable mammals from the three
sites are the predatory feliforms Nimravus, Dinaelurus, Patriofelis, and Pogonodon;
bison-sized entelodonts, such as Archaeotherium and Daeodon; the rhinoceros
Diceratherium; the chalicothere Moropus oregonensis; the dwarf brontothere
Xylotitan; the mouse-deer Hypertragulus; and amynodonts. See also Clarno Nut
Beds and Painted Hills. North America, middle Eocene to late Miocene.
Josephoartigasia: a genus of the largest known rodents, in the family Dinomyidae;
related to the modern pacarana, it is estimated at about 1 ton and 10 feet long;
known only from a very large and almost complete skull; South America, Pliocene
to early Pleistocene.
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Jugulator: a monospecific genus of North American eutriconodont mammals in

the proposed family Volaticotheria; slightly larger than Volaticotherium, J. amplis-
simus was probably also a glider; it is known only from teeth and jaw fragments
from the boundary between Early and Late Cretaceous.
Juramaia: a genus of basal therian mammals known from an almost entire skele-
ton found in 2011; the type species, J. sinensis, is dated at 160 Ma and is important
in determining the date when marsupials diverged from placental mammals; China,
Late Jurassic.
Jurassic: the second period of the Mesozoic Era, 201 to 146 Ma, divided into
Early, Middle, and Late Epochs. As the supercontinent Pangaea continued to sepa-
rate into northern and southern halves, more coastline appeared globally. The arid
climate of the Triassic gave way to an almost “greenhouse” climate worldwide, with
sea level rising and rain forests replacing deserts in many places. Gymnosperms
diversified widely, including conifers and many tropical genera. The separating land
masses were dominated by dinosaurs and pterosaurs, which had appeared in the
Late Triassic. The stem-mammaliaforms of the Late Triassic developed slowly at
first but then diversified rapidly in the Middle Jurassic. Almost unnoticed, the first
birds appeared near the end of the period. In the seas, plesiosaurs, ichthyosaurs, and
sharks were dominant large predators, and the largest fish known (such as
Leedsichthys) were among their prey. Marine invertebrates, especially ammonoids,
also proliferated during the period.
Juravenator: a monospecific genus of small, feathered theropod dinosaurs; similar
to Compsognathus but only 1–2 feet long, J. starki bore some feathers but had scaly
dinosaur skin on its tail; Germany, Late Jurassic.
K
Kaiparowits Formation: a sandstone and mudstone formation in Southern Utah,

comprising layers dating between 76.6 and 74.5 Ma. One of the most productive
sites for plants and animals of the Campanian age, it has yielded crocodiles, turtles,
and especially herbivorous horned dinosaurs such as Kosmoceratops richardsoni
(found only here) and Nasutoceratops titusi. Hadrosaurs, also herbivores, are com-
mon, as well as several other very large species. The tyrannosaurids Teratophoneus
curriei and Lythronax argestes are among the predators in the formation. Many of
the specimens are complete, such as several whole dinosaur skeletons and a turtle
that died with many eggs in her body.
Kangaroo Island: a South Australian site that has produced several fossils similar
to those of the Burgess Shale and the Chengjiang biota, including Anomalocaris and
Tuzoia; it is dated at 520 Ma, intermediate in time between the two more famous
localities.
Kaprosuchus saharicus: a species of crocodiles in the family Mahajangasuchidae,
characterized by three sets of large upper and lower tusks on each side of its jaws;
these boar-like tusks worked like scissors to slice the flesh of prey, possibly dino-
saurs; variously estimated between 11 and 20 feet long; Niger, Early Cretaceous.
Karanisia: a genus of lorisid primates found in 2002 in the Fayum Depression; the
genus includes at least one undisputed species, K. clarki, which seems to have a
toothcomb similar to modern lemurs; Karanisia represents the earliest evidence of
anthropoid radiation; Egypt, middle Eocene, about 40 Ma.
Karaurus: “Karatau tail,” a monospecific early genus of salamanders in the family
Karauridae; known from one complete specimen found in the Karatau Mountains of
Kazakhstan, K. sharovi was about 7 inches long, with a broad, frog-like head, a
short body and tail, and sturdy legs; Kazakhstan, Late Jurassic, 155 Ma.
Karkenia: the form genus of seeds associated with the Ginkgophyta; global distri-
bution, Jurassic to Eocene.

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Karoo Ice Age: see icehouse conditions.

Karoo Supergroup: an extensive series of formations spanning about 100 million
years, important especially for the formations from the time between the Permian
mass extinctions and the Early Triassic; South Africa, Late Carboniferous to Middle
Jurassic.
Kayentachelys: a genus of stem-turtles either in the order Testudines or in a more
primitive classification of its own; it seems to be a link between the primitive
Odontochelys and modern turtles; about a foot long, it possessed both a plastron and
a carapace; found in the Kayenta Formation of Utah and Arizona, Early Jurassic.
Kayentatherium: a genus of Early Jurassic cynodont mammaliamorphs in the fam-
ily Tritylodontidae; known from some anterior skeletal bones and several complete
skulls found in Arizona; United States, Early Jurassic.
Keichousaurus: a genus of sauropterygian marine reptiles in the suborder
Pachypleurosauria; up to about 9 feet long, with a long neck and tail; may have been
viviparous; Guizhou Province, China; Early to Late Triassic, 249–236 Ma.
Keilor skull: a skull found in 1940 in a late Pleistocene site in southern Australia,
classified as Homo sapiens; the skull, between 13,000 and 15,000 years old, led to
the discovery of human habitation in the area about 31,000 years ago; the site is
among the earliest locations of human habitation in Australia.
Kelenken: the largest of the phorusrhacids, a monospecific genus of large flightless
predatory birds; between 7 and 10 feet tall and weighing up to 500 pounds, K. guill-
ermoi had the largest head of any known bird; Argentina, middle Miocene; see also
Andalgalornis.
Kellwasser event: the extinction event that marks the boundary between the
Frasnian and Famennian Stages near the end of the Devonian.
Kem Kem beds: a site in southeastern Morocco that has produced many fossils
from the Late Cretaceous, including several crocodylomorphs, pterosaurs, and tur-
tles, and especially several large dinosaurs such as Carcharodontosaurus and
Spinosaurus.
Kennalestes: a genus of shrew-like eutherian mammals in the order Leptictida,
known from several specimens found in Mongolia; China, Late Cretaceous, 75 Ma.
Kentrosaurus: an African genus of ornithischian dinosaurs in the family
Stegosauridae; about 15 feet long but only 1 ton, the herbivorous Kentrosaurus was
armored by several pairs of plates and spikes along the spine, as well as a large spike
projecting from each shoulder; Tanzania, Late Jurassic.
Kenyanthropus platyops: the name tentatively given to poorly known fossils, a
skull and fragments of a hominin found in the late 1990s by Meave Leakey; Lake
Turkana, Kenya, 3.5 to 3.3 Ma; see also Lomekwian industry.
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Kenyapithecus: a middle Miocene hominoid primate, known from only a few East
African fossils. Although similar to Sivapithecus and other Asian pithecines,
Kenyapithecus is now thought more likely to be in the direct line of descent to the
Hominidae. Kenyapithecus fossils date to around 16 million years ago, about 12
million years earlier than any presently recognized hominins, and the fossil record
in between is incomplete.
keratin: a fibrous protein, similar to chitin, that is an important protective material
found in hooves, nails, skin, horns, feathers, beaks, claws, and whale baleen. It is
also the basic material of mammalian hair and is considered an important evolution-
ary contributor to the rise of mammals. Keratin has uncertain origins; a 2008 genetic
study claims that keratin genes originated in an unknown reptile ancestor from
about 300 Ma.
key bed: a thin but widely occurring stratum of sedimentary rock composed pri-
marily of a particular, easily identifiable material, sometimes of fossils; the prolific
nautiloid Orthoceras, for example, forms key Ordovician beds in several places; the
key bed most often referred to is the iridium layer that marks the end of the Mesozoic
Era.
Kimberella: an enigmatic genus of Ediacaran bilaterian animals; benthic and slug-
like, it is generally considered a primitive mollusk; known from hundreds of oval
specimens dated between 558 and 555 Ma, it ranged from less than an inch long to
6 inches; Australia and Eurasia, Ediacaran.
Kirkbyoidea: a small superfamily of reticulate, straight-hinged Paleozoic ostra-
cods; Devonian to Middle Permian.
Kirkidium: a genus of articulate brachiopods in the family Pentameridae; 3–4
inches long, biconvex and astrophic, with a large pedicle valve; North America and
Europe, Silurian.
Kirtland Formation: an important sedimentary formation of the Late Cretaceous,
located in the San Juan Basin of New Mexico and Colorado; it is known for many
specimens of dinosaurs of the Campanian, including Kritosaurus and Alamosaurus,
as well as many other vertebrates; United States, 75–73 Ma.
Kloedenelloidea: a large superfamily of Paleozoic ostracods in the extinct order
Palaeocopa; global distribution, Ordovician to Triassic but less abundant in the later
Paleozoic.
Knightia: a genus of clupeiform fish in the family Clupeidae; generally from 4 to
8 inches long and herring-like, Knightia occurs in great numbers in the Green River
Formation of Wyoming and is the most common fish fossil in shops around the
world; North and South America, early Eocene.
Knightoconus antarcticus: a species of monoplacophoran mollusks in the order
Archinacelloidea; it had a chambered, conical shell and is a possible ancestor of the
Cephalopoda; known only from Antarctica, middle Cambrian.
150 K
Koilops herma: a species of tetrapods described in 2016 as similar to Greererpeton;

Scotland, Early Carboniferous (Tournaisian).
Koolasuchus: a genus of temnospondyl amphibians in the suborder Stereospondyli
and family Chigutisauridae; known from several fragmentary specimens, it is esti-
mated to have been about 15 feet long, with a large, flat, triangular head; one of the
last temnospondyls; Australia, Late Cretaceous, 125–115 Ma.
Kosmoceras: a large genus of ammonitid ammonoids in the superfamily
Stephanocerataceae; most species characterized by a combination of ribs and tuber-
cles; global distribution in Callovian deposits, Middle Jurassic.
Kosmoceratops richardsoni: an ornithischian dinosaur in the family
Chasmosaurinae, notable for the 15 horns and spikes on its head and ornate neck-
frill; a herbivore dated at 75 Ma and about 15 feet long, it had a skull almost 6 feet
long; known from only two specimens found in the Kaiparowits Formation of
Southern Utah, it seems to be closely related to a Canadian dinosaur, Vagaceratops
irvinensis; North America, Late Cretaceous.
Kritosaurus: a genus of poorly known hadrosaurid ornithischian dinosaurs in the
subfamily Saurolophinae; first found in the Kirtland Formation and also known
from fragments found in Montana, Alberta, and Mexico; North America, Late
Kryptops: “hidden face,” a proposed monospecific genus of carnivorous sauris-
chian dinosaurs in the clade Ceratosauria, known from fragmentary remains
described in 2008; K. palaios was about 20 feet long and is possibly one of the earli-
est abelisaurids; some researchers believe, however, that it may be a chimera since
some fragments were found 50 feet from others and may belong to a different dino-
saur; Ténéré Desert, Niger, Early Cretaceous, 110 Ma.
Kuehneosauridae: a family of gliding, lizard-like lepidosauromorph diapsids in
the order Eolacertilia; lateral sails on either side of the body enabled them to glide
in the air, but they could not fly; North America and Europe, Early to Late Triassic.
Kuehneotherium: a genus of primitive shrew-sized mammals in the family
Kuehniotheriidae, known from abundant tooth, dental, and mandible specimens but
no complete skeletons; weaker jaws than the morganucodonts, probably subsisting
on small insects and worms; Europe and Greenland, Late Triassic to Early Jurassic.
Kulindroplax perissokomos: a species of mollusks in the class Aplacophora,
known only from a specimen found in England in 2012; the first unambiguous apla-
cophoran with chiton-like armor, for the first time connecting the generally shell-
less Aplacophora with the Polyplacophora (chitons); its underside has a fine
covering of small spicules; late Silurian.
Kunmingella: a genus of Cambrian crustaceans in the order Bradoriida; one of
several crustaceans in the Chengjiang biota with a bivalved carapace over head and
abdomen; known only from China, early Cambrian.
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Kutchicetus: a monospecific genus of slender ancestral whales in the family

Remingtonocetidae; K. minimus was about 8 feet long and weighed between 200
and 400 pounds; its four short, gracile limbs would give it only slight mobility on
land, while its elongated and robust tail vertebrae indicate that it used caudal undu-
lation for swimming, a transitional feature in the evolution of whales; however, it
did not have tail flukes; Pakistan and India, early Eocene.
Kutorginata: a small class of primitive brachiopods, one of the three extinct
classes in the subphylum Rhynchonelliformea; global distribution, early to middle
Cambrian.
Kutorginida: an order of brachiopods in the class Kutorginata; smooth, biconvex
calcareous shells and primitive articulation are similarities to early Cambrian mol-
lusks and hyoliths; early Cambrian.
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La Brea Tar Pits: a Konzentrat-Lagerstätte in Los Angeles that has produced

many complete specimens of mammals and birds of the late Pleistocene. The bones
are impregnated by asphalt, with little degradation of bone structure or proteins;
very little soft tissue is preserved, however. More fossil birds have been produced
here than anywhere else. United States, dated between 38,000 and 11,000 years ago.
labyrinthodonts: a paraphyletic group of early amphibian tetrapods; they
descended from the rhipidistian fish and were probably ancestral to all later amphib-
ians; characterized by a labyrinthine infolding of the dentine with conspicuous
fangs; global distribution, Early Devonian to Early Cretaceous.
Laccognathus: “pitted jaw,” a genus of lobe-finned sarcopterygian fish in the
extinct order Porolepiformes and family Holoptychiidae; characterized by three
large fossae on the jaw, of unknown function; probably amphibian; North America
and Europe, Middle to Late Devonian.
Laganosuchus thaumastos: a 20-foot-long crocodile notable for its 3-foot-long
flat head and snout; like modern crocodiles, its short legs extending to the side kept
it from moving well on land; Niger, Cretaceous, 110 Ma.
Lagenida: an order of benthic foraminiferans characterized by finely perforate
monolamellar walls; global distribution, late Silurian to Recent.
Lagerpeton: a monospecific genus of dinosauromorph archosaurs in the family
Lagerpetidae; known from fragmentary specimens, L. chanarensis is estimated to
have weighed only 10 pounds; a possibly lagerpetid ichnogenus from a trackway in
Poland indicates that the genus may have originated in the Early Triassic; see also
Dromomeron; Argentina, Middle Triassic.
Lagerstätte: a German mining term commonly used for a fossil location that con-
tains an exceptionally large number of specimens. Konzentrat-Lagerstätten contain
many specimens which are not notably well-preserved but are important for their
sheer numbers (such as nesting areas, bone beds, cave deposits, and coquinas).

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Konservat-Lagerstätten, on the other hand, are noted for quality rather than quantity,
containing significant numbers of fossils preserved in exceptional detail, sometimes
in three dimensions and sometimes showing soft tissue and fragile appendages.
Lagerstätten are especially useful in understanding the development and evolution
of fossil ecosystems.
“Laggania”: see Anomalocarididae.
Lagosuchus: a genus of small archosaurian reptiles of the Triassic. Although it is
incompletely known from only one partial specimen, it may be significant in having
an unreptilian hinged ankle joint that allowed it to walk erect with its legs under its
body, a characteristic of dinosaurs. Therefore, Lagosuchus may fit somewhere in the
ancestral line leading to dinosaurs. Argentina, Middle Triassic, about 235 Ma.
Lambdotherium: a contentious genus of perissodactyl mammals formerly placed
in the Brontotheriidae but now considered only a close early relative; although
poorly known, it was smaller than the earliest accepted brontothere, Eotitanops;
North America, early Eocene.
Lambeosaurinae: a sub-family of later duck-billed ornithopod dinosaurs in the
family Hadrosauridae. Ranging in length from 25 to 35 feet, the lambeosaurs are
characterized especially by a large, hollow cranial crest; Lambeosaurus lambei also
had a backward-pointing spike behind the hollow crest. They were herbivorous and
could move both bipedally and quadrupedally. Other genera include Corythosaurus,
Hypacrosaurus, and Olorotitan; North America and Asia, Late Cretaceous,
86–66 Ma.
Lamnidae: a still-extant family of sharks, including the great white shark,
Carcharodon carcharias, and its extinct larger ancestor C. megalodon (Miocene to
Pleistocene); global distribution, Cretaceous to Recent.
Lance Formation: a stratigraphic unit of the Maastrichtian Stage in Western North
America, coextensive in places with the Hell Creek Formation; it contains an impor-
tant group of dinosaurs and smaller vertebrates from between 69 and 66 Ma, just
before the end-Cretaceous extinction.
lancelets: see Cephalochordata.
land plants: evidence of the earliest true land plants is found in fossilized spores
from the Ordovician, and in possible spores from the Cambrian; the earliest fossil
plant specimens are known from the Silurian (see Cooksonia).
lantern: see Echinoidea.
Laramidia: the name given to the western part of North America which existed as
a narrow island continent in the Late Cretaceous, when North America was split by
tectonic activity and the rising water level of the oceans; the strata laid down in the
resulting Western Interior Seaway are rich with Cretaceous fossils.
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Las Hoyas Lagerstätte: a site in Eastern Spain dated at 126–125 Ma in the late
Barremian Stage of the Early Cretaceous; Las Hoyas is well-known for the only
known specimens of the enantiornithine birds Eoalulavis and Iberomesornis, impor-
tant in establishing the development of flight in birds; the thin-bedded limestone
strata, probably laid down in a freshwater environment, have also produced a great
variety of fish, spiders, insects, and plants.
Latimeria: an extant genus of lobe-finned fish in the order Coelacanthiformes and
family Latimeriidae. The coelacanths were long considered extinct since the
Cretaceous until the discovery of modern latimerians in the twentieth century. A
specimen of L. chalumnae was discovered in 1938 near the Comoros Islands, and
several others have been found along the East African coast since then; a second
modern species of coelacanth, L. menadoensis, was discovered in Indonesia in
1997. The latimerians have fleshy lobe fins and are up to 6 feet long. Their fossil
record is unclear, but some researchers believe they originated in the Triassic. See
Coelacanthidae and Tetrapoda. Triassic? to Recent.
Laugiidae: a family of sarcopterygian fish in the subclass Actinistia, related to the
modern coelacanths of Latimeria; Europe, Greenland, and China; Early Triassic to
Late Jurassic.
Laurasia: A term referring to the northern land masses that are now North
America, Europe and Northern Asia; see paleogeography for dates.
Laurasiatheria: a clade of placental mammals that appeared in the northern
supercontinent Laurasia in the Cenomanian, at the beginning of the Late Cretaceous;
it includes the carnivorans, ungulates, shrews, bats, whales, and pangolins; see also
Eutheria; Cretaceous to Recent.
Laurentia: the name given to the early North American land mass when it was a
separate continent from the Cambrian to the Silurian.
La Voulte-sur-Rhône: a marine Konservat-Lagerstätte of the Callovian that is
notable especially for the number of soft-bodied cephalopods preserved by pyritiza-
tion, such as Doliocaris ingens; see also Ophiopinna; France, upper Middle
Jurassic.
Law of the Unspecialized: see Unspecialized Descent.
Lazarus taxon: a taxon whose fossil record shows one or more significant gaps,
with no morphological change between the first and last specimens.
Leaellynasaura: “Leaellyn’s lizard,” a monospecific genus of small herbivorous
ornithischian dinosaurs in the order Ornithopoda. It was 7 to 10 feet long with a very
long tail, but only 2 feet tall. Dated between 118 and 110 Ma, L. amicagraphica is
known from two almost complete specimens found at Dinosaur Cove in Australia,
which was close to the South Pole in the middle of the Cretaceous. It is notable for
large optic lobes in its brain which indicate good night vision. Australia, Early
Cretaceous.
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Lebachia: a genus of tall cone-bearing trees in the order Coniferales, of the late
Paleozoic and possibly Mesozoic; similar in appearance to the modern Araucaria,
it is related to the earlier Cordaitaceae and may be ancestral to modern conifers;
global distribution, Late Pennsylvanian to Permian or Triassic.
lectotype: when the type of a taxon is identified by a set of syntypes, a single
name-bearing specimen may at some point be designated as the lectotype, a later-
selected holotype.
Leedsichthys problematicus: a species of actinopterygian fish in the extinct family
Pachycormidae; possibly the largest fish known, this filter-feeding giant is estimated
to have been more than 70 feet long; the pachycormids have only partly calcified
skeletons, making their fossils sometimes difficult to interpret; Europe and Chile,
Middle Jurassic.
Leehermania: see Staphylinidae.
Leioceras: a genus of ammonoids including L. opalinum and L. lineatum; the first
appearance of the assemblage marks the beginning of the Middle Jurassic at
174.1 Ma; Spain.
Lemuroidea: a still-extant superfamily of strepsirrhine primates that includes at
least the lemurs, and in some classification schemes the lorises and galagos also;
their origins are still disputed, but they probably arose from adapiforms or lorisoids
in the Eocene; Africa, Eocene to Recent.
Lenticulina: a large genus of foraminiferans in the family Nodosariidae; global
distribution, Permian to Miocene.
Leperditicopida an order of paleocopan ostracods; from 1 to 2 inches long, thick-
shelled and straight-backed; global distribution, Cambrian to Devonian.
Lepidocaris rhyniensis: the only species in the branchiopod order Lipostraca; pos-
sibly related to the similar modern fairy shrimp in the order Anostraca, this small
(0.12 inches) crustacean is the most common arthropod in the Rhynie Chert but is
known only there, where it lived in ephemeral pools created by hot springs; Scotland,
Middle Devonian.
Lepidodendron: “scale branch,” a genus of tree-like lycopodiophytes related to the
club mosses, in the order Lepidodendrales. One of the most common fossils in the
coal beds of the Pennsylvanian, they probably reached heights of 100 feet and diam-
eters of about 4 feet, with narrow, grass-like leaves as long as 3 feet. The surface of
the branches was a mat of closely spaced squares, and the branches ended in spore-
producing cone structures that grew 1–3 inches in diameter and up to a foot in
length. Global distribution, Early to Late Carboniferous.
Lepidophylloides: a form genus of the leaves of Lepidodendron; Carboniferous.
Lepidosauria: a large subclass of reptiles that includes the orders Rhynchocephalia
and Squamata, q.v.
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Lepidostrobus: a form genus of the large cones (strobili) of Lepidodendron, q.v.

Lepidotes: a genus of semionotid actinopterygian fish; generally about a foot long,
some were as long as 3 feet; they were covered with diamond-shaped ganoid scales
and preyed on mollusks; Europe and South America, Early Jurassic to Early
Cretaceous.
Lepospondyli: a diverse subclass of small tetrapods, generally considered amphib-
ians; the lepospondyls include five orders: Adelospondyli, Aistopoda, Lysorophia,
Microsauria, and Nectridea; distinguished from the labyrinthodonts by spool-
shaped centra in the vertebrae, the lack of fangs, and the lack of infolded dentine in
the teeth; the group almost disappeared early in the Permian, with only one species
surviving to the late Permian; Pennsylvanian to late Permian.
Leptaena: a genus of articulate brachiopods in the extinct order Strophomenida
and family Raphinesquinidae; its shells are notable for a high degree of wrinkling
and ornamentation; Ordovician to Early Carboniferous.
Leptictida: a possibly paraphyletic order of small eutherian mammals, presenting
an unusual combination of primitive and specialized characters; one of the earliest
branches from basal eutherians, they may be distantly related to the Euarchontoglires
(rodents and related groups); North America, Europe, and Asia; Late Cretaceous to
Oligocene.
Leptictidium: a genus of small mammals in the order Leptictida. It was 2–3 feet
nose to tail but weighed only about 5 pounds. Leptictidium had a tail longer than the
body; short, weak forelimbs; and strong, elongated hindlimbs. Its cranium and den-
tition are archaic, and it was omnivorous. Among the few completely bipedal mam-
mals besides humans, it used its tail for balance. Best-known from the Messel Shale
Pit; Europe, early to late Eocene.
Leptochoeridae: a family of early artiodactyl mammals in the superfamily
Dichobunoidea, related to and possibly ancestral to the Entelodontidae; North
America, middle Eocene to middle Oligocene.
Leptocleididae: “slender clavicle,” a family of small plesiosaurs in the clade
Leptocleidia; generally 8–10 feet long with a round body and short tail, they swam
in shallow seas worldwide; Early Cretaceous, 140–115 Ma.
Leptocyon: “slender dog,” a genus of fox-like mammals in the family Canidae and
subfamily Caninae, weighing about 7 pounds; see also Mesocyon; North America,
Oligocene through Miocene.
Leptodus: a genus of productid articulate brachiopods in the family Lyttoniidae;
characterized by an unusually thin and flattened brachial valve and an oyster-like
pedicle valve that was cemented to the substrate or other shells; global distribution,
Permian.
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leptograptid fauna: an important fossil assemblage of the Ordovician that

includes the slender planktic graptoloid Leptograptus, extinct at the end of the
Ordovician; see Graptolithina.
Leptolepis: “thin scale,” a long-lived genus of actinopterygian fish in the family
Leptolepidae. It was one of the earliest teleosts, with a skeleton made up completely
of bone, as opposed to the combination of bone and cartilage in Pholidophorus and
some other early teleosts. Leptolepis is named for its thin scales, which lacked the
dentine or cosmine covering of most earlier fish. The combination of bony skeleton
and thinner scales improved swimming ability. Global distribution, Middle Triassic
Leptomeryx: a genus of ungulate mammals in the family Leptomerycidae; it was a
small deerlike ruminant, weighing between 20 and 30 pounds, and is thought to be
one of the earliest cervids; North America, late Eocene to early Miocene.
Lesothosaurus: a monospecific genus of small, herbivorous ornithischian dino-
saurs; about 3 feet long, with small forelimbs and manus, L. diagnosticus was evi-
dently a fast runner; Lesotho, South Africa, Early Jurassic.
Lewisuchus: a genus of early archosaurian dinosauromorphs in the clade
Dinosauriformes and family Silesauridae; related to Pseudolagosuchus; about
3 feet long, it is known from several partial specimens dated at about 247 Ma;
Argentina, Middle Triassic.
Liaoceratops: a monospecific genus of early ceratopsian dinosaurs; L. yanzigouen-
sis is known only from two possibly juvenile horned skulls found in the Yixian
Formation; Liaoceratops, estimated to have weighed only 7 pounds, is dated at
130 Ma and thought to be a basal neoceratopsian, near the time when the neocera-
topsian branch of Ceratopsia diverged from the parrot-beaked Psittacosauridae;
China, Early Cretaceous.
Liaoning: see Jiufotang Formation.
Lias: a term long used for a 20-million-year series of strata in Northwest Europe
dated between the Rhaetian Stage of the Triassic and the Toarcian of the Jurassic;
the term also came to be used for that period of time, roughly the Early Jurassic;
modern dating revealed the inaccuracy of the references, and the term is now sel-
dom used.
Libellulium: a well-known genus of flying insects in the family Petaluridae; dis-
tantly related to the modern dragonfly and sometimes classified in its order, the
Odonata; wingspan 5–7 inches; North America and Europe, Jurassic.
Lichida: an order of spiny trilobites of the middle Paleozoic; the surface of the
exoskeleton was grainy, even tubercular; some spines were long, but most were
short and thick; Early Ordovician to Late Devonian.
Liliensternus: a genus of Late Triassic theropod dinosaurs; synonym Halticosaurus;
known from several fragmentary specimens found in Germany and Switzerland,
Liliensternus was a bipedal predator, about 17 feet long with strong hindlimbs and
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shorter forelimbs; in appearance it resembles an intermediate between the smaller

Coelophysis and the larger Dilophosaurus; Europe, Late Triassic, 228–201 Ma.
Limulus: “looking sideways,” one of two surviving genera of horseshoe crabs,
chelicerates in the order Xiphosurida and family Limulidae. Horseshoe crabs are
notable as an example of evolutionary stasis, the still-extant species being almost
identical to their earliest known ancestors. The genus Tachypleus is found on the
coast of Asia and Limulus polyphemus lives on the Atlantic Coast of North America.
More closely related to spiders than to crabs, the Limulidae have survived since the
Early Ordovician.
Limusaurus: a monospecific genus of slender theropod dinosaurs in the family
Noasauridae; bipedal with short arms and about 6 feet long, L. inextricabilis is
notable especially because hatchlings were born with several dozen teeth but lost
them and did not replace them; in the nineteen specimens known, adults were com-
pletely toothless, showing only traces of the tooth sockets; no other dinosaurs or
reptiles are known to follow this pattern, although many replace and increase the
number of their teeth as they grow older; China, Late Jurassic.
Linguatula serrata: an extremely simplified pentastome, among the important dis-
coveries in the 1990s of early Paleozoic specimens of this tongue worm, which is
common today as a dog parasite; phosphatized specimens were found in the upper
Cambrian strata of the Orsten Lagerstätten; it is roughly cone-shaped, with the
mouth and attachment hooks at the large end; Sweden, late Cambrian.
Lingula: a well-known genus of marine intertidal brachiopods in the order
Lingulida; about 2 inches long, with a 6-inch pedicle, Lingula has existed world-
wide and essentially unchanged in shape since the middle Cambrian.
Lingulella: an extinct genus of inarticulate brachiopods in the order Lingulida and
family Obolidae; L. waptaensis appears in the Burgess Shale; North America, mid-
dle Cambrian to Late Ordovician.
Lingulida: a still-extant order of inarticulate brachiopods in the class Lingulata;
the most conservative of brachiopod orders, lingulids burrow in sediment, attached
by a long pedicle; characterized by an oval, biconvex shape and a shell of calcium
phosphate; middle Cambrian to Recent.
Lingulina: a genus of generally tongue-shaped foraminiferans in the family
Nodosariidae; Permian to Miocene.
Liopleurodon: a genus of large but short-necked marine plesiosaurs in the family
Pliosauridae; L. ferox, the largest species, probably weighed between one and two
tons and grew to about 20 feet long, although some estimates are much higher; the
predatory Liopleurodon was a strong swimmer; Europe, Middle to Late Jurassic.
Lipostraca: a monospecific order of freshwater branchiopods in the subclass
Phyllopoda, containing the fairy shrimp Lepidocaris rhyniensis, known only from
the Rhynie Chert of the Early Devonian, one of the earliest preserved freshwater
crustaceans.
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Liriodendron: a still-extant genus of trees in the family Magnoliaceae, known

globally from the Cretaceous through the Eocene; its range shrank during periods of
late Cenozoic glaciation; also known in the form genus of the leaf, Liriophyllum;
Middle Cretaceous to Recent.
Lissamphibia: a subclass of amphibians that includes all living members of the
class: frogs, toads, salamanders, and caecilians; Early Triassic to Recent.
Lithistida: a still-extant polyphyletic group of demosponges whose massive skel-
eton of fused siliceous spicules is often preserved; global distribution, Cambrian to
Recent.
Lithostrotion: a genus of colonial rugose corals in the suborder Stauriida and fam-
ily Lithostrotionidae; especially important in Early Carboniferous North American
biostratigraphy; global distribution, Early Carboniferous to Permian.
Litopterna: an order of superficially horselike South American ungulate mam-
mals in the superorder Meridiungulata, possibly descended from condylarths; wide-
spread from the Paleocene to the Pliocene and present in South America in the late
Pliocene when true horses first appeared there; one family of litopterns, the
Macraucheniidae, survived into the Pleistocene.
Littorina littorea: the common periwinkle, a still-extant species of gastropods in
the superfamily Littorinoidea and family Littorinidae; global distribution, Eocene to
Recent.
Littorinoidea: a superfamily of gastropods previously placed in a supposed order
Mesogastropoda but now simply considered part of the very large clade
Caenogastropoda; Eocene to Recent.
Livyatan: a genus of odontocete whales in the superfamily Physeteroidea; L. mel-
villei is known from a middle-Miocene skull and mandible discovered in the Pisco
Formation of Peru in 2010; a powerful predator, it had some teeth more than a foot
long, longer than any other animal known; it was about the size of a sperm whale,
and its skull may have in fact contained a reservoir of spermaceti; a tooth found in
Australia in 2016 is thought to belong to the genus and was dated at 5–6 Ma;
Miocene to Pliocene.
lizard-hipped dinosaur: a general term referring to the saurischian dinosaurs.
lobe-fins: see Sarcopterygii.
Lomekwian industry: the name given to primitive tools found in Kenya near Lake
Turkana, dated about 3.3 Ma and evidently made by ancestors of the species Homo,
such as Australopithecines or the uncertainly identified Kenyanthropus platyops.
London Clay: a lower Eocene formation in Southern England that has produced a
large and diverse fauna and flora which indicate a tropical to subtropical climate in
the early Eocene; younger strata in the area, through the Eocene into the Oligocene,
clearly show a cooling of the climate.
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Longisquama: “long scales,” a monospecific genus of small four-legged reptiles

that bore long shafts, possibly feathered, on both sides of its body. Since it lived
around 225 Ma, shortly after the appearance of the earliest dinosaurs, it may be
evidence that birds did not descend from dinosaurs. The shafts of L. insignis (tenta-
tively called “non-avian feathers”) may not actually be feathers, and whether
Longisquama is related to birds is unknown. It is certainly not a dinosaur, but it does
have a wishbone identical to Archaeopteryx (150 Ma) and similar to that of modern
birds. Most authorities believe it is probably an ambiguous diapsid with no bearing
on the origin of birds. Madygen Formation, Kyrgyzstan, Middle to Late Triassic.
Lonsdaleia: a genus of colonial rugose corals in the order Stauriida, characterized
by their cerioid structure and the star-burst shape of the individual polyps; Asia,
Europe, and North America; Late Devonian to Permian.
Lophialetidae: an Asian family of ceratomorph perissodactyl mammals in the
superfamily Tapiroidea; the manus is tridactyl, as in the other Asian family, the
Deperetellidae; middle to late Eocene.
Lophiodontidae: a European family of perissodactyl ceratomorph mammals in
the superfamily Tapiroidea; early to late Eocene.
lophodont: “crested tooth,” having molar teeth with a hard chewing surface formed
by transverse, triangular, or ring-shaped ridges, common in herbivorous mammals
such as elephants, manatees, and tapirs.
Lophogastrida: a still-extant and widespread order of nektic malacostracan crus-
taceans; 2016 research identified the earliest-known species, Yunnanocopia grandis
and Y. longicauda, as part of the marine faunal radiation that followed the end-
Permian extinction; China, middle Triassic.
lophophorates: a large group of aquatic animals that feed by means of a lopho-
phore, including brachiopods, bryozoans, phoronids, and hyoliths; each is consid-
ered a phylum or a class, but their exact relationships are debatable.
lophophore: a coil-shaped, feathery feeding structure in lophophorates, varying
greatly in shape and length; cilia on the lophophore sweep and pump food particles
into the mouth of the animal.
Loxonema: a genus of gastropods in the order Murchisoniina; characterized by
slender, high-spired shells with fine axial ornamentation lines; global distribution,
Ordovician to Middle Jurassic.
LUCA: acronym for “last universal common ancestor,” an unidentified type of
prokaryotic organism thought to have been living almost 4 billion years ago, only
half a billion years after the formation of Earth; it was presumably simpler than its
offspring, probably single-celled with no distinct nucleus; the earliest definitive
organic fossil is dated at 3.7 billion years ago; see also Archaea.
Lufengosaurus: a genus of sauropodomorph dinosaurs in the family
Massospondylidae. Bipedal and about 30 feet long and weighing almost 2 tons,
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Lufengosaurus is notable for the preservation of collagen in some embryonic bones,

by far the earliest-known example of soft-tissue preservation in a fossil. Since the
proteins in collagen vary from species to species, “collagen fingerprinting” may be
important in phylogenetic research. The bones from at least twenty individual
embryos show that the embryos were growing faster than in other dinosaurs. China,
Early Jurassic, 197–190 Ma.
Lunataspis aurora: a species of chelicerates discovered in Manitoba in 2008, the
earliest-known xiphosuran; dated at 445 Ma and very similar to modern horseshoe
crabs; Canada, Late Ordovician.
Lycopodiophyta: the lycophytes, the oldest division of vascular plants, known
mainly for the large trees of the Devonian and Carboniferous but represented today
by numerous families of smaller plants such as club mosses; characterized by primi-
tive, narrow leaves known as microphylls, they reproduce by shedding spores;
global distribution, late Silurian or Early Devonian to Recent.
Lycoptera: a genus of small freshwater teleost fish in the still-extant superorder
Osteoglossomorpha; one of the earliest-known in the group, it includes species that
are index fossils for the end of the Jurassic and the Cretaceous in China; Mongolia
and China, Late Jurassic to Early Cretaceous.
Lyginopteris: a genus of early seed ferns in the order Lyginopteridales; like many
other Paleozoic plant names, Lyginopteris was originally the name of a form genus
(in this case for the stem) but is used now for the whole plant; small club-shaped
glands occur on the surface of all parts of the plant; North America and Europe, Late
Devonian to Late Carboniferous.
Lyginopteridales: the earliest pteridosperms, an order of climbing seed ferns in
the extinct division Pteridospermatophyta; superficially similar in appearance to
true ferns; Early Devonian to Carboniferous.
Lysorophia: an order of snake-like amphibians in the extinct class Lepospondyli;
tetrapods but characterized by a slender, elongate body with very small limbs; pos-
sibly related to the Microsauria; North America and Europe, Pennsylvanian to mid-
dle Permian.
Lystrosaurus: a genus of dicynodont therapsids in the infraorder Dicynodontia.
Surviving the end-Permian extinction, it became the most common large terrestrial
animal of the Early Triassic. Mammal-like and about the size of a large pig,
Lystrosaurus was herbivorous and semi-aquatic, with only two large (canine) teeth.
(Its species ranged in size from 2 to 8 feet long.) Lystrosaurus probably used its
robust forelimbs for digging and burrowing. Since its distribution encompassed
Antarctica, Southern Africa, and India in the Early Triassic, it is one of the larger
animals that provide information about the time of separation of Gondwana into its
now widely removed parts. Late Permian to Early Triassic.
Lythronax: a monospecific genus of theropod dinosaurs, known from one fairly
complete skeleton discovered in 2009; L. argestes is the oldest known true
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tyrannosaurid and is ancestral to T. rex; it grew as large as 25 feet long and 5,500
pounds; southern Utah, Late Cretaceous, 80 Ma.
Lytoceras: a genus of ammonitid ammonoids, the type genus of the family
Lytoceratidae; it was 3–5 inches in diameter and had a smooth shell and a complex
suture; global distribution, Early Jurassic to Early Cretaceous.
Lytoceratida: a conservative order of ammonoids in the order Ammonitida; nota-
ble for having fewer but longer-lived species than other ammonoids, the early
Lytoceratida evolved from phylloceratids and diversified into 12 families, the domi-
nant one being the Lytoceratidae; global distribution, Late Triassic to Late
Cretaceous.
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Machairoceratops: “saber-horned-face,” a monospecific genus of ornithischian

dinosaurs in the subfamily Centrosaurinae; known from a partial skull described in
2016, it is estimated to be a medium-sized ceratopsian about 15 feet long; like its
cousin Nasutoceratops, M. cronusi was marked by two long and curved eye-socket
horns; Utah, United States; Late Cretaceous, 82–80 Ma.
Machairodontinae: “saber-tooth,” an extinct subfamily of cats with varying
lengths of the canine teeth; the family is named for Machairodus and includes
Megantereon and the later Homotherium and Smilodon; North America, Europe,
Asia, and Africa; early Miocene to 10,000 years ago.
Machairodus: a genus of lion-sized cats in the subfamily Machairodontinae;
global distribution, late Miocene to middle Pleistocene.
Macoma: a still-extant and very large genus of deep-burrowing clams in the order
Veneroida and family Tellinidae; Pleistocene to Recent.
Macraucheniidae: a family of perissodactyl ungulates in the order Litopterna,
found only in South America; the Macraucheniidae appeared in the Eocene and
were the last surviving litopterns, disappearing about 12,000 years ago; research in
2014 on Macrauchenia patachonica seems to show that, although it looked like a
camel, it was most closely related to perissodactyls such as the horse, rhinoceros,
and tapir; South America, Eocene to Pleistocene.
macrofossil: a fossil large enough to be studied directly without a microscope.
Macropodidae: a still-extant family of marsupials in the order Diprotodontia; the
Macropodidae include kangaroos and other extant genera as well as several extinct
genera; Meganesia, late Oligocene to Recent.
Macroscelididae: a family of eutherian mammals in the order Macroscelida;
although often called elephant shrews because of a very long snout and shrew-like
shape, they are not closely related to either elephants or shrews; they are

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i nsectivorous, ranging from mouse-size to rabbit-size, and their very long hindlimbs
are adapted for jumping; widespread across Southern Africa, early Oligocene to
Recent.
Macrotaeniopteris: a genus of low-growing plants with long, lanceolate leaves;
global distribution, Early Triassic to Late Jurassic
Madygen Formation: a Triassic Konservat-Lagerstätte in Central Asia, dated
between 230 and 225 Ma; its strata were laid down in forested floodplains with
lakes, rivers, and swamps; it is known especially for its well-preserved insects and
flora, as well as the unusual gliding reptile Sharovipteryx (see also Longisquama);
especially important because of the paucity elsewhere of Early Triassic insect fos-
sils; Kyrgyzstan, Late Triassic.
Maevarano Formation: a Maastrichtian site in Madagascar where dozens of spe-
cies of dinosaurs (including Majungasaurus, Masiakasaurus, and Rapetosaurus),
reptiles, the large frog Beelzebufo, birds, and mammals have been found. Several
bone beds in successive strata have been explored. Recent studies point to toxic
algae as a possible cause of death of at least one group of different animals nestled
together. Northwestern Madagascar, Late Cretaceous, 70–66 Ma.
Maiacetus inuus: a species of ancestral whales described in 2009; males grew to
about 9 feet long, females 8 feet; a fetal skeleton found with several other sets of
remains is the only known fetal specimen of an ancestral whale; Maiacetus evi-
dently gave birth on land but spent most of its time in the water; notable for four
flipper-like legs better for swimming than walking on land; Pakistan, middle Eocene.
Maiasaura: a genus of hadrosaurid ornithopod dinosaurs in the subfamily
Saurolophinae; up to 30 feet long, it was a medium-sized hadrosaur, but it is notable
especially for indications that hatchlings were altricial, requiring some parental
care; see also Egg Mountain; North America, Late Cretaceous, 78–75 Ma.
Maiopatagium furculiferum: a species of early gliding mammals described in
2017, in the order Euharamyida and family Eleutherodontidae; about 9 inches long,
it had skeletal adaptations for gliding as well as a skin membrane attached to fore-
limbs and hindlimbs; not related to the earlier glider Volaticotherium; China, Late
Jurassic, 160 Ma.
Majungasaurus: a genus of large theropod dinosaurs in the family Abelisauridae;
dated at 70–66 Ma and known from several skulls and considerable skeletal mate-
rial, it is estimated to have been 20–23 feet long, weighing about a ton; it seems
closely related to abelisaurids from India but not Africa or South America; first
called Majungatholus; Madagascar, Late Cretaceous,
Malacostraca: the class of crustaceans that contains three subclasses: the
Phyllocarida, Hoplocarida, and Eumalacostraca, the latter containing most of the
familiar modern crustaceans; Cambrian to Recent.
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Mamenchisaurus: a genus of large sauropod dinosaurs in the family

Mamenchisauridae, discovered in 1952 by C.C. Young in the Sichuan province of
China and known from several species found since then. The type species is M.
constructus, notable for its very long neck, with large and elongated neck vertebrae;
only Sauroposeidon is known to have a longer neck. Described in 1993, M. sinoca-
nadorum grew up to 115 feet long, weighed between 50 and 75 tons, and had a neck
up to 59 feet long. The length of the neck gave the family a long reach, but it is
unclear whether they fed off high branches like a giraffe or kept their head close to
the ground. Possible relatives are the slightly larger Apatosaurus and Argentinosaurus.
China, Late Jurassic.
Mammalodontidae: a family of late Oligocene toothed mysticete whales; the two
genera in the family are Mammalodon and Janjucetus, both about 10 feet long; pos-
sessing synapomorphies of both odontocetes and mysticetes, the Mammalodontidae
seem to represent one of the transitional steps from toothed to baleen whales; see
also Aetiocetidae; Australia and New Zealand, known only from the late Oligocene.
Mammalia: the class of mammals; the earliest stem-mammaliaforms appeared in
the Late Triassic (see Agilodocodon, Castorocauda, Hadrocodium,
Kuehniotherium, and Morganucodonta). They experienced a burst of evolution-
ary diversification early in the Middle Jurassic and then spread worldwide with rela-
tively little diversification for 100 million years until the end of the Mesozoic. Then
in the Paleocene began the rapid diversification leading to their present dominance
in the world. The principal characteristics that differentiate them from other verte-
brates are mammary glands, hair, three middle ear bones, a squamosal-dentary jaw
joint, and a section of the brain called the neocortex.
mammoth: a general term used, sometimes interchangeably with mastodon, to
refer to the extinct elephants of the Pleistocene, which are classified in the still-
extant superfamily Elephantoidea and family Elephantidae. Some mammoths had
upper and lower tusks as long as 7 feet and strikingly curved; they generally had
more pointed skulls and more hair than modern elephants, and mammoth species
ranged greatly in size. Found throughout North America, Africa, Europe, and Asia
until recent times, the latest survived until about 7000 years ago.
Mammuthus trogonotherii: the steppe mammoth, one of several Pleistocene gen-
era of mammoths in the family Elephantidae that were larger than modern elephants;
also called Mammuthus armeniacus.
Mammutoidea: a suborder of proboscideans that includes the mastodons of the
families Mammutidae and Stegodontidae but are not directly related to modern
elephants; like the Elephantidae, they had only upper tusks; widespread in North
America, Europe, Asia, and Africa in the Miocene; early Oligocene to Recent.
Maniraptora: the clade that includes the birds and the dinosaurs most closely
related to them, principally the therizinosaurs, dromaeosaurs, and oviraptors; Late
Jurassic to Recent.
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Mantophasmatidae: a still-extant family of predatory insects in the order

Notoptera, formerly considered a small order in its own right, called
“Mantophasmatodea.” The family is now known only in Southwestern Africa but
was probably once widespread. It contains only half a dozen species, one a recently
discovered living African insect, and a similar one a few inches long, encased in
amber, from the Eocene in Europe. These insects bear characteristics of the larger
extant groups, the mantids and the phasmids. Europe and Africa, Eocene to Recent.
manus: plural manus, the distal part of the forelimb of vertebrates, corresponding
to the human hand; it includes the wrist and hand or carpus and forefoot.
Maotianshan Shales: see Chengjiang.
Mapusaurus roseae: a genus of carcharodontosaurid dinosaurs discovered in
Western Patagonia in the late 1990s; perhaps the largest carnivorous dinosaur,
Mapusaurus was about 40 feet long and probably preyed on Argentinosaurus, one
of the largest of all dinosaurs, possibly hunting in packs; Argentina, early Late
Cretaceous.
Marasuchus: a genus of archosaurian dinosauriform reptiles; bipedal and less than
2 feet long, Marasuchus is one of several early genera close to the line of descent
leading to the dinosaurs; it is known from two specimens dated about 235 Ma;
Argentina, Late Triassic.
marine transgression: a long-term invasion of the sea over a large land area as a
result of a warmer global climate and rising sea levels; such transgressions were
widespread in the Early Ordovician, and even more in the Cretaceous, when they
created large epicontinental seas like the Western Interior Seaway of North America.
Marrella splendens: a species of stem-group arthropods abundant in the Burgess
Shale assemblage, the best-known of the marrellomorphs; only about an inch long,
it superficially resembles trilobites but has significant gill and appendage differ-
ences; its appearance led Walcott to call it a “lace crab” because of several long
spines around its body; Canada, middle Cambrian, 508 Ma.
Marrellomorpha: a subphylum of early arthropods considered to include three
species of marrellids and several specimens called acercostracans; poorly known
because of their lack of hard parts that could fossilize easily; North America,
Europe, Asia, Africa, and Australia; early Cambrian to Devonian.
Marsupialia: “pouch animals,” mammals that bear young in a relatively undevel-
oped state, so that they must live for some time in an external pouch on the mother’s
chest, where they survive on milk secreted through the skin. Metatherian mammals,
the ancestors of marsupials, probably diverged from the placental mammals in the
Middle Jurassic, but the oldest definitive marsupial appeared at only about 65 Ma,
at the beginning of the Paleocene (see Peradectes). Marsupials originated in North
America and migrated to South America, where they diversified greatly while it was
still part of Gondwana. They reached Australia via Antarctica shortly before those
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two continents separated in the early or middle Eocene, probably in a single

dispersion event. Marsupials are extant in North and South America, Australia, and
a few islands north of Australia.
Marsupites: a genus of large, purse-shaped articulate crinoids in the small order
Uintacrinida; stemless, bowl-shaped, and with ten arms, M. testudinarius is the
boundary marker for the beginning of the Campanian Stage of the Cretaceous;
Europe, Late Cretaceous.
Masiakasaurus: a genus of small carnivorous dinosaurs; notable for being the only
known theropod with long, conical, hooked front teeth that protrude from the front
of its mouth; the teeth may have been used to stab small prey; coyote-sized, with a
long tail; Madagascar, Late Cretaceous, 70–66 Ma.
mass extinction: see extinction.
Massospondylus: a genus of prosauropod dinosaurs of the Early Jurassic. Bipedal,
13 feet long, and weighing more than a ton, it had a sharp thumb claw on its small
forelimbs. Researchers reported in 2012 that they had found a large nesting site of
this dinosaur that may be 100 million years older than the previous oldest dinosaur
nest site. The eggs are 2–3 inches long and were found in clutches of up to three
dozen eggs in ten nests. The discovery indicates earlier complex social behavior in
dinosaurs than previously thought. Southern Africa, 190 Ma.
mastodon: a term sometimes confused with mammoth, but referring more spe-
cifically to the slightly differentiated proboscideans of the extinct family
Mammutidae and suborder Mammutoidea; North and Central America, late
Miocene to late Pleistocene.
Mastodonsaurus: a genus of stereospondyl capitosaur temnospondyls; growing as
long as 20 feet, M. giganteus is the largest known temnospondyl, with a large head,
eyes on top and midway in the skull, and a short tail; Europe, Middle Triassic.
Mastotermes electrodominicus: a species of termite known from amber deposits
of the late Oligocene; analysis of its DNA supports the view that termites and cock-
roaches have a common ancestor from which they evolved separately; Dominican
Republic.
Materpiscis: see Gogo Formation.
Matoniaceae: a still-extant family of ferns in the order Gleicheniales; it appeared
in the Permian and spread globally in the Jurassic and Cretaceous; the family disap-
peared from the Northern Hemisphere at the end of the Cretaceous and was thought
to be a Lazarus taxon because of three surviving genera in Southeast Asia; but it is
now known from Tertiary specimens from Antarctica, Argentina, Australia, and
New Zealand; see also Weichselia; Permian to Recent.
Matthevia: an enigmatic genus of small, worm-like animals possibly related to
polyplacophorans; late Cambrian to Early Ordovician.
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Mawsonia: a genus of large coelacanth fish in the family Mawsoniidae; the largest,
M. gigas, was about 20 feet long and preyed on fish and large invertebrates; Africa
and Brazil, Cretaceous, 110–100 Ma.
Maxakalisaurus topai: a species of titanosaurian sauropod dinosaurs discovered
in Brazil and named in 2006; probably related to Saltasaurus, it is estimated to be
at least 40 feet long and weigh 9 tons; notable for its osteoderms and vertical spine
plates, it was herbivorous and had a long neck and tail, and a small head; Late
Cretaceous, about 80 Ma.
Mayomyzon pieckoensis: a species of small, lamprey-like ostracoderms in the
family Mayomyzontidae, known from the Pennsylvanian assemblage of the Mazon
Creek site; United States, Late Carboniferous.
Mazon Creek: a Konservat-Lagerstätte in Northeast Illinois where several coal-
mining sites, especially in the Francis Creek Shale, have produced very good fossils
of about 500 species from the Late Carboniferous. The Mazon Creek deposits, laid
down in a deltaic environment similar to the Grès à Voltzia, include terrestrial,
freshwater, and marginally marine organisms from about 306 Ma, generally inside
clay ironstone concretions. The most common fossils are insects, crustaceans, and
other arthropods, as well as mollusks, jellyfish, and a variety of plants and trees.
There are marine and freshwater bivalves but no corals or crinoids, and the only
brachiopod is Lingula. Mazon Creek is known especially for the famous but enig-
matic “Tully monster” (Tullimonstrum, q.v.). United States, Late Pennsylvanian.
Mecochirus: a genus of decapod crustaceans in the family Mecochiridae; M.
longimanatus, the type species, is notable for the extreme length of its first pair of
walking legs; several specimens in the Solnhofen deposits lie at the end of a short
trackway, evidently a “death trail”; cosmopolitan, Jurassic to Cretaceous.
Mecoptera: a still-extant order of neopteran insects, commonly called scorpion-
flies; they may have been pollinators of gymnosperm plants before the appearance
of the Hymenoptera in the early Mesozoic; global distribution, Early Carboniferous
to Recent.
Medullosaceae: a family of seed ferns in the extinct order Medullosales; the larg-
est known seed fern, growing more than 30 feet high, with large leaves and seeds;
North America, Europe, Asia, and North Africa; Carboniferous to Permian.
Megacerops: a genus of perissodactyl mammals in the family Brontotheriidae; one
of the larger brontotheres, the elephant-sized Megacerops stood about 8 feet high at
the shoulder; it is also notable for a pair of blunt horns extending side-by-side from
the nasal bone and the frontal bone; North America, late Eocene.
megafauna: term used for large terrestrial mammals — especially of the
Pleistocene — weighing more than 100 pounds.
Megaloceros: a genus of large deer in the subfamily Cervinae; M. giganteus, called
“Irish elk,” was almost 7 feet high at the shoulder and had a very wide (10 feet) and
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heavy set of antlers; the five species of Megaloceros inhabited Europe and Asia
from the Pliocene to the end of the Pleistocene.
Megalochelys atlas: an extinct species of cryptodiran tortoises in the still-extant
family Testudinidae; weighing more than a ton, M. atlas is the largest known land
tortoise; Asia, Miocene to Pleistocene.
Megalodon: see Carcharodon.
Megalograptus: a genus of large eurypterids in the family Megalograptidae; it was
4 feet long and had long, spiny claws and a scorpion-like tail but no stinger; North
America, Early to Late Ordovician.
Megalolamna paradoxodon: a proposed species of sharks in the family
Otodontidae; known from 2-inch teeth found in Peru, California, North Carolina,
and Japan; dated at 20 Ma, it seems to be a sister genus of the Paleocene-Eocene
Otodus; cosmopolitan, known only from the Miocene.
Megalonychidae: a family of large, bear-like ground sloths in the order Pilosa and
superorder Xenarthra; the type genus Megalonyx, including the huge M. jeffersonii,
could maneuver bipedally, standing 10 feet tall and weighing more than a ton; North
and South America, Oligocene to the end of the Pleistocene.
Megalosauridae: a family of tetanuran theropod dinosaurs; in addition to
Megalosaurus, the family includes Torvosaurus and Streptospondylus; North
America, Europe, and Asia; Triassic to Cretaceous.
Megalosaurus bucklandii: a species of large theropod dinosaurs, 23 feet long and
bipedal, with short but robust forelimbs; it was one of the first formally identified
dinosaurs, described in the 1820s and named by Georges Cuvier for William
Buckland, the early English paleontologist; England, Middle Jurassic.
Meganesia: a biogeographic name for the large land mass that comprised Australia,
Tasmania, and New Guinea during Pleistocene ice ages when the sea level was low
enough for land bridges joining them together; they have been separated for the last
10,000 years; also sometimes used for the region of the present lands.
Meganeura monyi: a species of giant predaceous griffinflies in the order
Meganisoptera; it had a wingspan of about 26 inches; Western Europe, Late
Carboniferous to early Permian, 305–299 Ma.
Meganeuropsis permiana: a species of giant griffinflies in the order Meganisoptera,
with a wingspan of 28 inches and body length of about 17 inches; North America,
early Permian.
Meganisoptera: an order of large-winged, predaceous insects, a Carboniferous
offshoot of the Odonata, the class that includes the modern dragonflies; formerly
called Protodonata; most were similar in size to modern dragonflies, but the order
includes the largest known flying insects, such as Meganeura and Meganeuropsis,
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which had wingspans up to 28 inches; global distribution, Early Carboniferous to

late Permian.
Megantereon: a genus of early sabertooth cats in the subfamily Machairodontinae,
probably ancestral to the later Smilodon; up to 330 pounds and bearing 3-inch
canine teeth; North America, Africa, Europe, and Asia; early Pliocene (perhaps late
Miocene) to middle Pleistocene.
Megatheriidae: the largest ground sloths, a large family of xenarthran mammals
in the still-extant order Pilosa; members of the family ranged in size from 6 to 20
feet long and weighed up to 4.5 tons; South America, Oligocene to Pleistocene.
Megatherium: a genus of large mammals in the still-extant order Pilosa and extinct
family Megatheriidae; it was the largest ground sloth, about 20 feet long and weigh-
ing more than 4 tons; it had three hooked claws on each foot and was primarily
quadrupedal; it became extinct about 11,000 years ago, after the arrival of humans;
South America, early Pliocene to late Pleistocene.
Megazostrodon: a genus of small early cynodont mammals in the family
Megazostrodontidae; 4–5 inches long, a scansorial insectivore, it seems to represent
one of the last transitional stages between the cynodonts and true mammals; Elliot
Formation, South Africa, Late Triassic to Early Jurassic.
Megistaspis: a genus of ptychopariid trilobites in the family Asaphidae; Australia,
Europe, Asia, and North America; Early to Middle Ordovician.
Megistaspis acuticauda: a large species of shovel-nosed burrowing ptychopariid
trilobites; related to the smaller Illaenus but more than a foot long and characterized
by a distinct and pointed cephalon; cosmopolitan, Early Ordovician.
Mei long: “sleeping dragon,” a species of small troodontid dinosaurs in the Jehol
biota; a basal troodontid, it is almost 3 feet long, with wing-like arms and long,
clawed fingers; known from two well-preserved specimens, both found in a bird-
like sleeping posture; neither has obvious feathers but may have had a feather-like
coating; China, Early Cretaceous, 125 Ma.
Melanopsis: a genus of Miocene gastropod mollusks notable for its evolutionary
stasis for 7 million years, followed by a period of rapid change for 2 million years
as the environment changed from marine to freshwater; in this period the shell size
increased and a distinctive shouldering pattern evolved; Europe and Asia, Miocene.
Melanorosaurus: a genus of early sauropodomorph dinosaurs in the family
Melanorosauridae; quadrupedal and one of the larger prosauropods, it was about 26
feet long and weighed a little more than a ton; Melanorosaurus is dated about 210
Ma, close to its relative Antetonitrus; South Africa, Late Triassic.
Melittosphex burmensis: a species of bees in the family Melittosphecidae, pre-
served in amber; the earliest-known specimen of a bee, dated at 99–100 Ma; Burma,
Early Cretaceous.
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Meniscotherium: a genus of herbivorous protoungulate mammals in the family

Phenacodontidae; ranging from 15 to 30 pounds, they had hooves and the selen-
odont teeth of ruminant herbivores; North America, Eocene.
Menoceras: a genus of perissodactyl mammals in the family Rhinocerotidae;
about 5 feet long, it was smaller than its relative Diceratherium but had the same
unusual pair of side by side nasal horns; North America as far south as Panama, late
Oligocene to early Miocene.
meraspis: the intermediate growth phase of trilobites, during which the thorax and
pygidium became distinct; see also protaspis and holaspis.
Merostomata: an obsolete term for a class of chelicerate arthropods that was con-
sidered to include the extinct order Eurypterida and the extant Xiphosura; the latter
are now sometimes regarded as a class in their own right; Middle Ordovician to
Recent.
Merychippus: a genus of three-toed Miocene horses in the family Equidae, the first
known grazing horse; at 3 feet high it was the tallest horse of the time, and it gave
rise to several other genera of grazers, such as Hipparion and Protohippus; North
America, early to late Miocene.
Merycoidodontidae: a large family of artiodactyl ruminants in the still-extant
suborder Tylopoda and extinct superfamily Merycoidodontoidea; previously called
Oreodontidae; they were browsers with a selenodont dentition and claws instead of
hooves; abundant in the Oligocene, they were pig-like in appearance but are gener-
ally considered camelids; known across North America from Oregon to Central
America, late Eocene to Miocene.
Mesacanthus: a large genus of primitive acanthodian fish in the family
Acanthodidae; ranging from a few inches to a foot in length, Mesacanthus is char-
acterized by pectoral, anal, pelvic, and dorsal fin spines, and it is named for an
unusual pair of pre-pelvic fins in the middle of the body; Scotland, Early to Middle
Devonian.
Mesocyon: a species of dog-like mammals in the family Canidae and extinct sub-
family Hesperocyoninae; see also Leptocyon; weighing about 16 pounds, it is
known only from sites in Oregon, California, Colorado, and Nebraska in Western
North America; early Oligocene to early Miocene.
Mesohippus: a genus of early Equidae in the extinct subfamily Anchitheriinae;
slightly larger than Hyracotherium but still only about 2 feet high at the shoulder; a
cousin, Anchitherium, migrated to Eurasia in the early Miocene, becoming extinct
in the Pliocene; North America, late Eocene to early Oligocene.
Mesolimulus: a genus of xiphosurans in the family Limulidae; almost identical to
the modern horseshoe crab, several specimens of Mesolimulus are known from the
Solnhofen quarries, including a famous one that died at the end of a well-preserved
30-foot trackway; Europe, Late Jurassic 151–146 Ma.
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Mesonychidae: a family of protoungulate mammals in the order Mesonychia,

related to a variety of modern mammals, including whales, hippopotamuses, and
dolphins. Although they were long thought to be direct ancestors of whales, their
relationship is now thought to be more distant. Ranging from 3 to 13 feet in length
and having four-clawed feet, the mesonychids encompassed a wide variety of herbi-
vores and carnivores. They declined in the Eocene, with only one genus, Mongolestes,
surviving into the Oligocene. Europe, Asia, and North America; early Paleocene to
early Oligocene.
Mesonyx: a genus of carnivorous mammals in the family Mesonychidae; wolf-like
in appearance and behavior although not directly related, Mesonyx grew to about 5
feet long, excluding its tail; North America and Asia, late Eocene.
Mesophyticum: term sometimes used for a paleobotanic division of time roughly
corresponding to the Mesozoic Era and characterized by the plant life of the
Mesozoic.
Mesosauria: a single-family order of small aquatic reptiles, also called
Proganosauria. Their taxonomic position is controversial, some considering them
basal “parareptiles,” others preferring to place them in the stem-reptile family
Captorhinidae, and some even claiming similarities to synapsids. They are the
earliest-known aquatic reptiles, possibly descended from terrestrial ancestors, and
may be the earliest-known viviparous reptiles. About 3 feet long, they probably
inhabited a saltwater basin where Southern Africa and South America were still
joined in the early Permian.
mesosuchians: a large and paraphyletic grouping of archaic terrestrial or semi-
aquatic reptiles, generally considered in the superorder Crocodylomorpha; charac-
terized by elongated and flattened skulls, and in some genera very long tails;
widespread in the Jurassic and Cretaceous.
Mesothelae: the primitive suborder of arachnids in the order Araneae, with only
one extant representative, the Liphistiidae of East Asia; the Mesothelae are among
the earliest spiders, known from North America and Europe from the Pennsylvanian
to the Permian; see also Palaeothele.
Mesozoic Marine Revolution: Early in the Mesozoic Era, several marine preda-
tors developed new, durophagous shell-crushing abilities: first the placodont marine
reptiles, then the ichthyosaurs, and in the Cretaceous the mosasaurs and ptychodon-
tid sharks. Because of the increase in durophagy, the populations of ammonoids and
other gastropods, crinoids, and brachiopods decreased rapidly, and the character of
the benthos was changed greatly, with many sessile and slow-moving taxa becom-
ing extinct. Some species developed new protections, such as lighter, streamlined
bodies that could escape by swimming or jet propulsion. Bivalves survived better
than brachiopods, mainly by burrowing and becoming infaunal; some, such as the
Pectinidae, developed the ability to jump away from predators.
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Messel Shale Pit: an Eocene Konservat-Lagerstätte near Frankfurt am Main.

Representing a complete lakeside ecosystem in a subtropical forest, it has yielded a
great variety of fossils, including especially mammals but also plants, insects and
other arthropods, and invertebrates. The thirty-five mammal species include
Darwinius masillae, possibly a very primitive primate, and the very unusual
Leptictidium. See also Palaeopython. Germany, early Eocene, 49 Ma.
Messinian Event: (also known as Messinian Salinity Crisis) the drying up of the
Mediterranean Sea at the end of the Messinian Age of the Miocene, from 5.96 to
5.33 Ma; the causes are unclear.
Metacopina: a suborder of ostracods in the order Podocopida; their carapaces
have features that seem to be transitional between the other two podocopid subor-
ders, the Platycopina and the Podocopina; global distribution, early Paleozoic to
Early Jurassic.
Metaplacenticeras: a genus of ammonoids in the order Ammonitida and family
Placenticeratidae; important as an index fossil for the late Campanian; Japan,
California, and Antarctica; Late Cretaceous.
Metasequoia: a genus of deciduous redwoods with one species still living in
China; originating in the Late Cretaceous, Metasequoia forests were found at high
latitudes across the Northern Hemisphere until the Miocene.
Metaspriggina: a monospecific genus of very primitive chordates known from the
Burgess Shale; it was so named because of a superficial similarity to the Ediacaran
genus Spriggina, but the two are now considered unrelated; M. walcotti may be a
craniate and possibly a primitive fish; Canada, middle Cambrian, 508 Ma.
Metatheria: an infraclass of therian mammals that includes all marsupials, extinct
and living. The earliest-known metatherian fossils, such as Sinodelphys szalayi,
have been found in Cretaceous sites in China. The first true marsupials are found in
lower Paleocene strata in North America (see Peradectes). Migrating into South
America by the end of the Cretaceous, marsupials then appeared in Europe and
North Africa in the Eocene. They reached Australia in the early or middle Eocene,
shortly before the island continent separated from Gondwana.
metazoan: an animal that has evolved beyond the one-celled stage, having a com-
plex cell structure and nervous system; although long known to be widespread and
diverse in early Cambrian deposits, only around 1950 were metazoans found in
upper Precambrian strata in the Ediacara Hills of Australia.
Metoposaurus: a genus of large temnospondyl amphibians probably close to the
ancestral stock of modern amphibians; up to 10 feet long, the salamander-like
Metoposaurus probably occupied a crocodile-like niche, feeding mainly on fish; the
genus includes M. diagnosticus algarvensis (discovered in Portugal in 2015) and
three other members; Europe, Late Triassic, 230–200 Ma.
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Metriorhynchidae: a family of marine crocodylomorphs in the clade

Thalattosuchia; the type species is Metriorhynchus (“moderate snout”), about 10
feet long with a streamlined body, paddle-like forelimbs, and a tail fin; North and
South America and Europe; Middle Jurassic to Early Cretaceous.
Miacis: a genus of arboreal carnivores in the extinct superfamily Miacoidea and
family Miacidae; close to the ancestry of modern carnivores, weasel-sized Miacis
descended from insectivores and had a larger brain than its cousins the creodonts;
North America, Europe, and Asia; late Paleocene to late Eocene.
Michelinia: a genus of colonial favositid corals in the extinct order Tabulata;
global distribution, Early Devonian to late Permian.
Micraster: an evolutionarily important genus of irregular atelostomate echinoids
in the still-extant subclass Euechinoidea and order Spatangoida. Characterized by
its heart shape and dense covering of small tubercles which supported a forest of
short spines, Micraster lived in a burrow in marine sediment. Study of the gradually
evolving changes in species of Micraster in 10 million years (almost 500 feet) of the
chalk beds of the Late Cretaceous made it well-known in the nineteenth century as
an example of evolution. North America, Europe, Africa, and Antarctica; Late
Cretaceous to Paleocene.
Microcystis: a genus of freshwater cyanobacteria that can produce algal blooms
with toxins that cause anoxia; global distribution, Permian to Recent.
Microdictyon: a genus of worm-like animals now known from several complete
fossil specimens and considered a possible basal onychophoran, but it may be unre-
lated to any later taxa; see M. sinicum; Tommotian biota, global distribution, early
Cambrian.
Microdictyon sinicum: a species of worm-like animals so far unrelated to later
taxa; its discovery in 1989 is remarkable for solving the mystery of small, round
phosphatic caps called Microdictyon that were one of the enigmatic elements of the
small, shelly fauna; these caps covered the joints where the ten pairs of leg-like
appendages joined the body of M. sinicum; China, Chengjiang biota, early Cambrian.
Microdomatidae: a family of marine gastropods in the superfamily Trochoidea;
this family of high-spired sea snails includes the subfamilies Microdomatinae and
Decorospirinae; North America, Asia, and Europe; Early Devonian to Early Triassic.
microfossil: a fossil whose typical form is microscopic in size; because so abun-
dant and rapidly evolving, many microfossils are extremely useful in biostratigra-
phy; some of the more important groups are the diatoms, Foraminifera, Radiolaria,
and other protists.
micropaleontology: the branch of paleontology dealing with fossils that require a
microscope to see their form or their important characteristics.
Microraptor: a genus of small (less than 3 feet long) carnivorous feathered dino-
saurs in the family Dromaeosauridae, known from several Aptian sites in Liaoning
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Province. It had feathers on forelimbs and hindlimbs, which may have been used for
short glides or flights. A 2012 study argues from comparisons of melanosomes with
modern birds that the feathers were probably iridescent. Synonym: Cryptovolans
pauli. China, Early Cretaceous, 125–113 Ma.
Microsauria: the most diverse and species-rich order of amphibians, in the sub-
class Lepospondyli; the microsaurs are all characterized by short tails and small legs
but were otherwise very different in form and habit, some being primarily aquatic
and others primarily terrestrial; North America and Europe; Late Carboniferous to
late Permian.
Microsyopidae: a family of early primates in the extinct suborder Plesiadapiformes;
the type genus is Microsyops; late Paleocene to middle Eocene.
Microtheriomys brevirhinus: a small species of beaver, living about 28 Ma, that
seems to be closely related to the modern beaver; discovered in the John Day Fossil
Beds in Oregon in 2014, where several other beaver species have been found; North
America, Oligocene.
Microtragulus: a genus of didelphoid marsupials in the extinct family
Argyrolagidae; the omnivorous Microtragulus resembled a small kangaroo; South
America, late Miocene to Pleistocene.
Middle Awash: an important site in Ethiopia for hominid and hominin fossils dur-
ing the period between 5 and 2.5 Ma. A series of important fossils have been found
here within a few surface miles of each other. The fossils, which have been found by
different collectors over the last few decades and may or may not be directly related
to each other, include several specimens each of the early Ardipithecus ramidus
(about 4.4 Ma) and the later Australopithecus anamensis (about 4.2 Ma) and Au.
garhi (2.5 Ma).
Milleporidae: a still-extant family of cnidarians in the class Hydrozoa and order
Capitata, commonly called fire corals for their powerful sting, but not true corals;
they form colonies of soft-bodied polyps but, like the stylasterines, differ from other
Hydrozoa in having calcareous parts that fossilize; global distribution, Cretaceous
to Recent.
mimicry: see Batesian mimicry.
Miocene: the fourth epoch of the Cenozoic, from 23 to 5.3 Ma. After the numerous
extinctions of archaic mammals at the end of the Oligocene, the Miocene saw the
almost global radiation of all modern mammals. If Ardipithecus kadabba is con-
firmed, it will mark the first appearance of hominins as coming at the end of the
Miocene, between 5.8 and 5.2 Ma.
Although grasses appeared much earlier, in the late Paleocene, they did not
spread rapidly until the end of the Oligocene. The early Miocene is characterized by
an accelerated change from lush forests toward open grassland, so that by the end of
the period open and wooded grassland communities were common worldwide.
Ungulates in the Miocene evolved in response, developing specialized teeth for
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grazing the tough grasses, as well as longer limbs to enable escape from predators
in the open country. Although the northern continents saw considerable interchange
of animals, South America and Australia continued to be isolated, developing many
eccentric groups.
Miocidaris: a genus of pencil echinoids in the still-extant order Cidaroida and the
type genus of the family Miocidaridae; one of the few cidaroids that survived the
end-Permian extinction; North America, Europe, and Asia; late Permian to Early
Jurassic.
Mioplosus: a genus of actinopterygian fish in the still-extant family Percidae; up to
20 inches long, they were predators; global distribution, early to late Eocene.
Miosireninae: one of several extinct subfamilies in the sirenian family Dugongidae;
Europe, Africa, and North America; Oligocene to Miocene.
Mississippian: see Carboniferous.
Mistaken Point: an important Konservat-Lagerstätte in exposed surfaces of argil-
lites on the Avalon Peninsula in Newfoundland, with a rich Ediacaran assemblage
of several types of rangeomorph specimens, including Fractofusus misrai; some are
similar to specimens found in Russia and Australia, while others are unknown
except at Mistaken Point; Canada, Ediacaran, 575–560 Ma.
mitosis: the process of cell division by which cells in living organisms multiply.
The process is almost identical in plants and animals, evidence that both are
descended from a common ancestor. Genetic evidence indicates that this common
ancestor may have been a one-celled organism that lived about 1.5 billion years ago.
Mixodectidae: a family of insectivorous mammals in the order Dermoptera; North
America and Europe, Cretaceous to Paleocene.
Mixosaurus: “mixed lizard,” a genus of ichthyosaurs in the family Mixosauridae;
a medium-sized ichthyosaur 3–6 feet long, the most common of the Triassic; prob-
ably transitional between earlier, eel-like forms and the porpoise-like ichthyosaurs
of the Late Triassic; global distribution, Middle Triassic.
Mixotoxodon: a genus of notoungulates in the family Toxodontidae; the size of a
rhinoceros and weighing about 4 tons, it was the largest notoungulate; South,
Central, and North America; middle to late Pleistocene, the latest specimen dated at
around 25,000 years ago.
moa: see Dinornithidae.
Mobergella: a problematic fossil in the “small, shelly fossil” group; a circular cap
or shell, known especially from Scandinavia, early Cambrian.
Modiolus: a still-extant genus of medium-sized bivalve mollusks in the family
Mytilidae and subclass Pteriomorphia; a burrowing mussel, attaching itself byssally
to debris in sediment; global distribution, Early Devonian to Recent.
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modular evolution: of organisms in which advanced anatomical components are

combined with primitive components; the wukongopterid pterosaurs are a good
example.
Moeritheriidae: a family of pig-like mammals in the order Proboscidea; although
the moeritheres resemble tapirs and pygmy hippopotamuses, they are not related;
they are probably not ancestors of elephants either, but a separate extinct branch of
proboscideans; they were trunkless but bore long canines that can be considered
small tusks; North Africa, late Eocene.
mold: an impression left by a dead organism in soft material that has solidified; an
external mold is formed by sediment or volcanic material creating an impression of
a dead organism; an internal mold is formed by material filling any void created by
decay of the soft parts of an organism.
molecular paleontology: study of fossilized remains at the molecular level, focus-
ing on genetic material such as DNA; investigators have identified possibly viable
genetic material in insects preserved in amber as old as the middle Paleocene;
molecular techniques have been important in research on such species as the
Quagga, on hominins such as Homo heidelbergensis and the Denisovans, on dino-
saurs such as Diplodocus hallorum, and even older organisms; some investigators
advocate trying to recreate extinct organisms from this genetic material.
Mollusca: a large and very diverse phylum of marine and freshwater organisms.
The most prominent extant classes are the Bivalvia, Gastropoda, Cephalopoda,
Polyplacophora, Monoplacophora, Aplacophora, and Scaphopoda; two other classes
are extinct and important in the fossil record: Rostroconchia and Helcionelloida.
The main defining characters of mollusks are a muscular mantle, a rasping radula,
and a definitive nervous system. Global distribution, Cambrian to Recent.
Mongolestes: a genus of protoungulate mammals in the family Mesonychidae; the
last surviving mesonychid, Mongolestes was about 7 feet long and had larger teeth
than other mesonychids; Mongolia, Eocene to early Oligocene.
Monobathrida: a large order of generally pelmatozoan monocyclic crinoids in the
order Camerata; global distribution, Early Ordovician to late Permian.
Monoclonius: a contentious genus of ceratopsian dinosaurs usually classified in
the subfamily Centrosaurinae; about 17 feet long and 6 feet high, it is characterized
by one long horn on its beaked snout; known only from incomplete remains, some
of which have been lost and some reclassified, Monoclonius is probably a nomen
dubium.
Monocyathida: a class of archaeocyathans with only a single porous wall, usually
solitary; Australia, Antarctica, North America, Asia, and Europe; middle Cambrian.
monograptid fauna: an important graptolite fauna, generally characterized by a
single stipe, representing the graptoloids’ tendency toward scandent growth and
fewer stipes; see Graptolithina; early Silurian to Early Devonian.
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Monograptus: a genus of graptoloids in the family Monograptidae; scandent, gen-

erally with a single straight stipe with hooked thecae; the first appearance of M.
uniformis is considered the marker of the GSSP at the base of the Lochkovian Epoch
(419.2 Ma), the beginning of the Early Devonian; global distribution, early Silurian
to Early Devonian.
Mononykus olecranus: “single claw,” a unique species of theropod dinosaurs in
the family Alvarezsauridae; 3 feet long and lightly built, with slender legs; very
short but robust forelimbs, equipped with only one claw, which was 3 inches long;
these may have been adaptations for hunting mound-living insects; known from
only one partial specimen dated about 70 Ma; Nemegt Formation, Mongolia, Late
Cretaceous.
monophyletic group: see polyphyletic group.
Monophyllites: a genus of ammonoids in the order Phylloceratida and family
Ussuritidae; thin and evolute, 3–5 inches across; Asia, Europe, North America, and
New Zealand; Middle to Late Triassic.
monophyodonty: see dentition.
Monoplacophora: a primitive but still-extant group of mollusks with single,
limpet-like shells. It is generally considered polyphyletic, but the fossil record is
sparse and leaves many questions open; in some classifications the monoplacopho-
rans are included in the Gastropoda despite considerable anatomical differences.
They also have some features similar to segmented worms. The living monopla-
cophorans, such as Neopilina, are bottom-feeders in deep-ocean water. Cosmopolitan,
early Cambrian to Recent.
Monotremata: “single-hole,” a still-extant order of egg-laying mammals; once
widespread, the monotremes are now restricted to the platypus and echidna of
Australia and New Guinea; characterized by reptilian, avian, and mammalian traits,
including a cloaca through which the urinary, intestinal, and reproductive tracts all
open to the outside, rather than with separate urogenital and intestinal openings;
Late Triassic to Recent.
Montanalestes: a poorly known genus of small mammals closely related to later
eutherians; known only from six teeth; United States, Early Cretaceous,
113–100 Ma.
Montceau-les-Mines: a Konservat-Lagerstätte in central France, noted for flat-
tened and three-dimensional fossils of a tropical biota, including pteridosperms,
insects, arachnids and other chelicerates, and tetrapods; France, Late Carboniferous.
Monticulipora: a genus of stenolaemate bryozoans in the order Trepostomata;
massive colonies, sometimes encrusting brachiopods; North and South America and
Europe, Late Ordovician.
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Montlivaltia: a genus of large, solitary scleractinian corals in the family

Montlivaltiidae; up to 4 inches across, cup-shaped to cylindrical; global distribu-
tion, Early Jurassic to Cretaceous.
Morganucodon: a genus of advanced cynodonts in the family Morganucodontidae,
one of the earliest true mammals; about 4 inches long, it was omnivorous and had a
double jaw joint; North America, Europe, and Asia; Late Triassic to Middle Jurassic.
Morganucodonta: an order of early mammaliaform cynodonts; generally consid-
ered true mammals, but their classification remains contentious; morganucodonts
were shrew-sized and slender, with a relatively large head, and they probably had a
good sense of smell; they had strong jaws that were able to crush insects with stiff
exoskeletons; but their specialized, heterodont dentiton indicates that they were also
carnivorous; global distribution, Late Triassic to Late Jurassic.
Moropus: “slow foot,” a genus of clawed perissodactyl mammals in the superfam-
ily Chalicotherioidea; found from Oregon to Florida through the Miocene, Moropus
was more similar to horses than other chalicotheres; as its name indicates, it walked
awkwardly because of its large forelimb claws; it stood about 8 feet high at the
shoulder and weighed up to 600 pounds; United States, Miocene.
Morotopithecus: a genus of Miocene hominoids described in 2004 and dated at
20.6 Ma or earlier; known from dental, hip, and shoulder fragments, Morotopithecus
may be transitional between the apes of the Miocene and more advanced homi-
noids; Uganda, early Miocene.
Morrison Formation: a large geologic formation in the Rocky Mountains of
North America, stretching west from Kansas to central Utah and from New Mexico
up to southern Alberta and Saskatchewan. The Morrison Basin fossils, most famous
for many species of dinosaurs, comprise a complete community of terrestrial life in
the Kimmeridgian and Tithonian Ages at the end of the Late Jurassic. Almost all
locations are fluviatile or lacustrine, with almost no marine strata. Important dino-
saur specimens have been found at many outcrops of the formation, but the largest
Konzentrat-Lagerstätten are Utah’s Dinosaur National Monument and Cleveland-
Lloyd Dinosaur Quarry, Wyoming’s Como Bluff, and Colorado’s Dry Mesa
Dinosaur Quarry.
Mosasauridae: a family of large marine lizards in the order Squamata and extinct
superfamily Mosasauroidea. They probably evolved from the smaller, less special-
ized aigialosaurs in the Early Cretaceous and diversified worldwide to become the
dominant marine predators, becoming extinct at the end of the period. First found in
1764 at Maastricht on the Meuse River, mosasaurs were air-breathing and ranged
from 3 to 60 feet long. Mosasaurus hoffmannii, the type species and among the last
of the mosasaurs, is dated between 70 and 66 Ma; its jaws were about 40 inches long
and its body about 33 feet. Powerful swimmers, they have the body shape of a
streamlined varanid, such as the modern monitor lizards. They preyed on fish of all
sizes, sharks, ammonoids, turtles, and sea birds. They had double-hinged jaws,
gulping down small prey whole; some species had two large teeth pointing back-
182 M
ward down the throat to prevent prey from escaping. Others, such as Globidens and
Prognathodon, had blunt, rounded teeth adapted to crushing ammonoids and
bivalves. They were covered with scales, and the presence of melanin in their skin
indicates possible countershading like many modern fish (dark above and light
below). Global distribution, Early to Late Cretaceous.
Moschops: a poorly known genus of dinocephalian therapsids in the family
Tapinocephalidae; massively built, especially the skull, with long, stout teeth
adapted for a herbivorous diet; known only from the Karoo Basin of South Africa,
middle to late Permian.
Mosineia: a genus of arthropods in the family Euthycarcinidae; three post-
abdominal exoskeletons are the only specimens of the famous M. macnaughtoni,
one of the body fossils of early arthropods found along with their trackways in
Cambrian strata in Wisconsin and Quebec; see also Blackberry Hill; North
America, middle or late Cambrian.
Moythomasia: a basal genus of small palaeoniscid actinopterygian fish; only 4
inches long, Moythomasia had a downward-pointing tail with a row of bony nodes
along its dorsal edge; like the much larger Cheirolepis, it had overlapping ganoid
scales connected in a peg and socket design; Europe and Australia, Middle to Late
Devonian.
Mucrospirifer: a well-known genus of brachiopods in the class Rhynchonellata
and order Spiriferida. Mucrospirifer was 1–2 inches long, characterized by a long
hinge line and a prominent fold and sulcus. Several species are index fossils for the
Middle and Late Devonian. Asia, North and South America, and Europe; Early
Devonian to Middle Mississippian.
Müllerian mimicry: see Batesian mimicry.
Multituberculata: a large and very successful order of protoeutherian mammals,
classified separately from the Theria and the monotremes, but closer to the Theria;
characterized by large incisors and two or three longitudinal rows of simple conical
cusps; the order comprises many families, ranging from a few inches to 4 feet long;
omnivorous but generally herbivorous, they include the earliest herbivorous mam-
mals; global distribution but principally in the northern continents, Late Jurassic to
early Oligocene.
Murchisonia: a very long-lived genus of prosobranch gastropods in the order
Archaeogastropoda and extinct superfamily Murchisonioidea; characterized by an
elongate, high-spired, and multi-whorled shell, generally with a slit band that runs
in a spiral ridge from bottom to top; global distribution, Early Ordovician to Late
Triassic.
Muridae: a family of small mammals including mice, rats, and gerbils; stem
murids appeared in the early Eocene, probably sharing the ancestry of the Dipodidae,
and true murids enter the fossil record in the early Miocene; the murids are the larg-
est mammal family, diversifying from the middle Miocene to the Pliocene, and
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since the Pleistocene thriving commensally with humans; global distribution, early
Eocene to Recent.
Mustelidae: a large and still-extant family of caniform carnivorous mammals in
the superfamily Musteloidea; the family includes the ancestors of modern weasels,
martens, otters, badgers, minks, ferrets, wolverines, fishers, and stoats; like other
Carnivora, the first mustelids probably descended from the Miacidae in the Eocene
and quickly spread worldwide.
Muttaburrasaurus: a genus of ornithischian dinosaurs in the family
Rhabdodontidae; omnivorous, up to 26 feet long and weighing 3 tons, it had signifi-
cant forelimbs but was probably mainly bipedal; see also Altirhinus; Australia,
Early Cretaceous, 112–100 Ma.
Myalinidae: a family of bivalve mollusks in the superfamily Ambonychioidea;
marine and freshwater genera are important stratigraphically, especially in the
Carboniferous; see also Naiadites; global distribution, Ordovician to Triassic.
Myanmarpithecus: a proposed genus of primates in the family Amphipithecidae,
found in 2001 in the Pondaung Hills of Burma; Eocene.
Mygalomorphae: a still-extant infraorder of spiders that includes the funnel and
trapdoor spiders; see also Rosamygale; cosmopolitan, Early Triassic to Recent.
Mylagaulidae: a family of sciuromorph rodents which includes the horned rodent
Ceratogaulus; possibly related to the Aplodontoidea, whose only extant member is
Aplodontia, the mountain beaver; North America and Asia, late Oligocene to early
Pliocene.
Myllokunmingia: a genus of basal chordates, generally considered the earliest-
known vertebrate; about an inch long, it has a notochord and a skull and skeletal
structures made of cartilage; China, early Cambrian, 530–525 Ma.
Mylodon: see Paramylodon.
Myophorella: a genus of marine bivalve mollusks in the order Trigoniida and
extinct subfamily Myophorellinae; the irregularly shaped shell is covered with rows
of tubercles; in some specimens the soft tissue has been mineralized and preserved;
global distribution, Early Jurassic to Early Cretaceous.
Myriapoda: a subphylum of arthropods that includes centipedes, millipedes,
arthropleurids, and other small groups; global distribution, Cambrian to Recent.
Mysticeti: the baleen whales, a suborder of generally toothless, filter-feeding
Cetacea that appeared at the end of the Eocene or beginning of the Oligocene. The
modern Mysticeti include fin, gray, right, and blue whales, the latter at 110 feet long
being the largest animal known to have existed. They evolved from an archaeocete
ancestor, probably separately from the odontocetes. Some early genera were par-
tially toothed, and the evolution of fully edentulous baleen whales is well docu-
mented in the fossil record. The North Pacific family Aetiocetidae of the early
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Oligocene was toothed but also had baleen structures in the jaw. The South Pacific
family Mammalodontidae of the late Oligocene, including Mammalodon and
Janjucetus, consists of early toothed mysticetes, some of them having a dentition
similar to that of their terrestrial ancestors. The size of baleen whales stayed rela-
tively constant through the Miocene but increased rapidly in the Pliocene for
unknown reasons. Early Oligocene to Recent.
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Nacholapithecus: the most complete specimen of a Miocene ape, a genus of homi-

noids dated at 15 Ma and tentatively placed in the family Proconsulidae; Kenya,
middle Miocene.
Nacimiento Formation: an early Paleocene heterogeneous non-marine formation
in Western New Mexico, composed of sandstone, shale, and siltstone; laid down
primarily in lacustrine and floodplain environments, it has produced many fossils
from the time after the end-Cretaceous extinction, including champsosaurs, taenio-
labidoids, and the early primate Torrejonia; United States, 65–61 Ma.
Naegele’s giant jaguar: see Panthera leo atrox.
Nahecaris: a genus of malacostracan crustaceans in the subclass Phyllocarida;
about 6 inches long, it has long antennae, a pointed tail, and several pairs of short
legs; Germany and Morocco, Early Devonian.
Naiadites: a genus of non-marine bivalve mollusks in the family Myalinidae; along
with Carbonicola and Anthraconaia, it is an index fossil in the British Coal
Measures of the Carboniferous; cosmopolitan, Late Mississippian to Early
Pennsylvanian.
Nama Group: rock formations in Namibia that have produced three-dimensional
casts of Ediacaran and early Cambrian organisms.
Nannippus: “dwarf horse,” a genus of perissodactyl mammals in the family
Equidae. It appeared in the late Miocene and co-existed with other North American
horses until it became extinct in the late Pliocene. It was small and slender, the larg-
est species as tall as a Shetland pony and resembling a gazelle. Differing from other
horses, it had 3 toes on each foot. North America and Eurasia.
Naraoia: a genus of unusual trilobite-like arthropods in the order Nektaspida and
family Naraoiidae; from 1 to 2 inches long, Naraoia had a soft exoskeleton and no
thoracic segments, only the cephalon and a pygidium (or thoracopygidium) twice as

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long as the cephalon; first discovered in the Burgess Shale, it was initially thought
to be a crustacean and then was classified as a trilobite; North America, Asia, and
Australia; early Cambrian to late Silurian.
Nassellaria: an order of marine protists in the subclass Radiolaria; their siliceous
skeletons are circular and generally have spiky pseudopods; important stratigraphi-
cally where calcarous skeletons have dissolved; global distribution, Miocene to
Recent.
Nasutoceratops titusi: a species of mid-sized North American ceratopsian dino-
saurs in the subfamily Centrosaurinae. Dated at 75 Ma and known only from a par-
tial skeleton, N. titusi is considered a basal centrosaurine. It has a pair of curved,
forward-facing horns, a large nose, and a skull 5–6 feet long. Kaiparowits Formation,
Utah; Late Cretaceous.
natural selection: the basic process by which evolution occurs, combined with
small genetic variations in individual organisms. Darwin, in his formulation of the
process of natural selection, said that a species improves through the accumulation
of many small inherited variations, each of which gives the individual possessing
that variation an advantage in the competition, among all the members of the spe-
cies, for the necessities of life. The offspring of these individuals will, over some
period of time, outnumber disadvantaged individuals and at some point become a
new species.
Nautiloidea: an important, still-extant subclass of externally shelled marine ceph-
alopod mollusks that first appeared in the late Cambrian and flourished in the
Ordovician. Nautiloids were the first large group of chambered cephalopods to
develop; the chambers were separated by septa, through which a tube called a
siphuncle passed to move water from one chamber to another. Most early nautiloids,
such as Orthoceras and Ellesmeroceras, are orthoconic (straight-shelled), but
curved and coiled shells also appeared early; the Tarphycerida of the Early
Ordovician were the first of the coiled cephalopods, which became more numerous
in the Devonian and Carboniferous. Orthoconic species declined in the Triassic and
disappeared early in the Cretaceous.
The nautiloids are ancestral to the ammonoids, which appeared in the Early
Devonian and also diversified rapidly. Nautiloids declined somewhat through the
Paleozoic (probably through competition with their close relatives the Ammonoidea
and Coleoidea), but they continued to diversify. Most became extinct at the end of
the Permian, but the order Nautilida persisted, although in decreasing numbers after
the Miocene; it is still represented by two genera, Nautilus and Allonautilus. For an
early nautiloid, see Plectronoceras.
Neandertal: the name of the valley in northwestern Germany where the first
Neanderthal skeleton was found in 1856.
Neanderthal: the common name of a group of hominins who lived in Europe,
Africa, and Asia in the late Pleistocene from more than more than 300,000 years ago
to about 30,000 years ago. They are now classified either as a separate species,
N 187
Homo neanderthalensis, or as a subspecies, Homo sapiens neanderthalensis. The

Neanderthals were less dimorphic and somewhat shorter and heavier-set than mod-
ern humans, and were distinguished by a prominent brow ridge, a receding chin, and
a slightly larger cranial capacity (1450 cc) than the modern average.
Necrolestidae: a monogeneric family of small pantothere metatherian mammals;
the insectivorous Necrolestes seems to be unrelated to any modern animal;
Argentina, early Miocene,
Nectridea: an order of aquatic amphibians in the extinct subclass Lepospondyli;
generally similar to large newts, the largest grew to 3 feet long; the nectrideans are
distinguished by large protrusions on either side of their flattened skull, which
resembles a thick boomerang; North America, Europe, and Africa; Late
Carboniferous to Permian.
nektic: referring to organisms that live in the water column, swimming actively in
open water and able to move independently of water currents, as opposed to plank-
ton, which drift with the current; see also pelagic, planktic, and benthic.
nektobenthic: swimming freely but just above the bottom.
Nemagraptus: a genus of early graptoloids in the family Nemagraptidae; generally
characterized by multiramous S-shaped, reclined stipes; cosmopolitan, Middle to
Late Ordovician.
Nemagraptus gracilis: a species of Nemagraptus whose first appearance marks the
beginning of the Late Ordovician at 458.4 Ma; Sweden.
Nematophyta: a poorly known group of plants with characteristics of both vascu-
lar plants and algae; sometimes classified as a phylum containing the type genus
Nematothallus and four other genera; North America and Europe, Silurian to Early
Devonian.
Nemegtosauridae: a family of titanosaurian saurischian dinosaurs in the order
Sauropodomorpha; classification disputed, possibly related to Rapetosaurus and
Saltasaurus; Mongolia, Early to Late Cretaceous.
Nemicolopterus crypticus: a contentious species of small, toothless pterosaurs in
the suborder Pterodactyloidea, possibly related to the Tapejaridae. It had curved
toes, indicating that it was essentially a tree-dweller. Probably insectivorous, it lived
in a forested area of Western China around 120 Ma, whereas most previously known
pterosaurs have been found in coastal environments. It may be a hatchling or juve-
nile, but if it is an adult, it is one of the smallest known pterosaurs, since the holo-
type is the size of a sparrow. China, Early Cretaceous.
Neocathartes: see Bathornithidae.
Neogene: The second period of the Cenozoic Era, including the Miocene and
Pliocene Epochs.
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Neognathae: the larger of the two groups of living birds, dating back to about
100 Ma and including all living birds except the Palaeognathae (the flightless ratites
and the tinamous); despite their name, they are believed to have appeared earlier
than the Palaeognathae.
Neoproterozoic: the most recent era of the Precambrian Eon, from 1000 to
541 Ma.
Neoptera: a major taxonomic group of insects in the subclass Pterygota, q.v.; the
Neoptera include almost all of the flying insects, specifically those that can flex their
wings back over their abdomen.
Neospirifer: a genus of articulate brachiopods in the family Trigonotretidae; up to
2 inches across, characterized by robust valves and a prominent sulcus and fold;
several genera are important index fossils; global distribution, Carboniferous to
Permian.
Neosuchia: a large and very diverse clade of crocodylomorphs that includes the
extant crocodiles and their relatives as far back as the Early Jurassic.
neotype: a type specimen selected as a replacement holotype when the original has
been lost or destroyed.
Neovison macrodon: the sea mink, one of several formerly terrestrial mammals
that have evolved into a saltwater habitat; formerly called Lutreola and Mustela
vison. It bore a pelt that was 30 inches long when processed, much larger and bulk-
ier than that of the terrestrial species. Neovison lived in coastal waters off northeast-
ern North America and was thus not truly marine. Since its remains date back only
about 5,000 years, it was probably a recently differentiated species. However, it is
poorly known because of being hunted to extinction in the mid-nineteenth century
before it could be studied thoroughly.
Nerinea: a high-spired genus of marine snails in the extinct family Nerineidae;
global distribution, Jurassic and Cretaceous.
Nesophontes: the only genus in the extinct soricomorph family Nesophontidae;
commonly called West Indies shrews, these small insectivorous mammals may have
survived until the early twentieth century; possibly related to the still-extant
Solenodontidae of Cuba and Haiti; known only from the Caribbean, late Pleistocene
to Holocene.
Neurankylus: a genus of baenid turtles widespread in Western North America in
the Cretaceous; possibly related to the moderate-sized Compsemys of the Paleocene;
freshwater, with only 9 vertebrae but an extra pair of ribs; North America and Asia,
Late Cretaceous to Paleocene.
New Zealand avifauna: see Harpagornis.
Ngandong fauna: the fossil remains of several hominins found between 1931 and
1933 at Ngandong, Java, commonly referred to as the "Solo man" fossils; the site is
N 189
about 100 km from Trinil, the site of the "Java man" find; because of a cranial
capacity range of 1035–1255 cc, Solo man may represent a transitional stage
between Homo erectus and H. sapiens; late Pleistocene, between 140,000 and
500,000 years ago.
Nimravidae: a family of carnivorous mammals; not true cats like Smilodon or the
earlier machairodonts, but still classified in the order Feliformia; among their non-
felid traits are non-retractable claws and several skeletal differences; North America,
Europe, and Asia; middle to late Miocene.
Nimravides: a genus of machairodont felids; 6 feet long and about 220 pounds, it
is a sabertooth cat and not classified in the Nimravidae, which are not true cats;
North America, middle to late Miocene.
Nimravus: a genus of small saber-toothed mammals in the family Nimravidae;
cat-like but not a true felid, Nimravus was 4 feet long and probably weighed less
than 70 pounds; Western North America, early to middle Oligocene.
Niobrara Sea: see Western Interior Seaway.
Nipponites: a genus of heteromorph ammonoids in the suborder Ancyloceratina
and family Nostoceratidae, q.v. Famous for a very unusual shell form, Nipponites
resembles a tangle of knots. The tubular shell reaches lengths of almost 2 feet if
extended and might reach a diameter of 10 inches if coiled planispirally in the nor-
mal ammonoid manner. But it coils on itself on different axes, creating a globular
shape only 3–4 inches in diameter. Japan, Asia, North America, and possibly
Madagascar; Late Cretaceous.
Nodosauridae: a large family of ornithischian dinosaurs in the suborder
Ankylosauria. Herbivorous, squat, and quadrupedal, they grew as long as 20 feet.
Nodosaurs were heavily armored, generally with large shoulder spikes, but without
a tail club. North America, Europe, Asia, and Antarctica; Late Jurassic to Late
Cretaceous.
Noeggerathiales: an unusual order of fern-like vascular plants in the division
Tracheophyta and extinct class Progymnospermopsida; they have some similarities
to modern conifers and araucarians, but they may have reproduced by spores; there
was a notable 2012 discovery of a specimen in a large (20 square km) forest buried
in lower-Permian volcanic ash in China, but because of gaps in their fossil record,
they are still poorly defined and may be related to the Sphenopsida or Pteropsida;
North America, Europe, and Asia; Late Devonian to Late Triassic.
Noguerornis: one of the oldest known enantiornithine birds, a poorly known spec-
imen from the beginning of the Cretaceous; much smaller than Archaeopteryx but
more capable of flight; El Montsec, Spain, Early Cretaceous, 130–125 Ma.
nomen dubium: a name of unknown or doubtful application, often referring to a
lost type specimen.
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Nostoceratidae: a diverse family of heteromorph ammonoids in the suborder

Ancyloceratina and superfamily Turrilitoidea; most genera coil in unusual direc-
tions, sometimes partly planispirally; Nipponites resembles a randomly coiling
knot, while the tubular Bostrychoceras and the type genus Nostoceras create a heli-
cal spire shape; global distribution, Late Cretaceous.
Nostolepis: a genus of acanthodian fish in the order Climatiiformes and family
Climatiidae; a “spiny shark,” known principally from scales but thought to be as
long as 4 inches; North America, Europe, Asia, and New Zealand; late Silurian to
Middle Devonian.
Notharctus: a genus of adapiform primates of North America; related to
Smilodectes and possibly the Amphipithecidae, the lemur-like Notharctus is a well-
known example of the evolutionary trend in primates toward greater brain size,
reduction of muzzle size, and the capability for stereoscopic vision — cephalic hall-
marks of advanced primates and of humans; early to middle Eocene.
Notharctus tenebrosus: a species of Notharctus found first in Wyoming in 1870,
dated at 50 Ma; evidently adapted to a fine-branch niche in its arboreal habitat, hav-
ing long fingers, an oppposable thumb, and nails instead of claws; North America,
middle Eocene.
Nothosauroidea: an order of seal-like aquatic sauropterygian reptiles with two
suborders, Nothosauria and Pachypleurosauria. From 8 to 13 feet long, the notho-
saurs evolved from the much smaller pachypleurosaurs, which were 8 inches to 3
feet long. The nothosaurs were related but probably not directly ancestral to the
marine plesiosaurs that replaced them at the end of the Triassic (see Pistosauria).
North America, Europe, Asia, and Africa; Early to Late Triassic.
Nothrotheriidae: an extinct family of ground sloths related to the larger
Megatheriidae; they migrated to North America at the beginning of the Pleistocene
as part of the Great American Biotic Interchange; South and North America, late
Miocene to 11,000 years ago.
Nothrotheriops shastensis: the Shasta ground sloth, a species of giant sloth in the
family Nothrotheriidae; N. shastensis appeared at the beginning of the Pleistocene
and became extinct about 11,000 years ago; known from Mexico to Oregon and
especially abundant in the La Brea Tar Pits; Western North America, Pleistocene.
Notiomastodontinae: a subfamily of proboscideans in the extinct suborder
Euelephantoidea and family Gomphotheriidae; South America, late Pleistocene.
Notioprogonia: a probably paraphyletic suborder of primitive mammals in the
extinct order Notoungulata, with a more primitive dentition than that of any other
notoungulate; 3–5 feet long, some genera resembled rabbits in appearance; two
families are currently recognized, the Henricosborniidae and the Notostylopidae;
South America; Paleocene to Eocene.
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notochord: a stiff, flexible dorsal rod, the main identifying character of chordates;
the notochord is the developmental and evolutionary precursor of the vertebrate
backbone.
Notocupoides: a genus of coleopteran insects in the family Ommatidae; 4–5 inches
long, it is known only from the Madygen Formation; Kyrgyzstan, Late Triassic,
230–225 Ma.
Notoryctidae: a still-extant family of marsupial moles in the order
Notoryctemorphia; they have no external ears, and their eyes are vestigial lenses
under the skin; poorly understood but possibly related to the fragmentary Miocene
mole Yalkaparidon coheni, found in Queensland in 1985; Australia, Miocene? to
Recent.
Notosuchus: a genus of crocodylomorph reptiles in the order Notosuchia; about 3
feet long and 80 pounds, Notosuchus preyed on small animals; poorly known and
restricted to the Coniacian and Santonian Ages, 90–84 Ma; South America, Late
Cretaceous.
Notoungulata: a large order of hoofed mammals in the superorder Meridiungulata;
some genera, such as Mixotoxodon and Toxodon, were quite large, but others filled
the same ecological niches as rabbits and hyraxes; basically South American, but
some genera (see Mixotoxodon) are found as far north as the Southern United
States; Paleocene to 11,000 years ago.
Nucleolitidae: a family of generally atelostomate echinoids in the superorder
Microstomata and order Cassiduloida; the most primitive genus is Pronucleolites;
North and South America, Europe, Africa, and Asia; Middle Jurassic to middle
Pleistocene.
Nummulitidae: a family of large foraminiferans in the order Rotaliida. Lenticular
and from half an inch to 4 inches in diameter, nummulites are named for their coin
shape. Characterized by narrow, helical coils of septa, they are found principally in
shallow-water deposits of the Paleogene, but they still exist in some tropical areas.
They appear in the limestone used for the Egyptian pyramids and were probably
used as jewelry and coins in Egypt. Cosmopolitan, Early Cretaceous to Recent.
Nuna: see Rodinia.
Nyasasaurus: a genus of early dinosauriform reptiles that may be the earliest dino-
saurs; the type species, N. parringtoni, is dated at 243 Ma, much earlier than any
other of the oldest dinosaurs known (see Eoraptor, Herrerasaurus, and
Staurikosaurus); described in 2013, it has not yet been definitively classified, but
research in 2017 supports its identification as a dinosaur; Tanzania, Early Triassic.
Nyctitheriidae: a family of soricomorph mammals; scansorial insectivores, the
family is generally considered eulipotyphlan (related to shrews, hedgehogs, and
moles) but claimed by some as euarchontan (closer to tree shrews and primates);
North America, Europe, Africa, and Asia; Late Cretaceous to early Miocene.
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Nyctosaurus: a genus of pterodactyloid pterosaurs, notable for a large crest sup-

ported by two antler-like spars 1–2 feet long; smaller than its relative Pteranodon,
its body was only a foot long, but Nyctosaurus had a wingspan of up to 7 feet;
known only from Kansas and probably short-lived; United States, Late Cretaceous,
86–84 Ma.
Nymphaeales: see Pluricarpellatia.
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Obolellata: one of the three extinct classes of brachiopods in the subphylum

Rhynchonelliformea; a small class formerly labeled inarticulate but now considered
primitively articulate; early to middle Cambrian.
Obolellida: “small coin,” an order of rhynchonelliform brachiopods in the class
Obolellata, characterized by impunctate biconvex shells. Among the earliest bra-
chiopods, they were a short-lived but widespread order that began to exhibit some
articulation in the later genera. They are considered part of the Tommotian biota, the
early Cambrian wave of evolution of invertebrate animals that included the archaeo-
cyathans, trilobites, and echinoderms. Cosmopolitan, early to middle Cambrian.
Obovothyris: a genus of terebratulid brachiopods in the family Zeilleriidae; it had
a long loop and a biconvex, punctate shell; Europe, Middle Jurassic.
Ocepeia: a genus of small, primitive mammals placed tentatively in the clade
Paenungulata; known from several skull fragments, it was a cat-sized herbivore;
Morocco, middle Paleocene.
Octocorallia: a subclass of anthozoan cnidarians including the soft corals and sea
pens; generally colonial polyps without a calcareous or aragonitic skeleton; cosmo-
politan, Cambrian to Recent.
Odobenidae: see Otarioidea.
Odonata: an order of pterygote carnivorous insects that includes the dragonflies
(Anisoptera) and damselflies (Zygoptera); possibly related to the giant Meganisoptera
of the Carboniferous and Permian; traditionally grouped with mayflies and several
extinct groups in the paraphyletic classification Palaeoptera; the dragonflies and
most damselflies cannot fold their wings back over their abdomen; global distribu-
tion, Triassic to Recent.
Odontoceti: a suborder of toothed whales, such as the modern orcas, sperm
whales, beaked whales, dolphins, and porpoises; the early odontocetes appeared in

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the late Eocene, the Agorophiidae being the most primitive; the Oligocene super-
family Squalodontoidea may be ancestral to most modern odontocetes; global dis-
tribution, late Eocene to Recent.
Odontochelys semitestacea: “half-shelled turtle with teeth,” a genus of stem-
turtles in the family Odontochelyidae; the oldest known turtle, dated about 220 Ma,
just after the time when the first dinosaurs appeared; about 16 inches long, with a
plastron covering its belly but only the knobby rudiments of a carapace on its back;
found in marine deposits but probably partly terrestrial; Guizhou Province, China,
Late Triassic.
Odontognathae: an obsolete term for a group of primitive toothed birds of the
Cretaceous, similar to modern birds in having a pygostyle; current classifications
have reorganized the group in the large clade Ornithurae; North and South America,
Cretaceous.
Odontogriphus omalus: a species of soft-bodied bilaterian mollusks in the family
Odontogriphidae. The oldest mollusk known, prominent in the Burgess Shale, it
continues to figure in current debates about the origins of mollusks, brachiopods,
and annelids. Odontogriphus has an ovoid, shell-less body up to eight inches long,
with simple gill-like structures surrounding the bottom of the foot on the underside.
Its stomach, intestines, outer membrane, mouth, and radula are also visible in the
specimens. Canada, middle Cambrian, 508 Ma.
Odontopleurida: an early order of trilobites related to the order Lichida; spinose,
especially the pygidium, which is smaller than the cephalon; global distribution,
middle Cambrian to Late Devonian.
Oepikellidae: a family of sexually dimorphic ostracods in the extinct superfamily
Oepikelloidea; generally characterized by biconvex shells without a sulcus; global
distribution, Ordovician to Devonian.
Ogygiocarella: a small genus of asaphid trilobites in the family Asaphidae; famous
as an early “flatfish” discovery, it was described in 1698 by Edward Lhywd; slightly
more than an inch in length, it is isopygous, flattened, and oval in shape; Wales and
South America, Middle Ordovician.
Öhningen site: a locality on the German border with Switzerland that has pro-
duced many fossils from the Miocene, including the 3-foot-long salamander Andrias
scheuchzeri, turtles, and many well-preserved flowers and insects; also spelled
Oeningen; Germany, late Miocene.
Oldhaminidina: a suborder of strophomenid articulate brachiopods characterized
by a lobed brachial valve and convex pedicle valve; cosmopolitan, late Paleozoic.
Olenellus: a stratigraphically important genus of trilobites in the order Redlichiida
and the only currently recognized genus in the subfamily Olenellinae; generally
characterized by a very large, semicircular cephalon, elongated thorax, and very
small pygidium; North America, in the Olenellus zone of the early Cambrian.
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Olenoides: a genus of trilobites in the order Corynexochida and family

Dorypygidae; up to 4 inches long, characterized by small eyes, a raised glabella
extending to the front of the cephalon, and many slender pygidial spines; more than
100 well-preserved specimens have been found in the Burgess Shale, including sev-
eral with soft-tissue preservation; North America and Asia, middle to late Cambrian.
Olenus: a genus of trilobites in the order Ptychopariida and family Olenidae; char-
acterized by 12–15 thoracic segments and pleurae with sharp recurved tips; an
important index fossil, Olenus was widespread in the late Cambrian; the name
comes from Olenus, a character in Greco-Roman mythology who was transformed
into a stone pillar; global distribution, late Cambrian.
Oligocene: the third epoch of the Cenozoic, from 34 to 23 Ma. New modern mam-
mals were relatively rare, a fact reflected in the name, “time of few recent groups.”
Among new animal orders were the first barnacles, pigs, rats, and deer. The world
climate continued to cool, causing some extinctions and fragmentation of marine
populations. Early in the epoch, Antarctica began to be covered with ice, and the
Grande Coupure in Eurasia caused several extinctions and large-scale rearrange-
ment of biozones. Several large mammals became extinct, such as the brontotheres
and embrithopods, but ungulates in general grew larger.
Oligopygoida: a small order of irregular echinoids with a well-developed lantern;
closely related and possibly ancestral to the Clypeasteroida of the late Paleocene;
Caribbean and Mexico, Eocene to Paleocene.
Olorotitan: a genus of lambeosaurine dinosaurs in the family Hadrosauridae,
known only from an almost complete specimen found in the Amur region of Russia;
it was about 26 feet long and had an unusual fan-shaped crest; Eastern Russia, Late
Omomyidae: an important early family of tarsier-like primates in the suborder
Haplorrhini and the infraorder Tarsiiformes; the earliest-known crown primates and
probably ancestral to modern tarsiers; North America, Europe, and Asia; early
Eocene to early Miocene.
Oncorhynchus rastrosus: the largest known species of fish in the family
Salmonidae; known from two specimens from Central Oregon, they were probably
anadromous like the modern salmon; notable especially for the two 1-inch fangs
protruding to the side like warthog tusks; they grew to about 9 feet long and weighed
about 1000 pounds; United States, Miocene to Pleistocene.
ontogeny: the growth and development of individual organisms.
Onychodus: “claw tooth,” a genus of basal sarcopterygian fish in the family
Onychodontidae; poorly known but up to 12 feet long, Onychodus is characterized
by a pair of retractable tooth whorls at the anterior end of the jaw; possibly related
to Psarolepis; Canada, Europe, and Australia, Middle to Late Devonian.
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Onychonycteris finneyi: the earliest-known primitive bat, in the family

Onychonycteridae; notable in having claws on all ten fingers but probably not able
to echolocate, so taken as evidence that bats could fly efficiently before some devel-
oped echolocation; North America, late Eocene, 52 Ma.
Onychophora: the “velvet worms,” a still-extant phylum of small, caterpillar-like,
predatory animals. The modern onychophorans live generally in moist terrestrial
habitats. Their fossil record is sparse, but discoveries of the 1990s show that they
were an important and diverse group of marine invertebrates in the Cambrian.
Paleozoic onychophorans are traditionally placed in their own class, the Xenusia;
they were entirely marine and had a terminal mouth, as opposed to the ventral mouth
of modern onychophorans. From the Burgess Shale assemblage, the famous
Hallucigenia is considered an onychophoran, and Aysheaia is also thought to be
related. The Carboniferous genus Helenodora is considered basal to modern ony-
chophorans. See also Peripatus. Global distribution, Cambrian to Recent.
Onychopterella: “claw wing,” a genus of middle-sized chelicerate arthropods in
the family Eurypteridae; its last walking legs are long, and its swimming legs end in
a small claw; known mainly from the Silurian of North America, smaller specimens
(about 6 inches long) have been found in the Gondwanan Soom Shale of South
Africa; Late Ordovician to Silurian.
oogonia: see charophyte.
Opabinia: a stem-arthropod genus of soft-bodied animals found only in the
Burgess Shale; O. regalis, the only known species, had a segmented body with lobes
on the sides, a fan-shaped tail, and a long and slender proboscis; more unusual are
its five eyes and a backward-facing mouth below the head; it is tentatively placed in
the order Radiodonta; see also Jianfengia; Canada, middle Cambrian, 508 Ma.
Opallionectes: a species of Cretaceous plesiosaurs identified in 2006; known only
from Central Australia, where it was found in company with the smaller plesiosaur
Umoonasaurus; Opallionectes was 16 to 18 feet long and is notable for its large
number of needle-like teeth adapted to trapping small fish and squid; Early
operculum: a structure in many animals, usually calcitic, whose function is usu-
ally to seal an aperture by opening and closing; a prominent feature in gastropods,
ammonoids, and other cephalopods.
Ophiacodon: “snake tooth,” a genus of primitive synapsid reptiles in the family
Ophiacodontidae, possibly the ancestor of all later synapsids. About 10 feet long, it
was much larger than earlier ophiacodontids and one of the largest early synapsids,
with a large snout containing more than 160 needle-like teeth. Ophiacodon was
partly aquatic and similar to crocodiles in some ways although only distantly related,
and so an example of convergent evolution. North America and Europe, Late
Pennsylvanian to early Permian.
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Ophiderpeton: an unusual genus of snakelike lepospondyl amphibians in the order

Aistopoda; 2–3 feet long and very slender, it was mostly terrestrial and probably
fossorial; it had 200 vertebrae in its spine and no limbs; North America and Europe,
Early Carboniferous to early Permian.
Ophiocistioidea: a class of stemless echinoderms in the still-extant subphylum
Echinozoa; similar to modern sea urchins and characterized by a flattened, dome-
shaped test; global distribution, Early Ordovician to Late Cretaceous.
Ophiopinna: a genus of brittle stars, ophiuroids in the family Ophiuridae; about an
inch across, hundreds of specimens are preserved at La Voulte-sur-Rhône, some
with paddle-shaped spines, possibly an adaptation for swimming; France, Middle
Jurassic.
Ophiurida: the brittle stars, a still-extant order of asterozoan echinoderms in the
class Ophiuroidea; characterized by a sharply defined circular central disc with five
(sometimes more) slender arms that can move horizontally for locomotion; global
distribution, Devonian to Recent.
Ophiuroidea: a class of echinoderms in the still-extant subphylum Asterozoa,
encompassing the brittle stars (Ophiurida) and basket stars (Euryalida); as in the sea
stars (Asteroidea), the central body extends into five or more arms, but the ophiuroid
body is sharply distinguished from the slender, very flexible arms, which are as
much as 20 times longer than the diameter of the body; since the ophiuroids, like sea
stars, disintegrate rapidly after death, complete skeletons are rare; global distribu-
tion, Ordovician to Recent.
Opiliones: a still-extant order of arachnids characterized principally by very long
legs; although similar to spiders, they are not closely related; known as harvestmen,
they originated in the Devonian, the oldest known being Eophalangium sheari
(Early Devonian, from the Rhynie Chert); global distribution, Devonian to Recent.
Opipeuterella: a genus of nektic proetid trilobites in the family Telephinidae; it is
remarkable for very large eyes that extend below the plane of the body, indicating
that it was free-swimming; the thorax is slender and the pygidium ends in a spine;
cosmopolitan, Early to Middle Ordovician.
Opisthothelae: a suborder of arachnids in the spider order Araneae, comprising
almost all true spiders; see also Mesothelae; global distribution, Carboniferous to
Recent.
opossum: see Didelphidae.
Orbiculoidea: a large and long-lived genus of inarticulate brachiopods in the order
Lingulida; phosphatic and generally only 1 to 2 inches in diameter, with a functional
pedicle; global distribution, Early Ordovician to Cretaceous.
Ordovician: the second period of the Paleozoic Era, from 485 to 444 Ma, divided
into Early, Middle, and Late Epochs. It was a time of high sea levels and marine
transgressions until the severe glaciation at the end of the period. During the
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Ordovician, the southern continent Gondwana coalesced and drifted south, and
Laurentia, Baltica, and other land masses became separated from each other.
Between 485 and 460 Ma, about 50 million years after the great radiation of life
of the early Cambrian, marine life again experienced great changes. In a rapid and
extensive diversification of organisms in the Early and Middle Ordovician, the num-
bers of biological orders and families more than doubled, especially in brachiopods,
bivalves, corals, crinoids, bryozoans, and gastropods. This development is referred
to as the Great Ordovician Biodiversification Event (GOBE). In the Middle
Ordovician, localized faunas became established in different parts of the world, and
these faunas remained relatively stable until the great Permian extinction ended the
Paleozoic Era.
The first vertebrates with bone appeared at the beginning of the Ordovician,
including the first agnathans, the jawless fish; and the first primitive jawed fish are
known from the middle of the period. In fact, all major groups of fish originated
during the Ordovician. Although vertebrates still appeared only in marine environ-
ments, there was greater overall ecological complexity than in the Cambrian. There
was great diversity of marine invertebrates, although relatively few new invertebrate
body plans appeared. The first definitive terrestrial plants appeared, and, despite a
lack of complete fossil specimens, there is some evidence of aquatic arthropods
venturing into terrestrial environments.
Ordovician extinction: the first of the five great extinctions, at the end of the
Ordovician Period, about 444 Ma. More than one hundred families of invertebrates
and as many as 60 percent of marine genera became extinct, making it second only
to the end-Permian extinction in the number of extinct genera. As in the other mass
extinctions of the Paleozoic, this one seems to have occurred in several pulses. But
at the end of the Ordovician, more than in the Devonian or Permian, these pulses
were strongly associated with several major glaciational events.
oreodonts: see Merycoidodontidae.
Oreopithecus: “mountain ape,” a genus of Miocene hominoids; it weighed between
60 and 80 pounds and had characteristics of hominids and of dryopithecines; also
possibly related to the parapithecines of the Oligocene; known only from European
and African sites of the late Miocene.
Ornithella: a genus of articulate brachiopods in the order Terebratulida and family
Zeilleriidae; pentagonal in outline and notable for its long loop; North America,
Europe, and Asia; Middle Jurassic.
Ornithischia: the “bird-hipped” dinosaurs, one of the two large orders.
Ornithischians are characterized by a four-pronged arrangement of pelvic bones
similar to that of birds, which are generally considered descendants of coelurosau-
rian theropods. Another typical ornithischian trait is the predentary, a horn-covered
beak-like structure at the front of the lower jaw bone that probably made it easier to
crop vegetation.
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Although birds have traditionally been considered saurischians, some classifica-

tions place them in the Ornithischia, and a proposed new classification of dinosaurs
in 2017 separates the theropods into a group of their own. Almost all ornithischians
are herbivorous, the principal groups being the ankylosaurs, ceratopsians, ornitho-
pods, hadrosaurs, and stegosaurs; one group known to be omnivorous is the heter-
odontosaurids. Global distribution, Late Triassic to Late Cretaceous.
Ornithocheirus simus: “bird hand,” a species of pterodactyloid pterosaurs known
only from fragmentary English specimens of the 1860s; medium-sized, with a
wingspan of about 8 feet, and with a distinctive crest at the tip of its beak; since the
1970s, several other species once assigned to Ornithocheirus have been reassigned
or questioned; O. simus seems similar to Brazilian pterosaurs like Anhanguera, but
relationships are still unclear; England, Cretaceous, 110 Ma.
Ornitholestes: “bird thief,” a monospecific genus of bipedal theropod dinosaurs;
known only from an almost-complete skeleton found at Como Bluff, Wyoming, in
1900; lightly built and about 7 feet long, Ornitholestes hermanni had long forelimbs
and long, clawed fingers, preying mainly on small mammals and lizards; United
States, Late Jurassic.
Ornithomimus: a genus of small, bird-like dinosaurs in the family Ornithomimidae
and order Saurischia; a fast runner, superficially similar to ostriches, Ornithomimus
was 12 feet long, weighed less than 400 pounds, and was probably covered with
feathers; synonym: Dromiceiomimus; Western North America, Late Cretaceous,
77–66 Ma.
Ornithopoda: a suborder of medium to large herbivorous dinosaurs in the order
Ornithischia. They include virtually all of the bipedal ornithischians (except the
Pachycephalosauridae). The early ornithopods were small (3–10 feet long), and
later ornithopods became semi-quadrupedal, the largest (such as the Iguanodontidae
and Hadrosauridae) weighing as much as 25 tons. The ornithopods generally had
three digits, especially the later genera. Global distribution but rare in the Southern
Hemisphere, Early Jurassic to Late Cretaceous.
Ornithorhynchus anatinus: the duck-billed platypus, one of the few living mono-
tremes; semi-aquatic and egg-laying but a mammal, the modern platypus dates back
to 100,000 years ago, closely related to monotremes of the Cretaceous; found only
in Eastern Australia, the adult is 1–2 feet long and toothless, but it possesses a poi-
sonous spur on the hind foot; see also Steropodon.
Ornithoscelida: a term referring to different groupings of dinosaurs; first pro-
posed by T.H. Huxley in 1869 in his argument for seeing birds as descendants of
dinosaurs, the term has been revived in a 2017 proposal for a large-scale revision of
dinosaur classification; in this proposal the Ornithoscelida includes the Ornithischia
and the Theropoda as sister groups, the Theropoda including the probable ancestors
of birds.
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Ornithurae: a proposed term for all modern birds and their immediate ancestors,
back to the common ancestor of Ichthyornis and Hesperornis; the basic modern trait
is a short, fused pygostyle tail, as opposed to the longer, straight tails of primitive
birds.
Orrorin tugenensis: a controversial species of hominins found in the late 1990s in
Kenya; it seems to be in the line of descent from chimpanzee to advanced hominins,
possibly to humans, but is known only from fragments; between 6.1 and 5.7 Ma, 1.5
million years earlier than similar hominins.
Orsten Konservat-Lagerstätten: upper Cambrian deposits in Sweden, part of the
Alum Shale Formation. The Orsten beds have yielded important fossils since the
late twentieth century, especially of arthropod larvae and rare animals such as pen-
tastomes and tardigrades. These fossils are typically phosphatized and preserved
inside calcareous nodules, so that when the nodules are dissolved, often a three-
dimensional shell is left that preserves body structures in fine detail. Some of the
Orsten fossils have no counterparts in the fossil record — the recently discovered
pentastomes are almost the only extinct pentastomes of which there is any record,
and yet these fossil species are very similar to modern members of the phylum.
Orthoidea: a superfamily of articulate brachiopods in the order Orthida and sub-
order Orthidina; characterized by impunctate shells, an open delthyrium, and a
functional pedicle; global distribution, early Cambrian to the end of the Permian.
Orthida: an order of brachiopods in the class Rhynchonellata; the Orthida, which
include the earliest-known articulate families, arose early in the Cambrian and
diversified rapidly into the Ordovician; global distribution, early Cambrian to mid-
dle Permian.
Orthoceras: “straight horn,” a genus, now considered monospecific, of nautiloid
cephalopods in the order Orthocerida. The type species O. regulare was orthoconic:
cone-shaped, thin, and about 6 inches long. The genus was long considered much
larger, extending from the late Cambrian to the Triassic, but most species have been
reassigned. Orthocerids form Ordovician key beds in North America, Europe, and
Africa. Similar to but not related to Baculites, q.v. Global distribution, Early
Ordovician to Early Devonian.
orthocones: cephalopods whose shell is an elongated, straight, tapering cone.
orthogenesis: an obsolete biological hypothesis that organisms have an innate ten-
dency to evolve in a certain way that is determined by an internal driving force,
rather than by external processes such as natural selection; advanced in the early
twentieth century, it was soon universally refuted and abandoned; see also Gryphaea.
Orthopsidae: a family of Mesozoic euechinoids in the order Orthopsida, charac-
terized by a camarodont lantern; global distribution, Jurassic and Cretaceous.
Orthoptera: an order of neopteran flying insects that includes the suborders
Ensifera, the crickets and katydids, and Caelifera, the grasshoppers. Adapted for
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saltation, orthopterans are chewing insects with two pairs of wings; most genera
have stridulatory organs on their wings, used to produce sound when rubbed with
the hindlegs. The earliest known are the Ensifera from the Late Carboniferous.
Global distribution, Late Carboniferous to Recent.
Oryctodromeus: “digging runner,” a genus of hypsilophodont ornithischian dino-
saurs in the family Thescelosauridae, one of the earlier thescelosaurids; herbivorous
and basically bipedal but with forelimbs that seem to be adapted to digging, it is the
only confirmed burrowing dinosaur; one site contains the remains of an adult and
two juveniles in a burrow almost 7 feet long; North America, Late Cretaceous,
Cenomanian, 99–97 Ma.
Osmunda: a still-extant genus of large ferns in the order Osmundales; character-
ized by erect rhizomes and naked sporangia; it evolved in Gondwana, in the Triassic
or possibly late Permian, and spread worldwide; O. claytoniana is an example of
evolutionary stasis, having remained almost unchanged 180 million years after its
appearance in the Triassic.
ossicle: a small bony structure; see dermal ossicles.
ossicone: a horn-like protuberance consisting of a bony core permanently covered
with skin; they are found on modern giraffes and okapis and their ancestors, such as
Sivatherium, and also on other mammals, such as Uintatherium.
Osteichthyes: the bony fishes, the latest major group of fish, appearing in the late
Silurian. The osteichthyans are generally considered a superclass and divided into
two main groups, the ray-fins (actinopterygians) and lobe-fins (sarcopterygians).
The earliest-known osteichthyan is the Early Devonian Guiyu oneiros, dated around
419 Ma, and the group developed slowly in the Devonian during the dominance of
the acanthodians and placoderms. (See also Psarolepis.) But by the Carboniferous
the osteichthyans and the chondrichthyans had supplanted the earlier groups, and
they now account for more than half of all vertebrate groups.
osteoderm: a thin bony plate that forms part of the protective skin covering of
many animals, extant and extinct, especially reptilians and amphibians, but found
also in many xenarthran mammals.
Osteoglossomorpha: a still-extant superorder of teleost bony fish, among the most
primitive teleosts. The group includes some of the largest of all living (Arapaima)
and extinct (Xiphactinus) fish. One of the earliest-known genera is Lycoptera, espe-
cially important in Mesozoic biostratigraphy in China. The osteoglossids are named
“bony tongue” because many of them employ a specialized biting technique that
involves the tongue and the roof of the mouth. Global distribution but now primarily
in the Southern Hemisphere, Jurassic to Recent.
Osteolepis: “bony scale,” a genus of primitive sarcopterygian fish in the subclass
Rhipidistia and family Osteolepidae, related to the stem-tetrapods that began to
colonize the land in the Devonian; 8 inches long, it is characterized by large square
scales covered with cosmine, with small nerve canals running through the dermal
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covering; see also Eusthenopteron and Tetrapoda; Europe, Asia, and Antarctica;
Late Devonian.
osteolith: a completely mineralized fossil bone.
Osteostraci: an order of small to medium-size, fish-like jawless marine vertebrates
characterized by a large, bony headshield and a rounded mouth on the underside of
the head; the osteostracans were among the most advanced agnathans; global distri-
bution, middle Silurian to Late Devonian.
Ostlingoceras: a genus of heteromorph ammonitid ammonoids in the family
Turrilitidae; its tightly coiled helical shell resembles a long, thin gastropod; cosmo-
politan, early Late Cretaceous, 100–94 Ma.
Ostracoda: a large and very old subclass of small crustacean arthropods, many
species of which have remained relatively unchanged since the Silurian; they have
two valves and superficially resemble bivalve mollusks, but most are microscopic,
with some freshwater species as large as 30 mm; both freshwater and marine, ostra-
cods are the most common arthropods in the fossil record; global distribution,
Cambrian to Recent.
ostracoderms: “shell skin,” an informal term for several groups of armored jaw-
less fish of the early Paleozoic, usually with a bony headshield; they were gradually
replaced by placoderms and other jawed fish in the Devonian; global distribution,
late Cambrian (but first recorded in Early Ordovician) to Devonian.
Ostrea: the common oyster, a large and still-extant genus of sessile bivalves in the
order Ostreoida; global distribution, Permian to Recent.
Otarioidea: a superfamily of pinnipeds that includes the Odobenidae (walruses
and sea lions) and the Otariidae (fur seals); extinct members are Enaliarctos and the
Desmatophocidae; see also Phocidae; cosmopolitan, middle Miocene to Recent.
Otavipithecus: an obscure genus of Miocene apes, the earliest hominoid found in
subequatorial Africa; Namibia, 13 Ma.
Otodus: an early genus of large sharks in the still-extant order Lamniformes and
family Otodontidae; known from teeth and vertebrae, it is estimated to have been
about 30 feet long and a macro-predator; see also Megalolamna; global distribu-
tion, Paleocene to Eocene or Miocene.
Ottoia: a genus of predatory burrowing worms in the family Ottoiidae, considered
a stem-group priapulid; 3–4 inches long, it used its muscular toothed proboscis to
prey on hyoliths and trilobites; known definitively only from the Burgess Shale;
Ovalocephalus: a genus of phacopid trilobites in the family Pliomeridae; the body
is oval, with a strongly pentagonal glabella, small eyes, 12–20 deeply furrowed
thoracic segments, and a small pygidium; Italy, Spain, and China; Early Ordovician,
about 475 Ma.
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Oviraptor: a genus of theropod saurischian dinosaurs in the family Oviraptoridae;

generally ostrich-size but with a shorter neck and a long tail. Its name is misleading.
A spectacular find in the Gobi Desert in 1993 of an oviraptor fossilized above a nest
of 15 to 22 oviraptor eggs provided direct evidence that oviraptors brooded their
eggs and exhibited other bird-like behaviors. Although the interpretation is disputed
and the nest has been disturbed by vertisol movement, this raptor was probably
brooding rather than preying on the eggs. China, Late Cretaceous, 76–72 Ma.
Oxyaenidae: a family of relatively small carnivorous mammals formerly placed in
the polyphyletic order Creodonta but now separated in the proposed superorder
Laurasiatheria; characterized by a broad head and large jaw, with teeth designed for
crushing; plantigrade, walking flat-footed; North America, Europe, and Asia;
Paleocene to Eocene.
Oxynoticeras: a genus of ammonoids in the order Ammonitida, characterized by a
smooth, laterally compressed shell with a sharp keel; O. oxynotum is an index fossil
in the Early Jurassic; North America, Europe, and Africa; Early to Middle Jurassic.
Ozarkodina: a genus of conodonts in the order Ozarkodinida that contains impor-
tant index fossils for the late Silurian, including O. crispa, O. bohemica, and O.
snajdri; the extinction of the genus is one of the indices marking the GSSP at the
end of the Frasnian Age of the Late Devonian; see also Ancyrodella rotundiloba;
global distribution.
Ozarkodinida: an order of conodonts in the unranked clade Prioniodontida; global
distribution, Ordovician to Middle Jurassic.
Ozawainellidae: a family of foraminiferans in the order Fusulinida and superfam-
ily Fusulinoidea; North America, Asia, and Europe; Carboniferous to late Permian.
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Pachycephalosauria: “thick head,” a suborder of relatively small ornithischian

dinosaurs; bipedal and generally herbivorous, they are characterized by a very thick
skull roof; they include the genera Stegoceras, Troodon, Pachycephalosaurus, and
possibly Dracorex; North America and Asia, Late Cretaceous.
Pachycrocuta brevirostris: the giant hyena of the late Pleistocene, the largest
known at 240 pounds; it preyed on the giant elk and woolly rhinoceros, and the
Homo erectus bones of the Longgushan Cave site in China (300,000–600,000 years
ago), evidently gnawed by a predator, may bear the tooth marks of the giant hyena;
China, Pliocene to middle Pleistocene.
Pachydyptes: a monospecific genus of very large penguins (family Spheniscidae);
P. ponderosus was a flightless bird with large flippers adapted to swimming, about
5 feet long; New Zealand, late Eocene.
Pachypleurosauria: a suborder of sauropterygian reptiles in the order
Nothosauroidea; generally less than 3 feet long, slender, and lightly built, but
Keichousaurus and some other genera grew up to 9 feet long; Europe and Asia,
Middle to Late Triassic.
Pachyrhinosaurus: “thick nose,” a genus of ceratopsian dinosaurs in the subfam-
ily Centrosaurinae. From 20 to 26 feet long and weighing up to 4 tons, it is notable
for thick and ornamented bosses on the head and nose, and it also had several horns
on a large neck-frill. Several specimens have been found in Northern Alaska and
date from a time when climatic conditions were somewhat warmer than at present,
but still colder in winter than previously thought possible for dinosaurs. Alaska and
Alberta, Late Cretaceous, 74–69 Ma.
paddlefish: see Polyodon.
Pagetia: a genus of very small agnostid trilobites in the family Eodiscidae; less
than half an inch long, it had eyes, only two thoracic segments, and an annulated

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pygidium with very prominent rings; well-known from the Burgess Shale but found
worldwide, middle to late Cambrian.
Painted Hills: an area in the John Day Basin of Central Oregon that preserves a
great variety of fossils of the Eocene and Oligocene in multi-colored layers of vol-
canic ash that accumulated in the large lakes of the area; the weathering of the vol-
canic deposits produced paleosols of different types and colors.
Pakasuchus: an unusual monospecific genus of small crocodylomorphs with
mammal-like dentition and other characteristics different from most crocodiles;
considered a notosuchian, P. kapilimai was the size of an elongated house cat;
Tanzania, Early Cretaceous, 105 Ma.
Pakicetus: a genus of archaeocete whales in the family Protocetidae. Probably the
most basal whale, the quadrupedal Pakicetus was about the size of a dog and lived
primarily on land, probably semi-aquatic. Characterized by whale-like ear bones,
Pakicetus is known from several skull specimens and skeletal fragments. Pakistan,
early Eocene.
Palaeacanthaspidae: the most widespread family of primitive placoderms in the
order Acanthothoraci; armored with plates and thick spines; North America, Europe,
and Asia; Early Devonian.
Palaechinoidea: a superfamily of regular echinoids in the subclass
Perischoechinoidea; characterized by thick, polygonal, tesselate ambulacral plates
that bevel over each other to form a rigid test; rare but widespread in North America
and Europe, Silurian to early Permian.
Palaeeudyptes: a genus of large early penguins (family Spheniscidae); between 4
and 5 feet long, it was slightly larger than the modern emperor penguin; New
Zealand, South America, Antarctica, and Australia; middle or late Eocene to late
Oligocene.
Palaeobatrachus: a genus of anurans in the extinct family Palaeobatrachidae; 3–4
inches long, it was an early and long-lived genus of frogs; Europe, Early Cretaceous
to Miocene or possibly Pleistocene.
Palaeocastor: an early land beaver that excavated vertical helical burrows up to 10
feet deep; see also Agate Fossil Beds; North America and Asia, Oligocene.
Palaeocharaceae: a group of charophytic algae descended from the Middle
Devonian Eochara, both characterized by sinistrally spiralled gyrogonites; global
distribution, Early Cretaceous.
Palaeocharinus: an early genus of trigonotarbid arachnids; only a tenth of an inch
long but equipped with fangs, it preyed on insects; Scotland, Early Devonian,
408 Ma.
Palaeocircus: a genus of predatory birds in the still-extant family Accipitridae;
France, late Eocene to Oligocene.
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Palaeocopida: an order of straight-hinged ostracods, including at least the subor-

der Beyrichicopina; the assignment of other groups continues to be debated; global
distribution, Ordovician to Cretaceous.
Palaeodictyoptera: a possibly paraphyletic order of medium-sized to large insects,
the largest of five orders in the superorder Palaeodictyopteroidea. They are charac-
terized by two pairs of dragonfly-like wings, sucking mouth parts, and a pair of
winglets (paranotal lobes) on the prothorax. Several species had wingspans of up to
22 inches; among the largest known are Moravia grandis, Lithomantis carbonarius,
and Mazothairos. See also Dunbaria. Cosmopolitan, Late Carboniferous to late
Permian.
Palaeodictyopteroidea: a superorder of beaked insects in the subclass Pterygota;
formerly Palaeodictyopterida. The first herbivorous insects and indeed the first ter-
restrial herbivores, they comprised 50 percent of the known Paleozoic species of
insects. Some species were second in size only to the largest Protodonata. Their
extinction at the end of the Permian left many niches to be filled by the insects of the
Triassic. Global distribution, Middle Carboniferous to end-Permian.
Palaeodonta: a discarded term for a polyphyletic grouping of some primitive
artiodactyl mammals such as the oreodonts, all of which are now classified in either
the suborder Tylopoda or the suborder Suina.
Palaeognathae: the smaller of the two groups of living birds; they first appeared
around 70 Ma and spread worldwide but survive only in the flightless ratites and the
tinamous; almost all extant flying birds are classified as Neognathae.
Palaeoisopus: a genus of pycnogonid arthropods, possibly related to the modern
pycnogonid sea spiders; known only from the Hunsrück Slate; Germany, Early
Devonian.
Palaeomastodontidae: a family of elephantoid proboscideans currently assigned
to the Afrotheria; probably ancestral to modern elephants, they had tusks and
weighed more than 2 tons; Kenya, late Eocene to early Oligocene.
Palaeomerycidae: a family of artiodactyl ungulates in the still-extant infraorder
Pecora and superfamily Cervoidea, close to the ancestry of modern deer and giraffes;
global distribution except Australia, Eocene to Miocene.
Palaeonisciformes: a paraphyletic group of primitive actinopterygian fish in the
subclass Chondrostei; important especially because they have features that became
common in most later actinopterygians; also basic to understanding the structure of
the sarcopterygians and their descendants the terrestrial vertebrates; global distribu-
tion, late Silurian to Late Cretaceous.
Palaeoniscum: a genus of freshwater actinopterygian fish in the extinct family
Palaeoniscidae; about a foot long, it had a blunt snout and heterocercal tail fin, and
was notable for its replaceable teeth; also called Palaeoniscus, Palaeothrissum, and
Geomichthys; Europe and Asia, late Permian.
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Palaeopantopoda: an order of sea spiders, marine arthropods in the class

Pycnogonida, not arachnids but closer to them than to other arthropods; Europe,
Devonian to Jurassic.
Palaeophonus: one of the oldest known arachnids, a genus of scorpions; between
3 and 4 inches long, it had gills and is thought to have lived in water and on land,
preying on small animals and fish; North America and Europe, Silurian to
Carboniferous.
Palaeoptera: a term traditionally applied to all groups of flying insects that cannot
fold their wings back over their abdomen; the classification of the palaeopterans is
unsettled, but the group is probably paraphyletic; most palaeopterans are extinct, the
only surviving orders being the Odonata and the Ephemeroptera.
Palaeopython fischeri: a species of Eocene snakes as long as 7 feet; notable for a
three-trophic-level fossil found at the Messel Shale Pit; inside the snake’s stomach
is a lizard (Geiseltaliellus maarius), inside which is an unidentified insect; Germany,
48 Ma.
Palaeoryctidae: a family of non-placental eutherian mammals tentatively placed
in the order Cimolesta; part of the placental evolutionary radiation late in the
Cretaceous, they were shrew-like and mainly insectivorous, but their dentition
resembles that of later carnivores; North America, Asia, and Africa, Late Cretaceous
to Paleocene.
Palaeosaniwa: a monospecific genus of carnivorous lizards in the family Varanidae;
about 10 feet long, P. canadensis resembled modern monitors, and its dentition is
essentially like that of a Komodo dragon; Western North America, Late Cretaceous,
75–70 Ma.
Palaeosmilia: a horn coral, a genus of solitary rugose corals traditionally placed in
the extinct order Stauriida and family Palaeosmilidae but considered a doubtful
genus by some; characterized by radially arranged septa; type species P. murchi-
soni; Europe, North Africa, Asia, and North America; Early Carboniferous to
Permian.
Palaeospondylus: a genus of enigmatic primitive fish-like vertebrates, tentatively
placed in the class Pteraspidomorphi; it is known only from the area of Caithness,
Scotland, where hundreds of specimens have been found; it was jawless, eel-like,
and 2–3 inches long, but whether its skeleton is calcified or cartilaginous is unclear;
Middle Devonian.
Palaeotarbus: a monospecific genus of the arachnid group Trigonotarbida; P.
jerami, originally called Eotarbus, was among the first terrestrial animals; England,
late Silurian.
Palaeothele montceauensis: a species of mesothele spiders in the order Araneae;
originally named Eothele, it is the earliest-known mesothele, known from two spec-
imens from Montceau-les-Mines; France, Late Carboniferous.
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Palaeotheriidae: a family of herbivorous perissodactyl hippomorphs, close to the

ancestry of the Equidae; the type genus Palaeotherium resembled a small rhinoc-
eros and was possibly in the line of descent to horses; Europe and Asia, Eocene
through Oligocene.
paleobiochemistry: the study of chemical processes and their evolution in the fos-
sil record.
paleobioclimatology: the study of the changes in climate in the geological past
and of the effects of these changes on animal and plant life.
paleobiocoenosis: an assemblage of fossils of an interrelated community of organ-
isms that lived together.
paleobiogeography: the study of the effects on organisms of the dynamic interac-
tion of geologic processes (such as plate tectonics) with biological developments
and climatic and environmental changes.
paleobiology: the branch of paleontology concerned with the biology of extinct
organisms and their relationship to modern organisms.
paleobotany: the study of plant fossils and their interrelationships with other
organisms in the geological past; also called paleophytology.
Paleocene: the first epoch in the Cenozoic Era, from 66 to 56 Ma. The GSSP mark-
ing the beginning of the era is the worldwide occurrence of an iridium layer associ-
ated with a large asteroid impact. The end-Cretaceous extinction affected plant life
less severely than animal life, so that in the Paleocene conifers and cycads continued
to dominate in the higher latitudes. Angiosperms spread, especially in the Southern
Hemisphere; the first grasses appeared toward the end of the period. The first mod-
ern sand dollars, clypeasteroid echinoids, appeared in the sea, and ants diversified
rapidly on land. Large flightless birds appeared, and mammals diversified rapidly,
filling the many ecological niches left vacant at the end of the Cretaceous. Most
modern fish groups were present and continued with relatively little change through
the Cenozoic.
paleoclimatology: the study of past climatic conditions throughout geologic time,
and of the causes of worldwide and local climatic variation; it involves especially
sedimentology, palynology, glaciology, limnology, pedology, phytogeography, and
the study of certain marine organisms such as coral.
paleodepth: the water depth at which a geologic stratum was deposited.
paleoecology: the study of the relationships between fossil organisms and their
environment.
Paleogene: the first period of the Cenozoic Era, including the Paleocene, Eocene,
and Oligocene Epochs.
paleogeography: the study of the changes in the distribution of the earth’s land
masses caused by tectonic plate shifting in the geological past.
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In the billions of years of earth’s existence, sections of the crust have moved
constantly in the shifting of tectonic plates, sometimes called continental drift.
Throughout Paleozoic time, the most constant large land mass was the southern sec-
tion, called Gondwana; it was located generally in the lower part of the Southern
Hemisphere and included what is now South America, Africa, Antarctica, Australia,
Madagascar, and India. The Northern Hemisphere long contained only smaller frag-
ments of land — what is now North America, Greenland, Europe, and Northern
Asia.
But around 400 Ma, in a great mountain-building event called the Variscan orog-
eny, the northern fragments began to coalesce into a land mass called Laurasia and
drift southward. By the end of the Carboniferous, around 300 Ma, all of the large
land masses were locked together in the most recent global supercontinent, Pangaea,
which extended in the Permian from the South Pole almost to the North Pole. Then,
during the Late Triassic and Early Jurassic, the two sections began to drift apart,
separating the organisms on the Laurasian segment from Gondwana. The three main
sections of Laurasia — North America, Europe, and Asia — continued to drift apart
from each other in the Cretaceous, isolating the dinosaurs and other land animals on
each segment.
Since the beginning of the Cretaceous, although sporadic and fluctuating, the
movement of the continents has generally been away from each other. Late in the
Cretaceous, Africa and India broke off from Gondwana, with India moving north-
ward; Antarctica separated from Australia at the end of the Eocene, and from South
America in the early Oligocene, isolating their animals and plants. All of the present
continents existed in the Paleocene, but as they moved, some have changed in size,
notably North America and Asia, which have grown by accretion; nevertheless, the
total global land mass today is comparable to that of the Triassic.
paleoichnology: the study of trace fossils showing evidence of the actions of
organisms in the geological past, such as fossilized footprints, tracks, and burrows;
increasingly important in research on fossil ecosystems, providing evidence on the
activities and interactions of organisms in the past.
paleomorphology: the study of the structure and form of extinct plants and
animals.
paleontology: the study of ancient life in the form of fossils, especially the evolu-
tion of new species and their distribution in space and time.
paleopalynology: the stratigraphic study of fossilized pollen, spores, particulate
organic matter, and other wind-borne palynomorphs; important in the study of plant
evolution and in the correlation of geological strata; also significant in studying
climate change.
Paleoparadoxia: a genus of large to medium-sized herbivorous mammals in the
extinct order Desmostylia. A large marine relative of elephants that appeared in the
Miocene and became extinct about 11,000 years ago, Paleoparadoxia had very
large forelimbs, and its eyes were near the top of the head. Notable for the backward
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angle of the teeth, it probably ate seaweed and sea grasses. It is now thought to have
been almost fully marine like the modern sirenians. Northern Pacific coastal regions
from Japan to Alaska and as far south as Baja California.
paleopedology: the study of paleosols, fossil soils of the geological past, either as
part of a sequence of sedimentary strata or as a persistent soil surface that is no
longer actively forming.
Paleophytic: term sometimes used for a paleobotanic division of time in the early
Paleozoic Era, a time of transition from algae to gymnosperms; it was a period when
pteridophytes were abundant, from the late Cambrian up to the time of the appear-
ance of gymnosperms in the Late Devonian.
paleosol: a fossil soil horizon in the geologic past, typically altered in place by
biological, chemical, and physical processes; paleosols vary according not only to
their geological structure but also to the community of plants and animals that occu-
pied them.
Paleothyris: a genus of primitive anapsid reptiles of the late Paleozoic, tentatively
placed in the Romeriida; similar to Hylonomus, which was also found in Nova
Scotia a few million years earlier; Paleothyris was about a foot long and probably
ate insects and other small animals; Nova Scotia, Canada, middle Pennsylvanian,
312–304 Ma.
palmate: of a leaf divided into separate lobes or leaflets spreading from the base
like fingers of a hand.
palynofacies: an assemblage of palynomorphs and other organic matter in a sedi-
mentary stratum.
palynomorphs: organic microfossils from 5 to 500 microns in size, consisting of
pollen, spores, sporopollenin, silica, chitin, or pseudochitin.
palynostratigraphy: the application of palynologic techniques to the study of
geological strata; wind-borne palynomorphs, spread over great distances, are espe-
cially useful in correlating marine and non-marine sediments.
Pampaphoneus biccai: “plains killer,” a species of dog-sized carnivorous dino-
cephalians in the family Anteosauridae, known from one specimen, a skull identi-
fied in 2012 in Brazil; middle Permian.
Pampatheriidae: ancestral armadillos, a family of large armored xenarthran mam-
mals in the still-extant order Cingulata. Originating in South America in the Eocene,
some weighed more than 400 pounds. When the Panama land bridge formed in the
Pliocene, some genera emigrated to North America. All the larger ones, along with
their relatives the glyptodonts, became extinct in the Quaternary extinctions. Eocene
to Pleistocene.
Pandanaceae: a group of tropical plants in the order Pandanales; although called
screwpines, they are mostly palm-like and of medium height, and some genera are
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lianescent; tropical areas from Oceania and Australia across the Indian Ocean to
West Africa, Cretaceous to Recent.
Panderichthyidae: a small family of Devonian fish characterized by two pairs of
lobed fins and an unusual pattern of skull bones; probably ancestral to all tetrapods;
Latvia and North America, Late Devonian.
Panderodus: an early genus of conodonts in the family Panderodontidae; its ele-
ments are generally simple and horn-shaped, with fine striations, some of which are
caused by radial lamellae; Ordovician to Silurian.
Pangaea: a term for the supercontinent created by the convergence of almost all
land masses between the Late Carboniferous and Early Jurassic. See
paleogeography.
Pannotia: “all southern,” the most common name for a short-lived (roughly 650–
560 Ma) Neoproterozoic supercontinent completely within the Southern
Hemisphere, with the African craton centered on the South Pole.
Panphagia: a monospecific genus of early sauropodomorph dinosaurs in the fam-
ily Guaibasauridae; found in a middle Carnian layer of the Ischigualasto Formation,
P. protos is one of the earliest dinosaurs; named “eating all” because its dentition
suggests an omnivorous diet, it may represent a transitional stage between the car-
nivorous theropods and the herbivorous sauropods; Argentina, Late Triassic,
231 Ma.
Panthalassa: the Panthalassic Ocean, surrounding the supercontinent Pangaea
from the Permian to the Jurassic Period.
Panthera leo atrox: a species of large, lion-like carnivores in the family Felidae,
also called Naegele’s giant jaguar; short-legged and heavy-bodied, 25 percent larger
than the modern African lion, it grew up to 4 feet at the shoulder and nine feet long
head to tail; related to the Eurasian cave lion; North America, late Pleistocene to
11,000 years ago.
Pantodonta: an order or suborder of generally herbivorous eutherian mammals,
generally considered cimolestids; early forms weighed about 20 pounds, and some
later species, such as Coryphodon, weighed more than 1,000 pounds; global distri-
bution except Australia, Paleocene to Eocene.
Pantolestidae: a family of semi-aquatic, non-placental eutherian mammals in the
order Cimolesta and suborder Pantolesta; among the largest protoeutherians, some
grew to 3 feet from nose to tail, the later species resembling otters; North America
and Europe, Late Cretaceous to Oligocene.
pantotheres: a polyphyletic group of mouse- and rat-sized mammals, some of
which may be close to the line of descent of primitive marsupial and placental mam-
mals, including the Dryolestidae, Amphitheriida, Necrolestidae, and others.
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Pan troglodytes: the larger of the modern chimpanzees, which split off from their
Bonobo cousins about 1 Ma; the chimpanzees branched off the human line of
descent about 6 Ma.
Papio angusticeps: the earliest-known baboon, identified from a skull fragment
found in the cave at Malapa, South Africa, where Australopithecus sediba was
found; this fossil, 2 million years old, differs very little from the modern baboon; the
fossil record of baboons is sparse, but they seem to have appeared very recently;
Pleistocene.
Pappochelys: a genus of small diapsid reptiles that seems to be intermediate
between earlier diapsids like Eunotosaurus and primitive turtles like Odontochelys.
Its body is 6 inches long and its tail another 6 inches. Pappochelys is characterized
by two pairs of temporal fenestrae and by flattened ribs and gastralia that are begin-
ning to fuse together, a step in the process that led to the carapace and plastron of
later turtles. Europe, Middle Triassic, 240 Ma.
Pappotherium: a monospecific genus of small early eutherian mammals, known
only from a jawbone with two teeth found in Texas; possibly in the metatherian
order Deltatheroida; United States, Early Cretaceous, 113–109 Ma.
Paraceratherium: a genus of very large hyracodont perissodactyl mammals in the
extinct subfamily Indricotheriinae; resembling a large, hornless rhinoceros, it was
one of the largest land mammals ever, 18 feet high at the shoulder, 26 feet long, and
weighing 25–30 tons; also called Indricotherium and Baluchitherium; Asia, early
Oligocene to early Miocene.
Paracrinoidea: “near crinoid,” a small class of blastozoan echinoderms in the sub-
phylum Crinozoa; they superficially resemble stalked crinoids in having a stem,
thecae, and arms; North America and Europe, Early Ordovician to early Silurian.
Paradoxides: a genus of large early trilobites in the order Redlichiida, the largest
species over 3 feet in length; also characterized by a very long thorax and two spines
longer than its body, extending backward from the cephalon; index fossil for the
Avalonian beds of the middle Cambrian; North America, Europe, and North Africa;
early to late Cambrian.
Parahagla: see Habrohagla.
Parakidograptus acuminatus: a species of pterobranch graptolites in the order
Graptoloidea; its first appearance marks the beginning of the Silurian at 443.8 Ma;
Great Britain.
Paralititan strömeri: a species of titanosaurian dinosaurs, an extremely massive
sauropod found in 1999 at the Bahariya oasis in Egypt, near where Ernst Strömer
had made important finds in the early twentieth century. A six-foot humerus and
other upper-body bones make it clear that it was one of the largest dinosaurs.
Estimated to be 85 feet long and weighing 65 tons, it may be almost as large in mass
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as another titanosaurian, Argentinosaurus. Egypt, Late Cretaceous, 94 Ma, not long

after Africa and South America separated.
Paramyidae: a family of the earliest-known rodents, in the suborder Sciuromorpha;
scansorial and squirrel-sized, they may have lived in trees and competed with early
primates; probably the basal stock of the order Rodentia; North America, Europe,
and Asia, late Paleocene to late Eocene.
Paramylodon: a genus of giant ground sloths in the order Pilosa and the extinct
family Mylodontidae; about 10 feet long and weighing more than a ton, it is also
notable for the dermal ossicles embedded in its thick skin; formerly called Mylodon,
it is known from widespread North American deposits from Canada to Central
America; late Pliocene to late Pleistocene.
Paranthropus: a genus of hominins similar to and possibly descended from gracile
australopithecines; known from several skull and other fragments dated between 2.3
and 1.2 Ma; East Africa.
Paranyctoides: a genus of Late Cretaceous eutherian mammals; the earliest-known
erinaceomorph, the scansorial Paranyctoides has some characteristics of true insec-
tivores; North America and Uzbekistan, 89–66 Ma.
Paranyroca: a genus of anseriform birds in the extinct family Paranyrocidae or
family Anatidae; known from two specimens found in North Dakota; United States,
middle Miocene.
Paraparchitoidea: a superfamily of robust ostracods in the extinct order
Kloedenellocopina; global distribution, Devonian to late Permian.
paraphyletic group: see polyphyletic group.
Parapithecidae: a family of primitive anthropoid primates in the superfamily
Parapithecoidea, known especially from the middle Oligocene Fayum deposits of
Egypt. The parapithecines are difficult to classify but may be the most primitive
advanced primates known. One member of the family, Apidium, may be ancestral to
Oreopithecus, but its affinities are still unclear. Africa and Asia, Eocene to Oligocene.
Parapuzosia: a genus of very large ammonoids in the order Ammonitida and fam-
ily Desmoceratidae; the largest known ammonoids, with fossil shells almost 7 feet
across; global distribution, Late Carboniferous.
Parasaurolophus: a genus of very large duck-billed herbivorous dinosaurs in the
family Hadrosauridae, characterized especially by the long, distinctive crest from
the nose to behind its head. Several theories try to explain the function of this
unusual crest, such as to generate trombone-like sounds, to support a decorative flap
of skin, or to provide a sign of sex recognition. These hadrosaurs are known from a
few specimens of three species; the type species, P. walkeri, was about 31 feet long
and weighed almost 3 tons. North America, 76 Ma.
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Paraschwagerina: a genus of fusulinid foraminiferans that are index fossils for the
early Permian; they occupied a deep marine environment, far offshore; North
America, Europe, and Asia; early to middle Permian.
Paraselkirkia: a genus of predatory priapulid worms in the family Selkirkiidae;
similar but larger (6 inches) than the Burgess Shale Selkirkia, q.v.; known only from
the Chengjiang biota, China, early Cambrian.
Parasemionotidae: a family of small, bony actinopterygian fish in the subclass
Neopterygii and extinct order Amiiformes; global distribution, Early to Middle
Triassic.
Parasuchia: see Phytosauria.
paratheres: see Xenarthra.
Parathuramminoidea: a superfamily of foraminiferans in the suborder Fusulinina;
characterized by a test with a globular or tubular chamber; global distribution,
Ordovician to Permian.
paratype: a specimen of a type species other than the holotype; several paratypes
may be used in identifying the species.
Parazoa: a subkingdom of eukaryotic multicellular animals whose only known
living members are the sponges and the placozoan Trichoplax; parazoans early
evolved into a line separate from all other animals, splitting off from the Eumetazoa
in the early Neoproterozoic, about 940 Ma; although they have differentiated cells,
they have no organs or tissues.
Pareiasauridae: a family of bulky, armored herbivorous reptiles in the suborder
Procolophonomorpha; formerly included in the paraphyletic order Captorhinida; as
long as 10 feet and as heavy as 1,300 pounds, some species are notable for a great
number of scutes covering the body and for heavy cephalic ridges and knobs; Africa,
Europe, and Asia; middle to late Permian.
Paris Canyon: a site in Idaho being developed in 2017, dated at only 1.3 million
years after the end-Permian extinction; its fossil assemblage may indicate a more
rapid recovery of marine life after the extinction than previously thought; United
States, Early Triassic.
Parkinsonia: a genus of ammonoids in the order Ammonitida and superfamily
Perisphinctoidea, with a laterally compressed and strongly ribbed shell; index fossil
for the Bajocian age; North America, Europe, and Asia; Middle Jurassic.
Parksosaurus: a genus of hypsilophodont ornithopod dinosaurs in the family
Thescelosauridae; it resembles Thescelosaurus, another ornithopod also from the
Maastrichtian; about 8 feet long and bipedal, it was a slender, fast, herbivorous ani-
mal; Alberta, Late Cretaceous, 70 Ma.
Parvancorina: “small anchor,” a genus of enigmatic Ediacaran animals with some
arthropod-like characters; known from circular molds, less than an inch wide, and
216 P
characterized by a central ridge with a raised semicircular arc at one end of the
ridge; similar in shape to trilobites but probably unrelated; found in marine sedi-
ments in Russia and Australia, and also on the underside of some Australian paleo-
sols; 560–555 Ma.
Patagonykus: a monospecific genus of theropod dinosaurs in the family
Alvarezsauridae; known from a partial skeleton, P. puertai was about 7 feet long;
Argentina, Late Cretaceous, 94–87 Ma.
Patagotitan mayorum: a species of large titanosaurian sauropod dinosaurs
described in 2017; known from partial skeletons of seven individuals, it is estimated
to have been 120 feet long and weighing 69 tons, but these estimates may be revised
downward; Argentina, Early Cretaceous, 102 Ma.
Patellina (foraminiferan): a still-extant genus of dish-shaped spirillinid forami-
niferans in the family Patellinidae; global distribution, Late Jurassic to Recent.
Patellidae (gastropod): a still-extant family of gastropods in the order
Eogastropoda; imperforate limpets, specialized for clinging to rocks under their
cap-shaped shell; global distribution, Permian to Recent.
Paterinida: an order of early inarticulate brachiopods in the subphylum
Linguliformea and class Paterinata; characterized by biconvex, rounded, generally
phosphatic shells; global distribution, early Cambrian to Middle Ordovician.
Patriomanis: a monospecific genus of pangolins in the suborder Eupholidota and
family Patriomanidae; P. americana, well-known from several partial specimens
found between 1970 and 2016, is the only known North American pangolin,
although modern pangolins are widespread in Asia and Africa; see also Eomanis;
late Eocene to Oligocene.
peak zone: a stratigraphic zone in which a specific organism occurs in its greatest
abundance, or which is dominated by an exceptional abundance of one or more
types of organisms.
Pecopteris: a common form genus of fossil foliage especially from the marattia-
lean fern Psaronius, but also from a seed fern and several filicalean ferns; especially
prominent in the Carboniferous and Permian; global distribution, Devonian to
Miocene.
Pectinidae: the scallops, a family of bivalves in the superfamily Pectinoidea. The
Pectinidae are characterized by a single adductor muscle and a basically triangular
shell, which is usually ridged and from 1 to 5 inches wide. They are benthic, but
most genera differ from other bivalves by swimming freely. Although the Pectinidae
were the most diverse Mesozoic bivalves, they became almost extinct at the end of
the Cretaceous but then again radiated rapidly in the Cenozoic. Global distribution,
Triassic to Recent.
Pederpes finneyi: a species of stem-tetrapod amphibians in the family
Whatcheeriidae; an important specimen for its occurrence in the period of time
P 217
called Romer’s Gap; about 3 feet long and with a large, somewhat triangular head,
this tetrapod has several features indicating greater terrestriality than any earlier
animal; Scotland, Early Carboniferous, about 348 Ma.
Pedetidae: a still-extant family of African rodents in the order Sciuromorpha; the
modern springhares differ only slightly from the earliest Pedetidae, which arose in
the Miocene.
pedicle: the stalk that is present in most brachiopods, attaching the pedicle valve to
a hard substrate, usually below the animal.
Peking Man: see Homo erectus pekinensis.
pelagic: living in open water, typically in the water column of the ocean, neither
near shore nor at the bottom; pelagic animals include birds as well as free-swimming
or floating organisms; see also nektic, planktic, and benthic.
Pelagornis: a widespread genus of large sea birds of the Neogene; one of the
Miocene species, P. sandersi, had probably the widest wingspan known in a bird, as
much as 24 feet (see also Argentavis magnificens); Pelagornis is notable for the
bony spikes in its mouth which it used to grasp its prey, live or dead fish; global
distribution.
pelmatozoan: “stem animal,” a term formerly referring to the paraphyletic sub-
phylum of echinoderms called Pelmatozoa, which were attached to the sea substrate
by a stem; the term is still used informally to refer to organisms with this habit;
global distribution, early Cambrian to Recent.
Peltephilus: a genus of fossorial horned armadillos, South American xenarthran
mammals in the still-extant order Cingulata and family Chlamyphoridae. It was
characterized by large claws and bony scutes on its head and bands of scutes across
its back; it also bore at least one pair of small horns on the top of its snout, forward
of the eyes. The function of the horns is unclear, although they offered at least some
defense for the eyes. These horns represent an example of convergent evolution in
two unrelated animals since the size and placement of the horns is similar to that of
Ceratogaulus, the horned rodent of the Neogene of North America and the only
other known fossorial horned mammal. Argentina, Chile, and Bolivia; Oligocene to
middle Miocene.
Peltobatrachus: “shielded frog,” a genus of armored temnospondyl amphibians in
the family Peltobatrachidae; about 2 feet long, it was armored with a series of nar-
row plates covering its entire body; Africa, late Permian.
pelycosaurs: “bowl lizard,” now an informal term for a paraphyletic stem group of
synapsids that were the dominant animals of the Carboniferous and Permian.
Among the earliest amniotes, pelycosaurs arose in the Pennsylvanian and are con-
sidered ancestral to therapsids and eventually to mammals. Their bowl-shaped pel-
vic structure gave them greater agility and speed on land than earlier reptiles like
Hylonomus. They are most noticeable for the high and partly vascularized sails
218 P
stretched on elongated neural spines down the back of several genera; in species like
Dimetrodon and Edaphosaurus, these formed a spectacular sail of several square
feet. Another important character of some pelycosaurs was one or two pairs of lon-
ger, grasping teeth at the front of the snout, the precursors of the canine teeth that
have been one of the hallmarks of the synapsid-mammal lineage ever since. North
America and possibly Europe, Early Pennsylvanian to late Permian.
Pengana: a genus of predatory birds in the still-extant family Accipitridae;
Australia, early Miocene.
Pennatulacea: the sea pens, an order of soft-bodied, generally feather-shaped
anthozoan cnidarians in the subclass Octocorallia; colonies of dimorphic polyps,
most of them attached to the substrate by a stalk, but some capable of movement;
global distribution, Cambrian to Recent.
Pennsylvanian: see Carboniferous.
Pentacrinites: a genus of articulate stemmed crinoids in the order Isocrinida, char-
acterized by a small cup and five long, repeatedly branching arms at the end of a
stem as long as 3 feet; the stem is pentagonal in cross-section; thought to have
grown on driftwood and thus be semi-planktic; global distribution, Middle Triassic
to Eocene.
Pentamerida: an order of biconvex articulate brachiopods in the class
Rhynchonellata, characterized by impunctate, somewhat pentagonal shells and a
short hinge line; global distribution, middle Cambrian to Late Devonian.
Pentastomida: the “tongue worms,” a small and controversial phylum of parasites
on terrestrial animals and on some fish. Characterized by internal plates separating
the body cavity pentamerally when seen in cross-section. Many biologists have con-
sidered them to be an offshoot of the branchiurans, a group of crustaceans. Since
1994, however, discoveries of several pentastomes in upper Cambrian strata in
Canada and the Orsten Lagerstätten of Sweden (see also Heymonsicambria and
Linguatula) have shown that the pentastomes have persisted almost unchanged
until the present, convincing many authorities that they should be accorded phylum
status. Global distribution, late Cambrian to Recent.
Pentecopterus decorahensis: the earliest-known species of eurypterids, sea scor-
pions, discovered in the early twenty-first century near Decorah, Iowa; as long as 6
feet, they were armed with a dozen clawed arms and had a three-foot tail used in
swimming; unlike some larger Paleozoic sea scorpions, they were efficient preda-
tors; North America, Middle to Late Ordovician, 460 Ma.
Pentoxylales: an order of gymnosperms that were small shrubs or trees. Their
affinities are unclear, but they are related to ginkgos and may be in the line of
descent to angiosperms. A stem genus Pentoxylon and a seed-bearing cone genus
Camoconites have been described for the genus Taeniopteris. The Pentoxylales
were an important part of the Gondwana flora in the Jurassic. Known only from
India and New Zealand, Jurassic and possibly Early Cretaceous.
P 219
Pentremites: a genus of blastoid echinoderms in the order Spiraculata and family

Pentremitidae; attached to the substrate by a slender stalk, from 4 to 12 inches long;
North America, Early Carboniferous.
peppered moth: see Biston betularia.
Peradectes: a genus of early marsupials in the family Peradectidae; the type spe-
cies, P. minor, was only 8 inches long from head to tail; thought to be a scansorial
omnivore, it appeared just after the end-Cretaceous extinction; North and South
America, early Paleocene.
periodic extinction: the theory that major extinctions have occurred since the
Permian at regular intervals of about 26 million years.
Peripatus: a still-extant genus of velvet worms (phylum Onychophora) in the fam-
ily Peripatidae; similar to Aysheaia of the middle Cambrian but Paleozoic collection
failure leaves the possible relationship unclear; Central and South America,
Myanmar, Australia, and New Zealand; Late Cretaceous to Recent.
Periptychidae: an early family of tapir-like protoungulate placental mammals;
characterized by vertical, wrinkling ridges on their molars, and ranging in size from
30 to 200 pounds, some periptychids were omnivorous and others were herbivo-
rous; North America and possibly Eastern Asia, Late Cretaceous through Paleocene.
perissodactyls: ungulates having an odd number of toes, especially herbivorous
terrestrial mammals having a large central digit on each foot, such as the horse,
tapir, and rhinoceros; the earliest perissodactyls appeared in the late Paleocene (see
Artiodactyla and Hyracotherium).
Perittodus apsconditus: “hidden odd tooth,” one of several new stem-tetrapods
discovered by Jennifer Clack in 2016; its dentition is complex and different from
other tetrapods; see also Tetrapoda; England, Late Devonian.
Permarachne novokshonovi: a species of proto-spiders in the order Uraraneida
and family Permarachnidae; known only from a Russian specimen (a molt) described
in 2005; see also Attercopus; Ural Mountains, Permian, 280 Ma.
Permian: the final period of the Paleozoic Era, from 299 to 251 Ma. Although it
was a relatively hot and dry time, sea levels sank, evidently because of ocean floor
changes accompanying the formation of the supercontinent Pangaea. Pelycosaur-
grade synapsid reptiles survived from the Late Carboniferous into the Permian, fol-
lowed by pareiasaurian reptiles and the first therapsids, which diversified rapidly
toward the end of the period. Cycads and conifers became widespread as the drying
climate gradually ended the coal swamps of the Carboniferous; the Glossopteris
flora dominated Gondwana plant life. The period ended with the most severe mass
extinction in Earth’s history, setting the stage for the dinosaurs of the Mesozoic.
Permian extinction: the most severe of the five great extinctions, marking the end
of the Paleozoic Era at 252.2 Ma. Like the four other mass extinctions, the Permian
event may have occurred in several phases. Two smaller extinction pulses occurred
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at 259.8 and 254.1 Ma, marking the end of the Capitanian and Wuchiapingian
Stages respectively; both may have contributed to the more extensive extinction at
252.2 Ma, referred to as the end-Permian extinction.
At least 90 percent of marine species became extinct, while the terrestrial extinc-
tions were less severe but still estimated to be 75 percent of species, especially the
large herbivores. Most lineages of insects were not severely affected except for the
pterygote superorder Palaeodictyopteroidea, which became extinct at about 252 Ma.
Among the marine groups that did not survive were the trilobites, rugose corals, and
eurypterids, and among those that survived in only one or two orders were the bra-
chiopods, crinoids, and nautiloids. After the Permian extinction, mollusks came to
rival arthropods for dominance in marine environments, and modern corals and
bryozoans replaced the Paleozoic types.
Several factors have been suggested as causes, but volcanic eruptions (see
Siberian Traps) are increasingly stressed as the primary cause, especially because
of high levels of carbon dioxide emitted into the atmosphere. There was clearly
global environmental degradation, including low levels of oxygen in the oceans,
evident in widespread sediments. Other possible causes are global cooling and the
disappearance of many nearshore environments as the continents coalesced into
Pangaea.
Peronopsis: a genus of trilobites in the order Agnostida and family Peronopsidae;
isopygous, similar to Itagnostus; global distribution, middle to late Cambrian.
Perudyptes: a genus of large penguins in the family Spheniscidae; flightless and
adapted for diving like all penguins, Perudyptes was about the size of the modern
emperor penguin and is one of the earliest definitive penguins known, dated at 42
Ma; Peru, middle Eocene.
pes: plural pedes, the distal part of the hindlimb of bipedal vertebrates, corre-
sponding to the human foot.
Petalichthyida: an order of flattened placoderms, characterized by cranial and
trunk shields, and a partly ossified skeleton; probably benthic; Australia, Asia,
Europe, and North America; Devonian to Carboniferous.
petalodont dentition: teeth that are specially hardened, flat-crowned, and embed-
ded in the palate, an adaptation for grazing on tough organisms like crinoids,
sponges, and other encrusting animals.
Petalodontidae: a family of primitive skate-like holocephalic fish in the subclass
Chondrichthyes and order Petalodontiformes; their classification is obscure because
they are known primarily from their petalodont teeth; Belantsea and Janassa are the
best-known genera; North America and Europe, Carboniferous to Permian.
Petrolacosaurus kansensis: the earliest-known diapsid, a species of North
American reptiles in the order Araeoscelida; about 16 inches long and insectivo-
rous; Kansas, Late Pennsylvanian, 302 Ma.
Peytoia: see Anomalocarididae.
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Pezophaps solitaria: the Rodrigues solitaire, a species of birds in the order

Columbiformes and extinct subfamily Raphinae; Pezophaps (“pedestrian pigeon”)
is a monospecific genus but is closely related to Didus solitarius, the dodo.
Pezophaps, the size of a swan and flightless, was endemic to the island of Rodrigues,
in the Indian Ocean east of Madagascar. It was discovered in 1691 and was hunted
to extinction within a century.
Pezosiren portelli: “walking sirenian,” the earliest-known fully quadrupedal sire-
nian, placed in the family Prorastomidae with the fully aquatic Prorastomus.
Probably amphibious and similar in habit to the modern hippopotamus, it represents
an important transitional stage between fully terrestrial mammals with four legs and
fully aquatic mammals with flippers. Jamaica, early Eocene.
phaceloid: of colonial corals, having tubular corallites with separate walls, joined
together at the base.
Phacopida: “lens-face,” a large and diverse order of trilobites with three subor-
ders: Phacopina (with schizochroal eyes), Cheirurina (holochroal eyes), and
Calymenina (holochroal eyes); global distribution, late Cambrian to the end of the
Devonian.
Phacops rana: a famous species of trilobites in the order Phacopida, notable for its
very large eyes and the warty surface of the glabella; North America and Africa,
Middle Devonian.
Phalangiotarbida: an order of early arachnids, considered a sister group of the
Opiliones but possibly related also to the Acari; Europe and North America,
Phanerorhynchus: a genus of chondrostean fish in the family Phanerorhynchidae;
characterized by large body scales, unjointed fin rays, and vertical jaw suspension;
Europe, Late Carboniferous.
Phanerozoic: “visible life,” the geologic eon from the beginning of the Cambrian
to the present.
Phareodus: a genus of freshwater osteoglossomorph fish in the family
Osteoglossidae; 6–7 inches long, it was known in two species, P. encaustus and P.
testis; North America, Europe, Asia, and Australia; Cretaceous to Eocene.
Phascolarctidae: “pouch bears,” a still-extant family of marsupial mammals in the
order Diprotodontia; it includes only one extant species, the modern koala
(Phascolarctos cinereus), but about eight extinct genera are known, including P.
stirtoni, which became extinct in the late Pleistocene; arboreal herbivores weighing
as much as 50 pounds, the family originated in the late Oligocene and is closely
related to the Vombatidae; Australia, Oligocene to Recent.
Phascolonus: a genus of herbivorous Australian marsupials in the order
Diprotodontia and family Vombatidae; one of several extinct genera of large
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wombats, Phascolonus weighed about 450 pounds; it is known from the middle
Pliocene to the middle or late Pleistocene.
Phenacodontidae: an early family of herbivorous mammals, possibly ancestral to
later mammals; their size increased from 15 to 120 pounds from the Paleocene to
the Eocene; the type genus Phenacodus, resembling Hyracotherium but slightly
stockier, was one of the earliest and most primitive of the ungulates; North America
and Europe, Paleocene and Eocene.
Phillipsastreidae: a family of generally colonial rugose corals in the order
Stauriida and suborder Columnariina; its corallites did not have separate walls, but
had radially arranged and sometimes curving septa, resulting in a star-shaped pat-
tern; global distribution, Early to Middle Devonian.
Phillipsia: a genus of small trilobites in the order Proetida and superfamily
Bathyuroidea; isopygous, with a large cephalon and a large pygidium; although not
numerous, it was one of the last trilobites; North America, Europe, and Asia;
Carboniferous to late Permian.
Phiomyidae: a small family of rodents in the suborder Hystricomorpha; Africa,
late Eocene to middle Miocene.
Phoberomys: a genus of giant rodents in the suborder Hystricomorpha and family
Dinomyidae. About the size of a buffalo, almost 10 feet long and weighing up to
1500 pounds, it was one of the largest rodents known. (See also Josephoartigasia.)
It had small forelimbs and teeth typical of herbivores, and was notable for a long tail
which it could use to balance on two feet to look over the vegetation of the swamp-
lands where it lived. Northern South America, known only from the late Miocene.
Phocidae: the true seals, a family of pinnipeds in the superfamily Phocoidea; prob-
ably descended from Enaliarctos and possibly related to the ursids; earless and
overall more specialized for aquatic life than their relatives, the Odobenidae and
Otariidae; global distribution, middle Miocene to Recent.
Pholadomya: “hole clam,” a still-extant genus of inarticulate marine bivalves in
the superorder Anomalodesmata and order Pholadomyoida, widespread in the
Mesozoic; deep-burrowing, up to 5 inches long; the shell gapes open at the posterior
end so that the siphon can be partially withdrawn; Late Triassic to Recent.
Pholidophorus: a genus of small, primitive teleost fish in the order
Pholidophoriformes; one of the earliest teleosts, it had ganoid scales and its skeleton
was partly bone and partly cartilage; South America, Europe, and Africa; Middle
Triassic to Late Jurassic.
Pholidota: the pangolins, an order of mammals once distributed globally and dat-
ing back to the Paleocene; now extant only in Africa and Southeast Asia.
Phoronida: a still-extant but very small phylum of lophophorates related to bra-
chiopods and bryozoans. They are tube-dwelling vermiforms that filter-feed by
means of the cilia on a crown of tentacles. From 1 to 20 inches long, they have no
P 223
hard parts, and there are no undisputed fossil specimens except as trace fossils in the
form of the chitinous tubes that they construct as burrows. Living phoronids burrow
into both soft and hard substrates. Global distribution, Early Devonian to Recent.
Phorusrhacidae: a family of large, flightless carnivorous birds in the order
Cariamiformes. Characterized by a large head and neck, with short wings used only
for balance, they averaged about 5 feet high. The phorusrhacids lived on the South
American continent while it was an island in the early Cenozoic. Probably seizing
and killing their prey by battering it on the ground, they were the principal carni-
vores of the continent until it joined North America in the late Pliocene, when saber-
toothed cats, jaguars, and wild dogs migrated southward. The largest of the
phorusrhacids, Kelenken, found in Patagonia in 2004, grew as high as 10 feet and
weighed as much as 500 pounds. South America, early Paleocene to late Pliocene.
Phorusrhacos longissimus: a genus of carnivorous birds in the family
Phorusrhacidae; notable for a long skull and long jaw with a hooked beak; South
America, Miocene.
phyletic gradualism: a model of evolution that stresses steady, gradual evolution-
ary change (see anagenesis) as the basic mode of speciation; this gradual evolution-
ary change is considered to occur throughout the history of the lineage.
Phyllocarida: a still-extant subclass of crustaceans in the class Malacostraca,
characterized by a folded bivalved carapace that covers almost the entire body; it
includes three orders: the extinct Hymenostraca and Archaeostraca, and the extant
Leptostraca, a small but long-lived group that appeared in the Cambrian; most other
crustaceans are classified in the Eumalacostraca; global distribution.
Phylloceras: a genus of nektic ammonoids in the order Phylloceratida; the smooth
shell, 4–10 inches long, is laterally flattened with no ornamentation; global distribu-
tion, Late Triassic to the end of the Cretaceous.
Phylloceratida: a conservative Mesozoic order (or suborder) of ammonoids; gen-
erally smooth or weakly ornamented, they are marked by a complex phylloid suture;
the phylloceratids are probably ancestral to the highly diverse Jurassic and
Cretaceous ammonoids and themselves persisted almost unchanged from the Early
Triassic to the Late Cretaceous; global distribution.
Phyllograptus: a colonial graptolite of the Ordovician in the order Graptoloidea
and family Phyllograptidae; distinguished by the oval, leaf-like shape of the colony,
which was 1–2 inches long; several species are index fossils for the Early Ordovician;
global distribution, Early to Late Ordovician.
phylloid: leaf-like, in the shape of a leaf.
Phyllolepida: “leaf scale,” an order of small, flattened placoderms, possibly related
to the arthrodires; the phyllolepids were freshwater predators with whole-plate
armor, a wide mouth, and probably only vestigial eyes; several genera in three fami-
lies are known globally from the Early and Middle Devonian, but Phyllolepis was
224 P
the only genus to survive into the Famennian, the final stage of the Devonian; Early
to Late Devonian.
Phyllotheca: a widespread genus of jointgrasses in the order Equisetales; global
distribution, Late Carboniferous to late Permian.
phylogeny: the evolutionary history of a biological organism and its relationship
to other organisms.
Physalis: see Solanaceae.
phytolith: “plant stone,” a microscopic siliceous structure persisting in plant tissue
after the decay of the plant; a common element in the fossil record, phytoliths may
identify changes in paleoenvironments, and they may also reveal information about
the diet of herbivorous animals; see also grasses.
Phytosauria: an order of semi-aquatic archosaurian reptiles, about 6 feet long,
heavily armored, and with an elongated snout; formerly called Parasuchia. An
example of convergent evolution, the phytosaurs closely resembled the unrelated
crocodiles, but their ancestry is obscure. They were widespread in the northern con-
tinents but unknown in Southern Africa, South America, Australia, and Antarctica.
Middle to Late Triassic, possibly Early Jurassic.
Pierolapithecus catalaunicus: an ape fossil discovered in 2004 near Barcelona
that had the upright posture of human beings, an ape-like body, and fingers like
those of chimpanzees; dated at 13 Ma, it is unique and a possible step in the evolu-
tion of ape to human; only one specimen has been found; Spain, Miocene.
Pikaia: an enigmatic genus of small primitive chordates known from the Burgess
Shale. It was first considered a polychaete worm or stem-arthropod but was later
identified as perhaps the oldest ancestor of chordates; it is now generally assigned
to the subphylum Cephalochordata. Laterally compressed and ribbon-shaped, the
soft-bodied P. gracilens was only about 2 inches long. Canada, middle Cambrian,
508 Ma.
Pilina: a genus of monoplacophorans in the family Tryblidiidae; United States and
Europe, Ordovician to Silurian.
“Piltdown Man”: a sophisticated hoax perpetrated in 1912 by Charles Dawson,
who claimed to have discovered a missing link between apes and humans; the fraud
was discovered forty years later when advanced dating techniques showed that the
specimen was actually only 50,000 years old; subsequent examination revealed that
it was a deliberate combination of a human cranium with the jaw of an ape, probably
an orangutan.
Pinacosaurus: a genus of ornithischian dinosaurs in the family Ankylosauridae;
herbivorous, 16–20 feet long with a spiked tail club, it lived between 80 and 75 Ma;
Mongolia, Late Cretaceous.
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Pinopsida: a still-extant class of cone-bearing gymnosperm trees, the most suc-

cessful group of gymnosperms still living; the earliest conifers of the Middle
Carboniferous and early Permian had sub-compact cones but may have been the
first plants whose seeds could remain dormant for a considerable time before germi-
nation; see also Araucaria and Lebachia; global distribution, Middle Carboniferous
to Recent.
Pisco Formation: an important Konzentrat-Lagerstätte on the southern coast of
Peru; known for the only definitive specimen of the large whale Livyatan as well as
the aquatic sloth Thalassocnus and other well-preserved specimens of whales, pen-
guins, marine crocodiles, and sharks; South America, middle Miocene to middle
Pliocene.
Pistosauria: an order of marine reptiles in the superorder Sauropterygia; about 10
feet long and characterized by an elongated neck and skull, they were probably
transitional between nothosaurs and the larger plesiosaurs; North America, Europe,
and Asia; Early Triassic to Late Cretaceous.
Pithecanthropus: an obsolete term, once applied to several hominins that are now
classified as Homo erectus.
Pityaceae: a family of plants of the Carboniferous long thought to belong to the
Cordaitales but now assigned to the Pteridospermophyta; global distribution, Early
Carboniferous.
Placenticeras: a genus of large ammonitid ammonoids in the family
Placenticeratidae and superfamily Hoplitoidea; relatively smooth-shelled, they
ranged from 1 to 3 feet across; global distribution, Late Cretaceous.
Placerias: a genus of dicynodont therapsids in the family Stahleckeriidae; it was
among the largest herbivores of the Triassic and one of the last surviving dicyn-
odonts; from 6 to 11 feet long and weighing up to 4,400 pounds, Placerias was
beaked with two strong upper tusks; its lifestyle was similar to that of a hippopota-
mus, spending much of its time in the water; Southwestern United States, Late
Triassic.
Placodermi: “plated skin,” the earliest class of gnathostome vertebrates, primitive
jawed fish of the middle Paleozoic; from a few inches to 30 feet long, they had
heavy bony armor covering their head, neck, and sometimes thorax; they were the
first fish with pelvic fins, the first evolutionary step toward hindlimbs in the tetra-
pods, and some of the first fish to colonize freshwater environments were placo-
derms (see Phyllolepida); global distribution, middle Silurian to the end of the
Devonian.
Placodontia: an order of diapsid marine reptiles in the order Sauropterygia. The
placodonts ranged from 3 to 10 feet long and used their flattened teeth to prey pri-
marily on mollusks. The early type genus Placodus resembled modern marine igua-
nas in body structure, but the later genera developed turtle-like body armor as a
defense against the growing numbers of nothosaurs and plesiosaurs toward the end
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of the Triassic; placodonts are not, however, related to turtles. Europe, Asia, and
Africa; Early to Late Triassic.
Plagiaulacidae: a family of small basal multituberculate mammals in the suborder
Plagiaulacida; North America, Europe, and Asia; Late Jurassic to Early Cretaceous.
Plagiosauroidea: “flat lizard,” an enigmatic superfamily of early labyrinthodont
amphibians in the order Temnospondyli; about 3 feet long, they were characterized
by completely cylindrical vertebrae and a very short, wide skull (about 6 inches
long and 9 inches wide in Gerrothorax); South Africa, South America, and Asia;
Early to Late Triassic.
planktic: referring to organisms that do not swim actively, independent of the
water current; they may swim weakly but generally drift with the current.
Planocephalosaurus: an early genus of rhynchocephalian diapsid reptiles in the
family Sphenodontidae; only about 8 inches long, it was similar in most other
respects to the modern tuatara; North America and Europe; Late Triassic,
210–200 Ma.
plastron: the hard ventral covering of turtles.
Plateosaurus: “broad lizard,” a genus of basal prosauropod dinosaurs; from 15 to
34 feet long and weighing up to 9,000 pounds, it was among the largest of its time;
it appeared at the end of the Triassic and survived into the Early Jurassic; Central
and Northern Europe.
Platybelodon: “flat tusk,” the most common shovel-tusked gomphothere, a genus
of large proboscideans in the family Amebelodontidae; Platybelodon was similar to
Amebelodon, another shovel-tusker; Africa, Europe, Asia, and North America;
Miocene.
Platyceratidae: a family of cap-shaped gastropods in the order Euomphalina,
often with irregular growth. Snail-like, they are notable for a symbiotic relationship
since they are found only clinging to crinoids, usually on the calyx. Sometimes
found near the anus, they may have been coprophagous, but their place of attach-
ment is marked by a scar in some specimens and even a small hole in others, so that
they may have been parasitic. Global distribution, Ordovician to Jurassic, and pos-
sibly later.
platypus: see Ornithorhynchus.
Platyrrhini: an infraorder of anthropoid primates that comprises the New-World
monkeys. They are characterized by nostrils that face sideways and by prehensile
tails. The Platyrrhini diverged from the Catarrhini in the late Eocene or early
Oligocene, but they have remained primarily alike, an example of parallel evolution.
South America, Eocene to Recent.
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Platysomidae: a family of palaeonisciform fish in the suborder Platysomoidei;

deep-bodied and vertically flattened, with ganoid scales; global distribution,
Carboniferous to Triassic.
Plectronoceras: “spur horn,” a genus of primitive chambered cephalopods in the
subclass Nautiloidea and order Ellesmerocerida; the earliest-known nautiloid, it is
known from only a few specimens and was evidently short-lived; its tapering coni-
cal shell was 1 inch high, with several septa and a siphuncle; its body plan was not
stet well-adapted to swimming as the orthoconic shells of later nautiloids; North
America, late Cambrian.
Pleistocene: the next-to-last epoch of the Cenozoic Era, variously considered to
have begun between 2.6 and 1.8 Ma and lasting to 11,700 years before 2000 A.D.
Plesiadapiformes: a paraphyletic suborder of the oldest primates, appearing in the
Cretaceous, diversifying rapidly in the late Paleocene, and then disappearing at the
end of the Eocene; the suborder is generally considered to include the families
Paromomyidae, Plesiadapidae, Microsyopidae, and Campodeidae; North America,
Europe, Asia, and Africa.
Plesiadapis: the type genus of the family Plesiadapidae, it is significant in being
the only primate until Homo sapiens to inhabit both the Old and the New World;
North America and Europe, late Paleocene to early Eocene.
Plesiocidaroida: an order of regular echinoids in the subclass Euechinoidea,
poorly known but one of the groups of euechinoids diversifying in the Late Triassic
after almost all echinoids became extinct at the end of the Permian; Italy and North
Africa, Late Triassic, 240–220 Ma.
plesiomorphy: a character shared by an organism with primitive ancestors.
Plesiosauridae: the earlier of the two families of large marine reptiles in the order
Plesiosauria (see Elasmosauridae); global distribution, Triassic to Late Cretaceous.
Plesiosauria: “near lizard,” an order of mainly marine reptiles in the superorder
Sauropterygia; plesiosaurs grew to more than 40 feet long and had longer necks
than tails; they used their four strong flippers to become especially efficient swim-
mers; appearing near the end of the Triassic, the plesiosaurs are related to the notho-
saurs but directly descended from the pistosaurs; they flourished worldwide in the
Jurassic and persisted to the end of the Cretaceous.
Pleuracanthodii: see Xenacanthida.
pleural lobes: the lateral segments of a trilobite’s body, on both sides of the
thorax.
Pleurodictyum: a genus of dome-shaped colonial tabulate corals in the family
Favositidae; characterized by slender polygonal corallites; P. problematicum is usu-
ally found with the tubes of the worm Hicetes at the base of the colony, evidently
228 P
built on the tubes and possibly commensal with the worm; global distribution, late
Silurian to Carboniferous.
Pleurodira: “side neck,” the smaller of the two still-extant suborders of turtles in
the order Testudines; pleurodires are different anatomically from the cryptodires in
several ways because of the lateral flexure of the head, withdrawing it to the side, in
front of one of the forelimbs; Australia, Africa, and South America; Jurassic to
Recent.
Pleuromeiaceae: a family of small trees 3–4 feet high, lycophytes that may be
related to the Lepidodendrales and possibly ancestral to the modern Isoetaceae;
incertae sedis, however, because of a scarcity of fossil specimens; global distribu-
tion, Carboniferous to Early Triassic.
pleuronectiform: having both eyes on one side of head, characteristic of many flat
fish and bottom feeders.
Pleurotomaria: a large extinct genus of Paleozoic sea snails, gastropod mollusks
in the still-extant superfamily Pleurotomarioidea; related to the modern abalone but
characterized by a coniform or trochiform shell; global distribution, Early Jurassic
Pliobates cataloniae: a proposed species of hominoid in the family Pliobatidae,
described in 2015 as being close to the divergence between lesser and great apes;
dated at 11.6 Ma, this gibbon-like tree-dweller weighed about 10 pounds and has
several skeletal similarities to hominids; Spain, middle Miocene.
Pliocene: the fifth and last epoch of the Cenozoic period, from 5.3 to about 2.6 Ma.
Early hominins had appeared at the end of the Miocene, and many variants are
found in Pliocene strata throughout Africa and Asia. The period saw the beginning
of the decline of the large mammals of the Miocene, a decline which culminated in
the Quaternary extinctions. The Panama land bridge joined the Americas about 3
Ma, resulting in great changes in the animal populations of the two continents (see
Great American Biotic Interchange).
Pliohippus: a North American horse descended from Merychippus and ancestral
to Equus; middle to late Pliocene.
Pliohyracidae: the largest family of early hyraxes in the order Hyracoidea; partly
arboreal but several genera cow-sized; Africa, Eocene to Pliocene.
Pliomera: a well-known genus of phacopid trilobites in the family Pliomeridae. It
is distinguished by a pentagonal glabella; small, raised eyes; up to two dozen tho-
racic segments; and a small pygidium. Argentina, Norway, Iran, Russia, and China;
Early Ordovician, 479–472 Ma.
Pliomeridae: a family of Ordovician phacopid trilobites in the suborder Cheirurina;
several subfamilies are monophyletic but others are uncertain; global distribution,
Early to Late Ordovician.
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Pliopithecus: a genus of early hominoid primates in the superfamily Pliopithecoidea;

Pliopithecus appeared in Asia in the early Miocene and spread into Europe before
disappearing about 10 Ma; although gibbon-like, it is not considered a direct ances-
tor of the gibbons; see Propliopithecus.
Pliosauroidea: a suborder of marine sauropterygian reptiles in the order
Plesiosauria, characterized by a relatively short neck and a massive skull; a 33-foot-
long specimen found in the early twenty-first century is the first entire intact skele-
ton of a pliosaur; it was recovered at Spitsbergen, 800 miles from the North Pole;
Europe, Africa, and Australia; Early Jurassic to Late Cretaceous.
Plotosaurus: “swimmer lizard,” a genus of mosasaurs in the order Squamata;
strong swimmers, it was similar to ichthyosaurs but more closely related to turtles;
known only from a complete skull and skeletal fragments found in Central
California, it is estimated to have grown to as much as 40 feet long; synonym:
Kolposaurus; United States, Late Cretaceous, 70–66 Ma.
Pluricarpellatia: a genus of flowering aquatic plants in the still-extant order
Nymphaeales; found in the Crato Formation of Brazil and dated about 112 Ma, it is
a basal angiosperm; with flowers, seeds, large leaves, and branching rhizomes, it
resembles modern water lilies; Brazil, Early Cretaceous.
Poaceae: the large family of grasses, in the order Poales; see grasses; Late
Cretaceous to Recent.
Podokesaurus: “swift-footed lizard,” a genus of very small saurischian dinosaurs
in the superfamily Coelophysoidea; characterized by tiny forelimbs and long, deli-
cate tail and neck, Podokesaurus was about 3 feet long; it is known only from two
partial specimens, and identification as a genus separate from Coelophysis is debat-
able, but it is among the earliest dinosaurs found in Eastern North America; Early
Jurassic.
Poebrotherium: “grass-eater,” a genus of even-toed ungulates in the family
Camelidae; at 3 feet high, Poebrotherium was about the size of a goat, larger than its
ancestor Protylopus; North America, late Eocene to early Oligocene.
Pogonodon: a genus of carnivorous feliform mammals in the family Nimravidae;
weighing 50–80 pounds, with 2-inch canines, Pogonodon probably hunted from
ambush; it was widespread in North America in the Oligocene and may have sur-
vived in Eurasia into the Miocene.
Pojetaia: a genus of small, primitive bivalve mollusks in the order Fordillida and
family Fordillidae; suboval, less than one-tenth of an inch long; global distribution,
early middle Cambrian.
Polacanthus: “many spines,” a contentious genus of ankylosaurian dinosaurs ten-
tatively placed in the family Nodosauridae; poorly known from body fragments, it
was quadrupedal, well-armored with dorsal spines, and estimated to be about 16
230 P
feet long; two nomina dubia associated with it are Hylaeosaurus and
Horshamosaurus; England, Early Cretaceous, 130–125.
polychaetes: see annelids and scolecodonts.
Polydolopidae: a family of small South American marsupials in the order
Paucituberculata, similar to the modern shrew opossums; South America and
Antarctica, middle Paleocene to middle Eocene or possibly Oligocene (see also
Antarctodolops).
Polygnathidae: a family of conodonts in the suborder Ozarkodinina; the family
includes several genera that are important as index fossils; global distribution,
Silurian to Late Triassic.
Polygnathus costatus partitus: a species of ozarkodinid conodonts in the family
Polygnathidae; the index fossil whose first appearance marks the beginning of the
Middle Devonian at 393.3 Ma; global distribution.
Polyodon spathula: the American paddlefish, a still-extant species of cartilaginous
acipenseriform fish in the family Polyodontidae; averaging about 5 feet long, it is
characterized especially by its paddle-shaped, elongated rostrum; the species is
known from the Late Cretaceous and the middle Eocene Green River formation;
North America, Cretaceous to Recent.
polyphyletic group: a group of organisms that do not derive from a single com-
mon ancestor in the group, so that the name of the group is used only informally. A
monophyletic group is a clade consisting of an ancestor and all of its descendants,
as opposed to a paraphyletic group, which consists of a common ancestor and some
but not all of its descendants. Paraphyletic groupings are still useful; for instance,
dinosaurs and birds are paraphyletic but obviously greatly different from each other.
polyphyodonty: see dentition.
Polyplacophora: the chitons, a still-extant class of marine mollusks known from
the Cambrian but with a sparse fossil record; synonym: Amphineura. They are char-
acterized by an oval, flattened body covered with a dorsal shell of 7 or 8 calcareous,
slightly overlapping plates. See also Wiwaxia. Global distribution, Cambrian to
Recent.
polyprotodont: an informal term used for a diverse grouping of primitive marsupi-
als that arose in the Cretaceous and was succeeded by the diprotodonts.
Pondaungia: a species of small, primitive haplorrhine primates in the family
Amphipithecidae; synonym: Amphipithecus; probably frugivorous, similar in some
ways to the adapiform primates of North America; Burma, early Eocene.
Porifera: the phylum of sponges. Whether the phylum is monophyletic is in dis-
pute, but it is generally considered to date from the late Precambrian, with definitive
specimens appearing in the early Cambrian. It contains several classes:
Hexactinellida, Calcarea, Sclerospongiae (polyphyletic), and Demospongiae. The
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Archaeocyatha, Heteractinida, Stromatoporoidea, and Receptaculitidae are also

considered sponges in some classifications.
Porocharaceae: a family of charophytic algae in the order Charales, probably
ancestral to the post-Paleozoic family Raskyellaceae and others; cosmopolitan,
Carboniferous to Cretaceous.
Poroxylaceae: a monogeneric family of pteridospermous plants formerly assigned
to the Cordaitales but now considered progymnosperms or seed ferns; global distri-
bution, Carboniferous.
Posidonia Shale: see Holzmaden.
Postosuchus: “crocodile from Post,” a genus of large archosauromorph reptiles in
the family Rauisuchidae. Most specimens come from the Post Quarry in West Texas,
but others have been found across the Southern United States. Up to 15 feet long
and 4 feet high, Postosuchus is variously estimated to have weighed between 600
and 1500 pounds and was one of the dominant predators of the Triassic in
Southwestern North America. Its forelimbs were somewhat shorter than the
hindlimbs but were robust enough to enable it to walk either bipedally or quadrupe-
dally. North America, Late Triassic, 222–201 Ma.
Precambrian Eon: the long period of geologic time preceding the Paleozoic Era,
up to 541 Ma. There are signs of organic life as far back as 4 billion years, and fos-
sils of bacteria as old as 3.5 billion years. The first structurally complex eukaryotes
appeared about 1.5 billion years ago, but the first multicellular animals came much
later, around 700 Ma. The last part of the Precambrian, the Ediacaran Period (630–
541 Ma), is marked by rapidly diversifying marine animals.
preservational environment: a depositional environment that fosters the preser-
vation of fossils, especially in relation to the degree of acidity in the soil or marine
strata, since high acidity tends to dissolve bone and other organic remains.
Priapulida: a still-extant phylum of burrowing marine worms; characterized by a
tubular shape, spines around the mouth, and an extensible, spiny feeding proboscis;
some species are tolerant of anoxia and hydrogen sulfide; stem-group priapulids are
known from Cambrian strata, with the earliest crown-group specimens appearing
only in the Late Pennsylvanian; see also Ottoia; global distribution, Late
Pricyclopyge: a genus of small deep-ocean trilobites in the order Asaphida and
family Cyclopygidae; characterized by a broad glabella and very large eyes, it may
have swum upside down; Europe and Asia, Early to Late Ordovician.
Primelephas: a genus of primitive elephants with four tusks in the subfamily
Elephantinae; one of the first elephants, probably ancestral to the Mammutioidea
and to modern elephants; similar to the gomphotheres; Africa, Miocene to Pliocene.
Primitiopsis: a genus of dimorphic ostracods in the order Paleocopida; it probably
lived in the deep subtidal zone; cosmopolitan, Late Devonian and Silurian.
232 P
primitive features: characteristics of an organism that are relatively close to those

of its ancestors, as opposed to advanced features.
Prioniodontida: a clade of conodonts with composite teeth, including the subor-
ders Ozarkodinina, Prioniodinina, and others; global distribution, Early Ordovician
to Triassic.
Prionocyclus: a genus of ammonoids in the order Ammonitida and family
Collignoniceratidae; ranging in size between 2 and 4 inches, several species are
index fossils for the Turonian Stage of the Late Cretaceous; global distribution,
94–90 Ma.
Prionosuchus: a genus of very large temnospondyls in the family Archegosauridae;
the largest amphibian at an estimated 30 feet long, it is known from two partial
specimens; Brazil, middle Permian.
Priscacara: a genus of freshwater actinopterygian fish in the still-extant family
Cichlidae; of medium size, it is characterized by a flattened oval body and spiny
dorsal, anal, and ventral fins; North America, early to late Eocene.
Priscansermarinus: “ancient sea goose,” a monospecific genus of crustacean
arthropods similar to the modern gooseneck barnacle and classified in its order,
Pedunculata; P. barnetti is known only from the Burgess Shale and is considered the
oldest known barnacle; Canada, middle Cambrian, 508 Ma.
Pristerodon: a genus of dicynodont therapsids, type species P. mackayi; about 2
feet long but only a few inches high, the herbivorous Pristerodon seems to be one of
the first animals able to hear airborne sounds, as opposed to vibrations in the ground;
Southern Africa and India, late Permian.
Proailurus: a genus of cat-like carnivores, close to the ancestry of the Felidae,
probably ancestral to Pseudaelurus; about the size of a lynx but with slender limbs
and a long tail; partially arboreal; Europe and Asia, late Oligocene to Miocene.
Proanura: an order of amphibians ancestral to the modern order Anura, the frogs;
a very sparse fossil record leaves their phylogeny questionable; the only recognized
Mesozoic proanuran is Triadobatrachus, a Triassic amphibian from Madagascar;
authorities disagree as to whether the Temnospondyli, the Lepospondyli, or some
other Paleozoic group may be the ancestors of the Anura, which appeared in the
Jurassic.
Probactrosaurus: a genus of Cenomanian ornithischian dinosaurs, one of the
smallest and earliest hadrosaurs; about 18 feet long and weighing up to a ton, it was
principally quadrupedal, with long and slender forelimbs; similar in several ways to
Iguanodon, the ancestor of the hadrosaurs, but with a thumb spike and weaker fore-
limbs; China, Late Cretaceous, 96–88 Ma.
Probainognathus: “advanced jaw,” a monospecific genus of small cynodont the-
rapsids; carnivorous and the size of a very small dog, P. jenseni is one of the earliest
therapsids to have a squamosal-dentary jaw joint similar to the typical mammalian
P 233
structure; it may be related to the Chiniquodontidae; Argentina and Brazil, Late

Probathyopsis: a basal uintathere, a small genus of early dinoceratan ungulates,
closely related to the Asian Prodinoceras; North America, late Paleocene to early
Eocene.
Probelesodon: a genus of early cynodonts in the family Chiniquodontidae; dog-
sized, one of the larger chiniquodontids; South America, Middle to Late Triassic.
Proboscidea: an order of mammals that originated in Africa in the early Eocene
and had achieved nearly global distribution by the middle Miocene. The earliest
proboscideans, such as Moeritherium, were trunkless, but Oligocene species such
as Paleomastodon had a rudimentary trunk and short tusks. The order includes the
two surviving species of elephants as well as the extinct mammoths, gomphotheres,
deinotheres, and possibly the desmostylians.
Procaviidae: the modern family of hyraxes, known principally from Africa, where
some of their ancestors in the Pliohyracidae had evolved into sirenians and ele-
phants; Africa, Europe, and Asia; Miocene to Recent.
Proceratops: see Ceratopsia.
Procolophonia: a suborder of small herbivorous reptiles related to the Pareiasauria;
formerly included in the paraphyletic order Captorhinida; the colophonids were
about a foot long with large eyes and were probably fossorial; they survived the
end-Permian extinction and diversified in the Triassic; South Africa, Antarctica, and
Brazil; late Permian to Late Triassic.
Proconsul: a genus of arboreal hominoid primates in the family Proconsulidae;
about the size of a modern baboon, Proconsul was sexually dimorphic and bore
characteristics of both monkeys and apes; the four named species differ mainly in
body size, the specimens ranging from 10 to 150 pounds; possibly ancestral to
Propliopithecus and later primates, including humans; Kenya and Uganda, early
Miocene, 23–25 Ma.
Procoptodon: a genus of short-faced kangaroos in the family Macropodidae and
extinct subfamily Sthenurinae; the genus includes the smallest of the Sthenurinae,
P. gilli, while P. goliah was about the same size as Sthenurus, weighing more than
500 pounds; they probably moved more slowly than modern kangaroos; Australia,
Pliocene to late Pleistocene.
Procynosuchus: “before dog-crocodile,” a genus of basal cynodont therapsids in
the family Procynosuchidae; the size of a cat, it was quadrupedal and was probably
semi-aquatic; it was notable for a very primitive dentition and a long tail; Europe
and Africa, Late Permian.
Prodinoceras: a genus of basal uintathere ungulates in the order Dinocerata; they
were about 10 feet long and very similar to Probathyopsis of North America;
Mongolia, late Paleocene.
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Prodryas persephone: a species of lepidopterans in the family Nymphalidae, a

butterfly known from a single, unusually well-preserved specimen found in the
Florissant Fossil Beds; about an inch wide, the specimen shows the wing venation
and even the original color patterns; United States, late Eocene.
Productida: a large order of late Paleozoic articulate brachiopods; a very diverse
order containing a wide range of shapes, generally characterized by concave-convex
shells and by spines on the surface; global distribution, Ordovician to Triassic.
Proetida: an order of generally small trilobites, very similar to the Ptychopariida;
characterized by a narrow rostral plate and a high, sometimes pitted, glabella, most
were less than an inch long; the only order of trilobites to survive the Devonian
extinction, the Proetida diversified and persisted through the Carboniferous and
Permian, but they disappeared from the fossil record shortly before the end-Permian
extinction; global distribution, Early Ordovician to Permian.
Proganochelys: a genus of stem-turtles in the order Testudines; dated at 210 Ma, it
is the oldest turtle found with a complete shell, carapace and plastron; it was differ-
ent in several ways from modern turtles; about 3 feet long, it had a spiked tail ending
in a club and was probably mainly terrestrial; Europe and Asia, Late Triassic.
Proganosauria: see Mesosauria.
Prognathodon: a genus of squamatan reptiles in the family Mosasauridae. Several
species of these large mosasaurs, ranging from 20 to more than 40 feet long, were
widespread at the end of the Cretaceous, preying on fish, ammonoids, other shell-
fish, and turtles. Skulls and teeth from phosphate deposits near Khouribga, Morocco,
are well-known as commercial specimens. Two almost-complete specimens from
Canada show fins and stomach contents, including a 5-foot-long fish. Prognathodon
is similar to Globidens in its shape and in a robust skull, and its teeth also are spe-
cialized for crushing shellfish, although sharper-pointed than the blunt teeth of
Globidens. North America, Eurasia, Africa, and New Zealand; 71–66 Ma.
progymnosperms: woody, spore-bearing plants intermediate between ferns and
gymnosperms; global distribution, Middle Devonian to Mississippian.
Prokennalestes: a poorly known genus of basal eutherian mammals, considered a
possible ancestor of Kennalestes; known only from an individual ear bone; China,
Prolacertiformes: see Protorosauria.
Prolecanitida: one of the longer-lived and most speciose orders of ammonoids,
with more than 1200 species; discoidal or thinly lenticular, they are notable for the
increasing complexity of their suture from the Devonian to the Permian; global
distribution, Late Devonian to Late Triassic.
Promicroceras: a small genus of ammonoids in the order Ammonitida and family
Eoderoceratidae; characterized by strong ribbing and small spines; one of the
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c ommon fossils of the Sinemurian stratum on the Dorset coast, between 199 and
192 Ma; England, Early Jurassic.
Promissum pulchrum: a species of conodonts in the suborder Prioniodinina; one
of the few known fossil imprints of the animal and the largest known at 16 inches
long; Soom Shale, South Africa; Late Ordovician.
pronghorn: see Antilocapridae.
Pronucleolites: see Nucleolitidae.
Propalaeotherium: a genus of very small perissodactyl mammals at first classified
in Palaeotheriidae but in 2004 placed in Equidae; it was found in the Messel Shale
Pit in 1849 and later across Europe and in China; a forest browser on leaves and
berries (found with its stomach contents), it stood 1–2 feet high at the shoulder and
weighed only about 22 pounds; hoofless, it had nail-like hooflets at the ends of the
toes, four on the manus and three on the pedes; Europe and Asia, Eocene.
Proplebeia dominicana: a species of stingless bee found in amber deposits in the
Dominican Republic; its successfully reconstituted and copied DNA, 40 million
years old, is the oldest yet recovered for study; Caribbean, late Eocene.
Propliopithecus: a genus of early anthropoid primates in the family Pliopithecidae,
possibly ancestral to the early hominins; only 1.5 feet long and with stereoscopic
vision, it was more advanced than the earliest primates; it was probably ancestral to
Pliopithecus, and was closely related to the early Oligocene Aegyptopithecus; North
Africa, middle to late Oligocene.
Prorastomidae: a family of primitive sirenians, possibly ancestral to the modern
dugongs and manatees; known only from two Jamaican specimens, the partly ter-
restrial Pezosiren of the early Eocene and the fully aquatic Prorastomus of the mid-
dle Eocene.
Prorichthofenia: an unusual genus of reef-forming strophomenid brachiopods in
the order Productida and family Richthofeniidae; characterized by their conical,
horn-shaped pedicle valve, with the upper valve acting as a lid covering the cone;
North America and Europe, middle to late Permian.
Prorubidgea: a synonym of Aelurognathus.
Prosaurolophus: a genus of duck-billed dinosaurs in the subfamily Saurolophinae;
medium-sized hadrosaurids, they were about 30 feet long and had a prominent crest
above the eyes; North America, Late Cretaceous, 76–73 Ma.
prosauropods: the first sauropod dinosaurs, which were among the first dinosaurs
to evolve from the theropods at the end of the Middle Triassic. Prosauropods devel-
oped and spread rapidly, the large Plateosaurus and others surviving the Triassic
extinction, followed by Anchisaurus in the Early Jurassic and then by the sauropods
themselves, which eventually achieved global distribution and some of which were
236 P
among the largest of all dinosaurs. See also Nyasasaurus and Herrerasaurus.
Triassic to Cretaceous.
Prosimii: see Strepsirrhini.
Protarchaeopteryx: a species of coelurian theropod dinosaurs in the infraorder
Oviraptorosauria; probably flightless although it had long, bird-like legs and true
feathers on body, wings, and tail; more than 3 feet long, it was larger and more
primitive than Archaeopteryx but lived 10 million years later; one of several feath-
ered dinosaurs in the Jehol biota; China, Early Cretaceous, 125 Ma.
protaspis: the hatching ontogenetic phase of trilobites; see also holaspis and
meraspis.
Proteroctopus: a monospecific genus of cephalopods tentatively placed in the fam-
ily Palaeoctopodidae; if P. ribeti is an octopus, it is the oldest known; known only
from one specimen found at La Voulte-sur-Rhône; France, Middle Jurassic.
Proterosuchidae: a family of reptiles, probably paraphyletic but considered basal
archosaurs. When most large therapsids became extinct at the end of the Permian,
the proterosuchids became the largest carnivores and flourished at the beginning of
the Triassic; they were replaced by phytosaurs and crocodilians in the Middle
Triassic. Global distribution except North America, late Permian to Middle Triassic.
Proterosuchus: one of the earliest archosaurs, in overall appearance similar to
modern crocodiles except for its distinctive hooked premaxilla; up to 7 feet long
with long jaws, it was one of the largest land reptiles of its time; first found in the
Karoo Basin of South Africa and now known also from China; Early Triassic,
252–250 Ma.
Proterotheriidae: a diverse family of horse-like South American ungulates in the
order Litopterna; the proterotheres arose in the Paleocene and flourished in the
Eocene before declining in the Miocene; most genera became extinct in the Pliocene
at about the time that true horses arrived in South America; one genus, however,
survived in Argentina and Uruguay into the Pleistocene.
Protichnites: see Diplichnites.
protists: an informal group of unrelated simple, eukaryotic organisms that are not
animals, plants, or fungi; they are microscopic and chiefly aquatic.
Protoavis: a contentious specimen with a bird-like skull and other bird-like fea-
tures. Dated at 225 Ma, it has been proposed as an early avian evolutionary branch
from the time of the earliest dinosaurs, but since it is known from only an assort-
ment of fragments, its reconstruction is questionable. The tentative reconstruction
indicates an animal 1–2 feet long. Although it is unclear whether it had a true wish-
bone, Protoavis seems unlikely to be ancestral to modern birds. Post Quarry, Texas,
Middle to Late Triassic.
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Protoceratidae: a long-lived family of herbivorous artiodactyl ungulates; the

males were notable for a pair of rostral horns near the nose as well as a pair of horns
at the back of the head; they had relatively long limbs and resembled deer and elk,
growing up to 6 feet long; North America, middle Eocene to early Pliocene.
Protoceratopsidae: a family of ornithischian dinosaurs in the suborder Ceratopsia;
herbivorous, with the characteristic ceratopsian neck frill and large skull, but only
up to 9 feet long; known only from Asia, Late Cretaceous, 100–70 Ma.
Protocetus: a genus of archaeocete cetaceans in the family Protocetidae, one of the
earliest oceangoing whales, fully aquatic and bearing only vestigial limbs that prob-
ably could not have supported its weight on land. Like the deep-ocean Rodhocetus,
Protocetus had unfused sacral vertebrae, so that it could use its tail efficiently for
propulsion in the water. In addition, it had the advanced character of vestigial
hindlimbs unattached to the backbone. Egypt, middle Eocene.
protoconodonts: an obscure group of small animals that may be either the earliest
conodonts or a stem group of Chaetognatha; they seem at least quite different from
the euconodonts; cosmopolitan, late Precambrian to Early Ordovician.
Protodonata: see Meganisoptera.
Protoeumalacostraca: an obscure group of early crustaceans that are thought to
be the stem group of the subclass Eumalacostraca, which includes most of the living
crustaceans; the eumalacostracans appeared in the Devonian, but their predecessors
remain in doubt.
Protopteridales: an order of fern-like plants in the class Polypodiatae; eusporan-
giate, producing sporangia at the edges of their leaves, and herbaceous or woody;
now considered to be related to the Rhyniales; cosmopolitan, Devonian.
Protorosauria: an order of archosauromorph reptiles that flourished in the Triassic;
synonym: Prolacertiformes. They are generally characterized by very long necks
and short bodies (see Tanystropheus). Some families, such as the Tanystropheidae,
are 15–20 feet long, while the Drepanosauridae are only 1–2 feet long. The
Protorosauridae of the late Permian, which were about 7 feet long, are the earliest-
known archosauromorphs. South America, Europe, and Asia; late Permian to Late
Triassic.
Protosalvinia: a genus of early land plants; similar to some phaeophytes (brown
algae) but generally considered one of the earliest bryophytes, probably a liverwort;
known from several Late Devonian specimens found in North and South America.
Protosirenidae: a monogeneric family of aquatic mammals in the order Sirenia;
like their descendants the manatees, the protosirenians inhabited temperate to tropi-
cal waters; Protosiren had hindlimbs, which have disappeared in the modern spe-
cies; the type species of the genus is Protosiren fraasi; North America, Europe, and
Asia; Eocene.
238 P
Protosuchia: a group of small to medium-sized archosaurian reptiles in the super-

order Crocodylomorpha; about 3 feet long, they were slender and terrestrial; global
distribution, Late Triassic to Early Cretaceous.
Protosuchus: a genus of basal crocodylomorphs that appeared near the boundary
of the Triassic and Jurassic; primarily terrestrial, Protosuchus was 3 feet long and
moderately armored, and its hindlimbs were somewhat longer than its forelimbs;
North America and Africa, Early Jurassic.
Prototaxites: an enigmatic genus of large, mid-Paleozoic tree-like structures; esti-
mated to be as much as 3 feet in diameter and 26 feet high, they were named “early
yew” because of the long, intertwined tubes that formed the trunk, resembling the
proto-vascular structure of other early plants such as the Calamitales or Nematophyta;
now considered probably fungal or algal; North America, Europe, and Saudi Arabia;
Late Silurian to Late Devonian, 420–370 Ma.
Prototheria: an obsolescent term for a subclass of mammals that includes the
order Monotremata; Late Triassic to Recent.
Protungulatum: a genus of early ungulate mammals, considered a non-placental
eutherian; rat-sized, Protungulatum had a distinctive foot structure and crushing
molars designed for eating plants; probably related to the arctocyonids; North
America, Late Cretaceous to Paleocene.
Protura: “primitive tail,” an enigmatic still-extant order of very small soil-dwelling
insect-like animals tentatively placed in the phylum Hexapoda; less than 2 mm long
and eyeless, they have no antennae or pigmentation; they are characterized by six
legs, a cylindrical body, and a conical head; their fossil record is very sparse, but
they are thought to be primitive hexapods, going back perhaps to the Early Devonian.
Protylopus: a genus of even-toed ungulates in the family Camelidae; one of the
earliest camelids, it was about the size of a rabbit; North America, middle Eocene.
proximal: referring to a body part of a bilaterian animal that is closer to the center
of the body than a distal part; in plants, referring to an area near the point of
attachment.
Psammodontidae: a family of holocephalic fish; characterized by bradyodont
dentition consisting of rectangular toothplates in two rows meeting at the midline;
North America and Europe, late Devonian to late Permian.
Psammosteidae: a family of heterostracan fish of the Devonian; with a round and
flattened body, they ranged in size from the early genus Drepanaspis (about 1 foot
long) to some at the end of the Devonian that were 6 feet long; they were the only
heterostracans that survived the Frasnian extinction, but were finally extinguished in
the Hangenberg Event at the end of the Famennian Stage; global distribution, Early
to Late Devonian.
Psarolepis: an enigmatic early genus of lobe-finned fish; it exhibits a combination
of osteichthyan and non-osteichthyan gnathostome characters; perhaps a basal oste-
P 239
ichthyan, close to the common ancestry of sarcopterygians and actinopterygians;

China, dated as early as 418 Ma, at the beginning of the Devonian.
Psaronius: a genus of tree ferns in the still-extant order Marattiales; the name orig-
inally referred to only the stem. Growing more than 30 feet high, Psaronius has
trunk specimens as large as 3 feet in diameter at the base. It is notable for the
structure of its trunk, which consisted of many thin roots intertwined with stems,
somewhat similar to the manner of Tempskya. The leaves grew in a crown at the top
of the trunk and are known mainly as the form genus Pecopteris, with the leaves of
some species classified as the form genus Sphenopteris. Global distribution, Late
Carboniferous to late Permian.
Pseudaelurus: a genus of Miocene mammals in the family Felidae, the first felid
to appear in North America; early species were the size of a domestic cat, while later
ones reached the size of a cougar; North America, Europe, and Asia, early to late
Miocene.
Pseudoborniales: an early order of sphenopsid plants in the class Equisetopsida,
growing to 60–75 feet; cosmopolitan, known only from the Late Devonian.
Pseudocidaris: a genus of sea urchins, Mesozoic euechinoids in the family
Hemicidaridae; P. mammosa is famous for its bulbous, club-shaped spines; Europe,
Africa, and Asia; Middle Jurassic to Early Cretaceous.
Pseudosuchia: a clade now considered to include living crocodilians as well as the
reptiles that arose in the Early Triassic and dominated the terrestrial carnivorous
niches of the Late Triassic, including the rauisuchians; all except the Sphenosuchia
and Crocodyliformes were extinguished in the Triassic extinction; cosmopolitan,
Triassic to Recent.
Psigraptus jacksoni: a species of anisograptid graptolites, notable for specimens
of three-dimensional pyritized remains of zooids; the specimens show, however, no
tentacles or other morphological details; Australia and Korea, Early Ordovician.
Psiloceras spela group: a group of smooth ammonitid ammonoids in the family
Psiloceratidae; its first appearance marks the beginning of the Early Jurassic at
201.3 Ma; Austria.
Psiloceratidae: a family of ammonoids in the order Ammonitida, probably
descended from the Ussuritidae; psiloceratids generally have a smooth surface or
blunt ribbing; cosmopolitan, Early Jurassic.
Psilophyton: a genus of fern-like tracheophytic plants in the class Trimerophytopsida
and family Psilophytaceae; among the first vascular plants, they grew from under-
ground rhizomes, with naked branches and a spore-producing structure at the tips of
branches; P. forbesii is the largest known at about 2 feet, but most were 1 foot high;
North America, Europe, and Asia; Late Silurian to Late Carboniferous.
Psittacosaurus: a speciose genus of Asian ceratopsian ornithischian dinosaurs in
the family Psittacosauridae. It includes more species than any other dinosaur genus
240 P
and is also one of the most completely known dinosaur genera because known from
several complete skeletons and hundreds of individual specimens. It is the most
common dinosaur in the Jehol biota. Psittacosaurus was bipedal, generally 6–7 feet
long, and characterized by a large, parrot-like beak. Early in the Cretaceous, prob-
ably around 130 Ma, the psittacosaurids split off from the neoceratopsian branch
which led to the familiar horned and frilled ceratopsians of the Late Cretaceous. See
also Liaoceratops. China, Mongolia, and Siberia; Early Cretaceous, 123–100 Ma.
Pteranodon: “toothless wing,” a genus of large pterosaurs in the suborder
Pterodactyloidea; they were toothless and had a large beak and a large cranial crest,
and the largest species had a wingspan up to 23 feet; they are especially well-known,
with more well-preserved specimens known than any other pterosaur; North
America and Europe, Late Cretaceous, 86–84 Ma.
Pteraspis: “wing shield,” a genus of small, jawless fishlike vertebrates in the sub-
class Heterostraci and order Pteraspidiformes. They were finless except for small
wing-like protrusions at the sides of the shield over the gills, which may have aided
in swimming. They are also characterized by a striking medial dorsal spine extend-
ing backward from the wide shield that protected the forward half of the body; the
rest of the 7-inch-long body was covered by small scales. North America and
Europe, Early Devonian.
Pterichthyodes: a genus of freshwater antiarch placocoderms in the family
Asterolepidae; distinguished by jointed wing-like appendages extending outward
on both sides of the head; it had a heterocercal tail fin and eyes on top of the head-
shield; up to a foot long, it had heavy anterior armor and scales on the posterior,
differing from its cousin Bothriolepis, which was posteriorly naked, with no scales;
it may have used its pectoral appendages to pull itself along the bottom; known only
from Scotland, Middle Devonian.
Pteridospermatophyta: the seed ferns, a large extinct group of gymnosperms
similar to tree ferns except that they bore seeds and pollen-bearing structures on the
fronds. Currently four orders are recognized: Callistophytales, Lyginopteridales,
Medullosales, and Peltaspermales. Some similar plants survived to the Eocene, but
the relationship is unclear. The seed ferns have also been called Cycadofilicales and
sometimes Pteridospermae. Global distribution, Late Devonian to Cretaceous, but
an especially important part of Carboniferous floras.
Pterobranchia: a still-extant class of small, worm-like colonial animals in the
subphylum Hemichordata. Phylogenetic analysis indicates that they are related to
the graptolites, but they differ in morphology and habits. The benthic pterobranchs
usually secrete tube-stalks which they extend to filter-feed. Families: Atubaridae,
Cephalodiscidae, and Rhabdopleuridae. Global distribution, early Cambrian to
Recent.
Pterodactyloidea: a large suborder of flying archosaurs in the order Pterosauria;
generally tail-less and much larger than the basal pterosaurs of the Late Triassic;
several later genera were toothless; many had large crests, and their wingspan
P 241
ranges from a few inches to as much as 40 feet (see Quetzalcoatlus); Europe and
China, Middle Jurassic to Late Cretaceous.
Pterodactylus: “finger wing,” the type genus of flying pterosaurs of the suborder
Pterodactyloidea; Europe and Africa, Jurassic and Cretaceous.
Pterosauria: an order of archosaurs distinguished by the leathery membranes
stretched between extremely elongated fingers that enabled them to glide or soar.
They are traditionally divided into two suborders, the rhamphorhynchoids and the
Pterodactyloidea, but only the latter is monophyletic. The wingspan of the ptero-
saurs ranged from a few inches to as much as 45 feet. Pterosaurs were carnivorous
and especially piscivorous, and many families were toothless.
They were not dinosaurs and not birds, but evolved along with dinosaurs from a
common ancestor in the Late Triassic. Because their bones were bird-like — thin-
walled and slender — the fossil record is less complete for some pterosaurs than for
most archosaurs. Their eggs are extremely rare, but several discoveries since 2004
have shown that the parents may have buried eggs; whether they were altricial or not
is also unclear. The exact shape of their “wings” and their mode of flying is still
unclear. Some investigators believe they had unusually large flocculi, brain lobes
that link brain activity with body movement, and that their flocculi may have enabled
them to fly more efficiently than modern birds and bats.
About 100 species are known, the earliest being dated at the Carnian-Norian
boundary in the Late Triassic, about 230 Ma. They flourished worldwide during the
Jurassic but were dying out by the late Cretaceous.
Pterygotidae: a family of marine arthropods in the extinct order Eurypterida.
Large sea scorpions, they had 2 pairs of eyes and a short, flattened telson ending in
a short spine. They swam with their tail and two paddle-like appendages, for which
they were named. The largest known is Jaekelopterus, estimated at 8 feet, and a
2009 discovery, Ciurcopterus, may be larger. Global distribution, Silurian to Late
Devonian.
Pterygota: the winged insects, a subclass of the class Insecta; the pterygote insects
are unrelated to the Pterygotidae, sea scorpions, except as arthropods; traditionally
divided into the infraclass Neoptera and the paraphyletic group Palaeoptera but now
reclassified in different ways which are still unsettled.
pterygote: referring to winged or secondarily apterous insects.
pterygotid: referring to the marine arthropods in the family Pterygotidae.
Pterygotus: a genus of large marine arthropods in the family Pterygotidae; its spec-
imens are 5–6 feet long; North and South America, Europe, and Australia; Silurian
to Devonian.
Ptilodontoidea: a group of multituberculate mammals in the suborder Cimolodonta;
generally small and rodent-like, poorly known but probably derived from the
Plagiaulacida; North America and Europe, Late Cretaceous to Eocene.
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Ptilodus: a North American multituberculate in the family Ptilodontidae; squirrel-

like and 1–2 feet long, it was widespread across North America, early to late
Paleocene.
Ptychagnostus: a genus of very small, blind agnostid trilobites in the family
Ptychagnostidae; less than half an inch long, it is characterized by 2 segments in the
glabella and 3 in the pygidium; it was nektobenthic, generally living in the water
column; global distribution, middle Cambrian.
Ptychodus: “layer tooth,” a genus of early sharks in the extinct superfamily
Hybodontoidea and family Ptychodontidae; about 30 feet long, Ptychodus was
characterized by its unusual dentition, which consisted of thousands of flat molar
teeth, evidently used to crush bivalves and crustaceans; cosmopolitan, Cretaceous to
Paleocene.
Ptychopariida: a large early order of trilobites that appeared, along with the
Redlichiida and Corynexochida, early in the Cambrian; diverse but generally char-
acterized by an oval shape and small pygidium; global distribution, early Cambrian
to Late Ordovician.
Ptyctodontidae: “folded teeth,” a family of placoderm fish in the monotypic order
Ptyctodontida, related to the Arthrodira and Phyllolepida; weakly armored on head
and shoulders and the only sexually dimorphic placoderms, the males having hook-
like claspers on their pelvic fins; they had large eyes, heads, and bodies; now con-
sidered the most derived placoderms, they may be ancestral to the jawed vertebrates;
cosmopolitan, Middle to Late Devonian.
Puertasaurus: a genus of titanosaurian sauropodomorph dinosaurs closely related
to Argentinosaurus; known only from skeletal fragments of one individual, it is
estimated to have been massive and 100–130 feet long; Argentina, Late Cretaceous,
Cenomanian or Maastrichtian.
Pugnax: a genus of articulate rhynchonellid brachiopods in the family Pugnacidae;
about 1 inch wide, generally characterized by a sub-triangular shape and strong
central ribs, often with a smooth shell; global distribution, Devonian to Permian.
Puijila: a genus of ancestral pinnipeds, the most primitive relative of the Phocidae,
the earless seals; otter-like and semi-aquatic, it had a tail and four legs with enlarged
feet; see also Enaliarctos; Canada, late Oligocene to Miocene.
Pulmonoscorpius kirktonensis: a species of giant terrestrial scorpions more than
2 feet long; Scotland, Carboniferous, 320 Ma.
punctate: pierced by pores; in brachiopods, a condition in which elongated, regu-
larly arranged cavities extend from the inside of the shell to the outside; although all
terebratulids are punctate and all atrypids and pentamerids are impunctate, the con-
dition cuts across the other systematic boundaries.
P 243
punctuated equilibrium: the evolutionary model that sees most change as occur-
ring in brief bursts of evolutionary diversification, with longer periods of stasis in
between; see also phyletic gradualism, anagenesis, and uniformitarianism.
Purgatorius: a genus of small mammals in the order Plesiadapiformes; possibly
the last common ancestor of primates and plesiadapids; North America, Late
Cretaceous to early Paleocene.
Purpuroidea: a genus of Mesozoic prosobranch gastropods in the clade
Littorinimorpha and extinct family Purpurinidae; traditionally classified in the
Mesogastropoda; generally characterized by a thick, robust shell almost 3 inches
high, with a series of short, blunt spines on the top half and fine spiral ribs on the
bottom; cosmopolitan, Jurassic and Cretaceous.
Pycnodontiformes: “flattened tooth,” an order of laterally compressed actinopter-
ygian fish; flourishing in the Jurassic and Cretaceous, the pycnodonts were small to
medium-sized fish, characterized by their almost circular shape and round and flat-
tened teeth; global distribution, Late Triassic to Eocene.
Pycnogonida: the sea spiders, a class of marine arthropods usually placed in the
subphylum Chelicerata; not arachnids but closer to them than to other arthropods,
they have small bodies and generally 4 pairs of long legs; their size ranges from less
than an inch to 3 feet; cosmopolitan but especially large in the Antarctic, late
Cambrian to Recent.
Pygasteridae: Mesozoic sea urchins, a family of irregular euechinoids in the
extinct order Pygasteroida; in the type genus Pygaster, the large periproct is
keyhole-shaped; Europe, Middle Jurassic to Late Cretaceous.
pygidium: the tail segment of an arthropod, such as a trilobite.
pygostyle: the ploughshare-shaped element of fused bones at the base of the tail of
modern birds.
Pyrotheria: an order of large South American ungulates that includes two small
families, the Pyrotheriidae and the Colombitheriidae; sometimes called “false ele-
phants,” they were characterized by short, columnar limbs and snouts a foot long;
Paleocene to Oligocene.
Q
Quadrijugatoridae: a family of palaeocopan ostracods in the order Beyrichicopina

and superfamily Hollinoidea; Canada, Late Ordovician.
Quagga: a subspecies of African plains zebras, perissodactyls in the genus Equus,
that became extinct in the middle of the nineteenth century.
Quaternary extinctions: the relatively abrupt extinctions of most of the large
mammals in all continents during the middle and late Pleistocene; most of these
extinctions occurred after the arrival of hominins, although some researchers have
pointed also to climate change or other factors as partial causes.
Quaternary Period: the current division of geologic time, including essentially
the Pleistocene Epoch and the current historical epoch, which is often referred to as
the Holocene and dated from about 12,000 years before the present. The Quaternary
is of course marked by the spread of modern hominins, and by the extinction of
many large terrestrial animals.
Quetzalcoatlus: a genus of large pterosaurs in the suborder Pterodactyloidea and
family Azhdarchidae; characterized by a very long neck and a sharp-pointed beak,
Quetzalcoatlus had a wingspan of 40 feet; it closely resembles the Romanian dis-
covery Hatzegopteryx; North America, Late Cretaceous, 68–66 Ma.

R
radiation: the evolutionary pattern of rapid appearance and dispersal of many

diverse species of a lineage.
Radinskya: an obscure genus of primitive mammals generally placed in the
Laurasiatheria; known only from a partial skull and jaw, it was the size of a small
dog and is thought to be evidence of the radiation of the perissodactyls from Asia;
China, late Paleocene.
Radiolaria: a subclass of marine protists in the phylum Retaria, from 0.1 to
0.2 mm in diameter; radiolarians are important stratigraphically in the absence of
calcareous microfossils, but even their siliceous skeletons are dissolved by high
concentrations of carbon dioxide in the ocean; global distribution, Cambrian to
Recent.
Radiolites: a genus of rudistid bivalves in the order Hippuritoida and family
Radiolitidae; up to a foot long, with valves that were unequal in size and shape, it
was one of the unusual horn-shaped bivalves that were important reef-forming
organisms in the Cretaceous; Europe, Asia, and North and South America; Early to
Late Cretaceous.
radiometric dating: the technique of using the known rate of decay of radioactive
isotopes to establish, within certain limits of certainty, the absolute age of strata of
Earth’s surface; used with biostratigraphic information to establish the geologic
time scale.
radula: a row of rasping chitinous teeth on a tongue-like structure behind the man-
dible of mollusks, a synapomorphy of the phylum.
Ramapithecus: see Sivapithecus.
rangeomorphs: a group of fern-shaped Ediacaran forms which may have been
either the first true animals or an extinct stem group related to animals. Dated at
575–560 Ma, just before the Cambrian Explosion of multivariate animal forms, they

248 R
may also be related to the even earlier sponges. At least two described species,
Rangea schneiderhoehoni and Fractofusus misrai, are considered valid. Global
occurrences, notably Russia, Canada, and Australia; Precambrian,
Raoellidae: a still poorly known family of cetartiodactyl mammals in the clade
Whippomorpha; the semi-aquatic Indohyus, the best-known genus, may represent a
step in the transition of whale ancestors back to an aquatic environment; Southeast
Asia, Eocene.
Rapetosaurus: a monospecific genus of titanosaurian sauropod dinosaurs in the
family Nemegtosauridae; R. krausei is known from an unusually complete skeleton
of a juvenile and parts of three other individuals; adults are estimated to have been
up to about 50 feet long, somewhat smaller than the largest titanosaurs; the juvenile
is estimated to have grown fast, probably without any parental care, from about 8
pounds at hatching to about 90 pounds when it died about two months later, proba-
bly from starvation; Madagascar, Late Cretaceous, 70–66 Ma.
Raphidonema: a genus of calcareous sponges in the still-extant subclass Calcaronea
and order Pharetronida; cup-shaped and often twinned, Raphidonema forms large
sponge beds in the Early Cretaceous of England; Europe and Asia, Cretaceous to
Eocene.
Raphinae: a subfamily of birds in the order Columbiformes that became extinct
less than 300 years ago with the disappearance of the dodo (Raphus cucullata) and
the Rodrigues solitaire (Pezophaps solitaria) from islands in the Western Indian
Ocean.
Raphiophoridae: a family of blind trilobites in the order Asaphida and superfam-
ily Trinucleoidea; characterized by very long, trailing genal spines and a single
spine extending forward from the triangular cephalon; cosmopolitan, Ordovician to
middle Silurian.
Raptorex kriegsteini: a dubious species of dinosaurs known from a single fossil,
once considered a possible ancestor of Tyrannosaurus rex but now thought to be an
unclassifiable juvenile specimen, possibly related to Tarbosaurus; China, Late
Cretaceous.
Rastrites: a short-lived genus of scandent graptolites in the order Graptoloidea;
characterized by long, slender, isolated thecae standing out from a single linear
stipe, each theca hooked at the tip; global distribution, early Silurian.
ratites: large, flightless birds of the still-extant infraclass Palaeognathae. Mostly of
Gondwanan origin, they include the modern rheas in South America, ostriches in
Africa, emus in Australia, and probably the extinct moas of New Zealand. The name
is derived from the Latin ratis (raft), chosen because ratites have a flat breastbone
with no keel on their sternum, and significant because the keel is crucial to anchor-
ing wing muscles for flight. Although no fossils are known from the Cretaceous,
genetic research indicates that the order probably originated before the
end-Cretaceous extinction. The earliest-known ratites appeared in the Paleocene
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(see Diogenornis). Several giant genera, such as Dinornis and Aepyornis, became
extinct after the arrival of humans in their habitats.
rauisuchians: a group of crocodilian archosaurs of the Triassic, included in the
clade Pseudosuchia; up to 20 feet long, they were generally carnivorous and among
the largest predators of the period; North and South America and Europe, Early to
Late Triassic.
Razanandrongobe: a monospecific genus of sebecosuchian crocodylians described
in 2017 as a mesoeucrocodylian and the earliest notosuchian; on the basis of several
cranial fragments and several large, serrated teeth, R. sakalavae is estimated to have
been large enough to prey on large dinosaurs; Madagascar, Middle Jurassic, 170 Ma.
Receptaculitidae: a family of benthic marine organisms, currently classified as an
alga; in the past it has been referred to the Porifera and is still controversial; see e.g.
Ischadites; global distribution, late Cambrian to Early Triassic.
Red Gulch Dinosaur Track Site: a location in the Sundance Formation in
Northern Wyoming, displaying numerous footprints made by Middle Jurassic cera-
tosaurs walking on a beach of fine, well-sorted sand.
Redlichiida: an early order of relatively flattened trilobites; characterized by a
large cephalon, a many-segmented thorax, and a small pygidium; the Redlichiida
appeared at about the same time as the Ptychopariida and Corynexochida; global
distribution, early to late Cambrian.
Remingtonocetidae: a family of archaeocete whales in the family Protocetidae;
similar in appearance to amphibians like the crocodile, they had short limbs and a
long, narrow skull with nostrils near the front; they could walk on land but were
mainly aquatic; see also Ambulocetus and Kutchicetus; Asia, early to middle
Eocene.
Repenomamus robustus: a small eutriconodont mammal in the family
Gobiconodontidae. About the size of an opossum or badger, it is especially notable
for a Jehol specimen of R. robustus with the remains of a baby psittacosaur in its
stomach, the first direct evidence that mammals were competing for food with dino-
saurs at the beginning of the Cretaceous. Bearing characteristics of reptiles as well
as mammals, Repenomamus existed also as a much larger species, R. giganticus,
which was 3 feet long and weighed more than 30 pounds. China, Early Cretaceous,
130 Ma.
replacement: the complete supplanting of minerals in a fossil by different miner-
als coming from the surrounding sediment, usually resulting in increased mass and
hardness, and often preserving anatomical structure in detail; in exceptional circum-
stances, the organism may be replicated in three-dimensional detail, especially by
pyrite, marcasite, calcium phosphate, kaolinite, or other relatively soft minerals; see
also taphonomy.
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Reptilia: a paraphyletic class of tetrapod animals that includes snakes, lizards,

crocodiles, and turtles; reptiles are characterized by a tough, horny skin covered
with scales, scutes, or plates; the class excludes some descendants of tetrapods,
namely mammals and birds; global distribution, Late Carboniferous to Recent.
reticulate or reticulated: having veins, threads, fibers, scales, or pigment creating
a net-like appearance.
Reticuloceratidae: an important family of ammonoids in the order Goniatitida
and superfamily Gastrioceratoidea; several species of the genus Reticuloceras are
index fossils in the Carboniferous; see also Arkanites; North America, Europe, and
Asia; Late Carboniferous.
Revueltosaurus: a genus of pseudosuchians originally considered an ornithischian
dinosaur but now classified as a sister taxon of the aetosaurian suchians; well-
armored with a coat of bony plates and a tail club, it is known principally from
several partial skeletons found in or near the Revueltosaurus Quarry in Petrified
Forest National Park; Southwestern United States, Late Triassic, 230–205 Ma.
Rhabdinopora: a genus of planktic colonial graptolites in the early order
Dendroidea and family Anisograptidae; Rhabdinopora is thought to be the ancestor
of the large order Graptoloidea; characterized by a conical reticulate colony with
many parallel stipes that branch and bifurcate; the stipes are connected by dissepi-
ments, forming a net-like surface and superficially resembling the bryozoan
Fenestella; global distribution, Early Ordovician.
Rhabdinopora flabelliformis: “fan-shaped,” a well-known species of graptolites
long considered benthic and classified as Dictyonema flabelliforme but now known
to be planktic; important index fossil in determining the date of the Cambrian-
Ordovician boundary; Europe, Early Ordovician.
Rhabdopleura: a still-extant genus of small benthic colonial animals in the class
Pterobranchia and family Rhabdopleuridae; it is a worm-like, sessile suspension
feeder, probably related to the extinct graptolites; global distribution, Cambrian to
Recent.
Rhachitomi: a group of late Paleozoic amphibians in the order Temnospondyli;
probably derived from the Eutemnospondyli and ancestral to the Stereospondyli;
cosmopolitan, Early Carboniferous to Early Cretaceous.
rhamphorhynchoids: the basal pterosaurs that gave rise to the pterydactyloids in
the Middle Jurassic. Previously considered a suborder, the group is paraphyletic, but
the term is still used to refer in general to the early pterosaurs. Although all could
fly, they were generally much smaller than the pterydactyloids, and they also dif-
fered in having a long tail; most were well-toothed. The rhamphorhynchoids
appeared about 230 Ma; among the earliest species are Preondactylus, Peteinosaurus,
Dimorphodon (q.v.), and possibly the poorly known Faxinalipterus. The best known
of the group is Rhamphorhynchus, which appeared later, about 151 Ma. Global dis-
tribution, Late Triassic (Norian) to Cretaceous.
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Rhamphorhynchus: a genus of fish-eating pterosaurs in the family

Rhamphorhynchidae; Rhamphorhynchus was only slightly larger than a crow but
had a wingspan of 4–5 feet and a very long tail, which ended in a characteristic
diamond-shaped vane; Europe and Africa, Late Jurassic, 152–150 Ma.
Rhenanida: an order of scaly marine placoderms; flattened and ray-like, they were
bottom-dwelling predators, with armor consisting only of unfused tubercles and
scales; their fossil record is sparse but worldwide, Early to Middle Devonian.
Rhenops: an unusual genus of phacopid trilobites in the large family Acastidae;
characterized by stout genal spines, protruding eyes, and two dozen pairs of legs;
Europe and South America, Middle to Late Devonian.
Rhineuridae: a still-extant family of burrowing squamatans in the suborder
Amphisbaenia; this family of worm-like lizards is found in Cenozoic strata across
North America but is now known only as one genus in Florida; middle Paleocene to
Recent.
Rhinocerotidae: a still-extant family of large perissodactyl mammals in the subor-
der Ceratomorpha and superfamily Rhinocerotoidea; this large family comprises
about 40 extinct genera as well as the modern rhinoceroses; late in their evolution,
the latter group developed horns; North America, Europe, Asia, and Africa; Eocene
to Recent.
Rhipidistia: a still-extant subclass of lobe-finned bony fishes in the class
Sarcopterygii; it includes the lungfishes, which are still represented by several gen-
era, and also includes numerous diverse extinct orders; probably ancestral to the
tetrapods; Early Devonian to Recent.
Rhizobiales: “root life,” an order of symbiotic bacteria, one of several groups
involved in the process of producing the ammonium and nitrates used in fertilizers
and gunpowder. These bacteria benefit certain plant subfamilies of the order Fabales
(principally the Papilionoideae and Mimosoideae) by enabling them to fix atmo-
spheric nitrogen in the form of ammonium and nitrates, eventually enriching the
soil; the bacteria benefit from flavonoids secreted by the plants. The rhizobia are
known to have existed in the Precambrian, possibly for billions of years, and the
type species Rhizobium leguminosarum has been thoroughly studied. Worldwide,
Precambrian to Recent.
Rhizodontida: an order of large predatory freshwater fish in the class Sarcopterygii;
one species, Rhizodus hibberti, was about 23 feet long, probably the largest fresh-
water fish known; North America and Europe; Middle Devonian to Late
Pennsylvanian.
rhizome: a root-like plant stem that generally grows horizontally, on or under the
ground.
Rhizomyinae: a still-extant subfamily of myomorph rodents in the family
Spalacidae, including the Asian bamboo rats and some African mole rats; Miocene
to Recent.
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Rhombifera: a class of blastozoan echinoderms in the superclass Cystoidea, char-

acterized by perforated plates in the calyx and rhombic thecal plates shared equally
between adjacent plates; similar to the Diploporita; North America, Europe, and
Asia; Early Ordovician to Late Devonian.
Rhynchocephalia: an order of lizard-like reptiles in the subclass Lepidosauria; it
includes the once-numerous sphenodonts, which are now extant only in the tuatara
(two species of Sphenodon) of New Zealand; global distribution, Middle Triassic to
Recent.
Rhyncholepis: a genus of early jawed fish in the order Anaspida; named “beaked
scale” for its thick, bony scales; Scandinavia, late Silurian, 426–423 Ma.
Rhynchonellata: a large, still-extant class of articulate brachiopods in the subphy-
lum Rhynchonelliformea, generally with biconvex calcite shells; global distribu-
tion, early Cambrian to Recent.
Rhynchonellida: a still-extant order of articulate brachiopods in the class
Rhynchonellata and superfamily Rhynchonelloidea; characterized by the absence
of brachidia and by astrophic hinges; generally wedge-shaped, often resembling a
nut; global distribution, Ordovician to Recent.
Rhynchonelliformea: term now given to the articulate brachiopods, replacing the
class Articulata and revised as a subphylum comprising the extinct classes Chileata,
Kutorginata, Obolellata, and Strophomenata, and the extant Rhynchonellata; in
addition to articulation, they are characterized by a fibrous secondary shell layer.
Rhynchosauria: “beak lizard,” an order of herbivorous diapsid reptiles, primitive
archosauromorphs; the early genera were small and lizard-like, while some toward
the end of the period grew to more than 6 feet long; these later genera had bulky
bodies, a strong beak, and wide skulls, with some teeth becoming broad toothplates
for cutting up plant material; global distribution, Early to Late Triassic.
Rhynia: a genus of early vascular land plants in the class Rhyniopsida; known only
from the Rhynie Chert, where it is a common rhizomatous plant with stems up to 8
inches high; Scotland, Early Devonian.
Rhynie Chert: a Lower Devonian Konservat-Lagerstätte that contains many early
arthropod and plant fossils. It is especially important because it represents an early
stage of the colonization of land by marine life, even the first evidence of herbivory
by sap-sucking mites. A short turf a few inches high was created by early land
plants, including bryophytes and the first vascular plants (see Rhynia), as well as
pre-vascular plants with mixed bryophytic characteristics, such as Aglaophyton and
Horneophyton. The informal term “rhyniophytes” is often used to refer to the
Rhynie flora in general. The Rhynie fossils are mainly found in chert deposited
around hot springs, and many are three-dimensional. Some of the significant ani-
mals are the earliest-known insect (Rhyniognatha hirsti), crustaceans (see
Lepidocaris), trigonotarbids, the oldest mites (see Acari), and harvestmen (see
Opiliones). Scotland, 410–405 Ma.
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Rhyniella praecursor: a famous hexapodal arthropod of the Rhynie Chert, placed

in the order Collembola; among the earliest-known proto-insects, 2 mm long and
wingless; Scotland, Early Devonian.
Rhyniognatha hirsti: the oldest known insect, a species known from specimens
found in the Rhynie Chert, especially a dicondylic pair of mandibles; the presence
of condyles and the short, triangular mandibles may indicate that R. hirsti had
wings; Scotland, Early Devonian.
Rhyniopsida: a class of primitive leafless plants in the division Tracheophyta;
among the simplest of all vascular plants, they bore sporangia near the tips of
branches; first known from the Rhynie Chert assemblage; global distribution, Early
Devonian.
Richthofenia: a genus of articulate strophomenid brachiopods in the order
Productida and family Richthofeniidae; similar to cup-shaped corals in shape and
also in its reef-forming habit, with specializations for encrustation on hard sub-
strates; North America, especially Texas, early Permian.
Riojasaurus: a genus of middle-sized early sauropod dinosaurs in the family
Riojasauridae, probably related to the slightly smaller melanorosaurs of South
Africa. Herbivorous and with a heavy body and massive legs, Riojasaurus was one
of the very few sauropods in South America. About 33 feet long and weighing about
10 tons, it was mainly quadrupedal but also bipedal. Argentina, Late Triassic,
230–210 Ma.
Rissooidea: a still-extant paraphyletic superfamily of very small marine and fresh-
water gastropods in the clade Littorinimorpha; generally cylindrical or conical in
shape, the group is extremely diverse and widespread; global distribution, Jurassic
to Recent.
Riversleigh: a Cenozoic Konservat-Lagerstätte in northwestern Queensland that
has produced exceptionally well-preserved specimens of bats, marsupials, insects,
and other Australian fauna and flora; many of the fossils are three-dimensional, hav-
ing been preserved in soft, uncompressed limestone; Australia, late Oligocene to
late Miocene.
Robertia: an early monospecific genus of herbivorous dicynodont therapsids in the
family Pylaecephalidae; about a foot long, R. broomiana had two sharp canine teeth
and a horny beak, probably used in digging; related to Diictodon; South Africa,
middle to late Permian, 265–260 Ma.
rock varnish: a dark biogeochemical encrustation on exposed rock in almost any
terrestrial weathering environment, but most commonly found as desert varnish in
arid or semi-arid sites; consisting primarily of iron and manganese oxides, clay, and
carbonates secreted by bacteria, it is useful in identifying paleoenvironments.
Rodhocetus kasrani: a species of archaeocete whales in the family Protocetidae,
an important recent discovery in strata deposited in deep water about 50 Ma,
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representing the earliest-known deep-ocean whale. Notable for its unfused verte-
brae that allowed it to swim principally with its tail, as do modern whales, Rodhocetus
still bore short but powerful hindlimbs and a pelvis connected to its sacrum. Similar
to the slightly more recent Protocetus except that the latter could not have supported
its own weight on land, whereas Rodhocetus may have been able to do so and there-
fore represents the farthest-known evolution of ancestral whales before they lost all
useful terrestrial characteristics. It is also notable because its blowhole was high on
its head, a step in the direction of modern whales, all of which have a blowhole on
top of the head. Pakistan, early Eocene.
Rodinia: “Motherland,” name given to a Proterozoic supercontinent that existed
between 1300 and 650 Ma; it consisted of fragments from an earlier supercontinent
called Columbia or Nuna that had drifted apart between 1800 and 1500 Ma; Rodinia
centered on Laurentia, which was on the Equator.
Rodrigues solitaire: see Pezophaps solitaria and Raphinae.
Romer’s Gap: the paucity of tetrapod and arthropod fossils in the period between
361 and 340 Ma at the beginning of the Carboniferous period, also called the
Tournaisian Gap. The first terrestrial vertebrates appeared at the end of the Devonian,
around 370 Ma, but their fossil record in the early Mississippian is very thin. (See
also Tetrapoda and Tulerpeton.) Terrestrial vertebrates appear in great numbers in
the middle Mississippian, so their radiation at the beginning of the period must have
been quite rapid. This radiation has drawn many researchers since Alfred Romer
himself, especially Robert Carroll, Per Ahlberg, and Jennifer Clack at the end of the
twentieth century.
Rosamygale grauvogeli: a species of funnel spiders in the infraorder
Mygalomorphae and extant family Hexathelidae; less than half an inch long, R.
grauvogeli is the oldest known mygalomorph spider; known only from the Grès à
Voltzia, France, Early Triassic.
Rostroconchia: a class of mollusks superficially similar to bivalves and once pro-
posed as ancestral to them; both, however, along with the brachiopods, appeared
close to the same time in the Cambrian Explosion near the beginning of the Paleozoic
Era, and their inter-relationships remain unclear. From 1 to 6 inches wide, the ros-
troconchs had a taco-like shell and a rostrum, a tube extending outside the shell,
possibly to filter the water. They diversified rapidly worldwide through the
Ordovician, then declined in comparison to the bivalves, and became extinct at the
end of the Permian.
Rotalipora globotruncanoides: a species of planktic foraminiferans whose first
appearance marks the beginning of the Late Cretaceous at 100.5 Ma; France.
Rubeosaurus: a genus of ceratopsian dinosaurs in the subfamily Centrosaurinae;
named “thornbush” for an array of horns and spikes, with a single short horn on its
snout and longer horns on its neck frill; known from two specimens of skull frag-
ments, it is estimated at about 19 feet and 2 tons; North America, Late Cretaceous,
75 Ma.
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Rubidgea: a monospecific genus of therapsids in the family Gorgonopsidae; as

long as 11 feet, R. atrox is notable for canine teeth longer than those of Tyrannosaurus
rex; known only from two skulls; South Africa, late Permian.
rudists: see Hippuritoida.
Rugops primus: a species of ceratosaurian theropod dinosaurs in the family
Abelisauridae; known only from a single skull, it is estimated at 15–20 feet long and
thought to be carnivorous but probably a scavenger; Niger, Late Cretaceous,
96–94 Ma.
Rugosa: the “wrinkled” corals, a large order of anthozoan cnidarians in the sub-
class Hexacorallia; also called Tetracoralla. The Rugosa and the Tabulata were the
most diverse and widespread coral reef-builders of the Paleozoic. Mainly solitary
and conical or horn-shaped, but also often colonial, the rugose corals were com-
posed of calcite, as opposed to the more easily dissolved aragonite of the modern
Scleractinia. The first undoubted Rugosa appeared in the Middle Ordovician, and
they became extinct along with the Tabulata at the end of the Permian.
Rugosodon eurasiaticus: a species of multituberculate mammaliaforms in the
order Paulchoffatiidae; found in the Tiaojishan Formation and dated at about
160 Ma, Rugosodon is so called because of its distinctively wrinkled teeth; it was
omnivorous, and its chipmunk-sized body was 6–7 inches long and adapted to tree-
dwelling; China, Late Jurassic.
Ruminantia: a suborder of artiodactyl ungulates that have a four-part stomach to
digest tough plant material. The Ruminantia include cattle, sheep, goats, antelopes,
deer, pronghorns, giraffes, and many extinct pecorans. A smaller group of ungulates
that includes tylopods (the camelids), and macropods (kangaroos and wallabies) are
considered “pseudo-ruminants” with a slightly different digestive system. Global
distribution, early Eocene to Recent.
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Sacabambaspis: a genus of early pteraspidimorph craniates in the order

Arandaspidiformes, closely related to Arandaspis and among the earliest whole-
body specimens of ostracoderms. Jawless and 10 inches long, it had a laterally
flattened body and an extended tail with a small fin web at the end, but no paired
fins. Its head was armored with bony plates and the body was covered with scales.
First found in Bolivia, it is known from several Gondwanan locations: Argentina,
Australia, and the Arabian Peninsula; Middle to Late Ordovician.
Saccorhytus coronarius: the oldest known species of deuterostomes, described in
2017 from a group of 45 individuals; only 1 mm long, its globular body seems to
have only one opening, which is surrounded by conical protuberances; China, early
Cambrian, 540 Ma.
saddle: an anterior-pointing fold in the suture line of an ammonoid.
Sagenocrinites: a genus of crinoids in the order Sagenocrinida; characterized by
an ovoid crown and a conical cup half the height of the crown and about an inch in
diameter, with branching arms well differentiated from the calyx; North America
and Europe, Silurian.
Sagenopteris: a genus of seed-fern leaves in the family Caytoniaceae; global dis-
tribution, Early Triassic to Early Cretaceous.
Saghatherium: a genus of primitive hyracoid mammals in the extinct family
Pliohyracidae and subfamily Saghatheriinae; it was scansorial and herbivorous, and
somewhat similar to the modern hyraxes of the Procaviidae; Fayum, Egypt, late
Eocene or early Oligocene.
Saharagalago: a genus of lorisid primates in the suborder Strepsirrhini and family
Galagidae; discovered in 2002 in Fayum, Egypt; dated between 40 and 37 Ma, the
early bushbabies Saharagalago and a cousin Karanisia pushed back the date of the
earliest lorisiforms by 20 million years; they represent the earliest-known examples
of the characteristic strepsirrhine toothcomb; Fayum, Egypt, middle Eocene.

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Sahelanthropus tchadensis: a species of hominids found near Lake Chad in 2001;

this early hominid lies near the divergence of humans from chimpanzees, although
probably predating it slightly; it shows a mix of human and chimpanzee characters
and at 6.9 million years old is the oldest known hominid.
St. George Dinosaur Discovery Site: a set of exceptionally well-preserved Early
Jurassic dinosaur tracks now enclosed in a museum on the Johnson Farm in south-
western Utah.
Salientia: “jumpers,” the clade that includes the modern frogs, Anura, and their
most closely-related ancestors.
Saltasaurus loricatus: a species of titanosaurian sauropod dinosaurs in the mono-
specific family Saltasauridae; about 40 feet long and weighing 8 tons, it was unusual
for a sauropod in having osteoderms and armor plates covering its upper body and
sides; Argentina, Late Cretaceous, 71–69 Ma.
Salterella: an enigmatic small, conical fossil, widespread in North America in the
Cambrian; its stratified walls are composed of agglutinated grains of calcite; it
superficially resembles some early cephalopods but is now tentatively referred to
the proposed short-lived phylum Agmata; North and South America and Europe,
early Cambrian to Late Ordovician.
Saltoposuchus connectens: the type species of a genus of relatively small, bipedal
sphenosuchian reptiles in the superorder Crocodylomorpha, 3–5 feet long and
weighing less than 35 pounds; Europe, Middle Triassic.
Saltopus: “leaping foot,” a monospecific genus of early dinosauriform reptiles,
placed by some in the family Silesauridae. The enigmatic S. elginensis is poorly
known from a partial skeleton, mainly cast impressions. It has been studied inten-
sively since its discovery in 1910 and classified variously as a stem-dinosaur or a
dinosauriform outside the clade Dinosauria. In a re-classification proposal in 2017,
it figures as possibly ancestral to dinosaurs. Scotland, Late Triassic, 228 Ma.
Samotherium: a genus of early giraffes similar in appearance to the modern okapi,
with two ossicones extending upward from the skull; larger than the okapi but
smaller than modern giraffes, Samotherium is a good example of a transitional
genus; Europe, Asia, and Africa; Miocene to Pliocene.
Sanmiguelia lewisii: a species of early plants that may belong to a stem-group of
angiosperms, known from several localities in Western North America. One espe-
cially well-preserved autochthonous specimen was found in a Late Triassic stream-
side paleosol near Palo Duro Canyon in Northwest Texas. Buried in its growth
position, it has several structures similar to angiosperms. North America, Late
Triassic to Early Jurassic.
Santa Maria Formation: a Triassic site in Southern Brazil with strata dating from
the Ladinian to Carnian Stages, similar to those of the Chañares Formation of north-
western Argentina; notable for specimens of early dinosaurs, aetosaurs, and cyn-
odonts; Middle to Late Triassic.
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Santana Formation: a Lower Cretaceous Konzentrat-Lagerstätte in the Araripe

Basin of Northern Brazil. It is dated uncertainly between 112 and 106 Ma, slightly
later than the neighboring Crato Formation. The Santana fossils, occurring inside
limestone concretions, include complete specimens of fish in great numbers; partial
specimens of pterosaurs with soft-tissue preservation of webbing; dinosaurs, includ-
ing spinosaurids and coelurosaurs; reptiles (see Araripesuchus); and a few insects
and other invertebrates.
Sarcopterygii: “flesh fin,” lobe-finned fish in the superclass Osteichthyes, the
bony fishes. Previously called Crossopterygii, the Sarcopterygii appeared in the late
Silurian along with other osteichthyans. They dwindled during the Mesozoic,
eclipsed by the many very successful osteichthyans and chondrichthyans. The
Sarcopterygii are still represented by several genera, including the coelacanths in
the subclass Actinistia and the lungfishes in the subclass Rhipidistia. Among the
many extinct sarcopterygian taxa, the closely related tetrapod lobefins of the Late
Devonian are especially significant evolutionarily as the first terrestrial vertebrates.
See also Actinopterygii and Tetrapoda.
Sarcosuchus imperator: a very large crocodilian of the Cretaceous, first discov-
ered in 1964 in Niger in the Ténéré Desert. From 37 to 40 feet long, with a head
more than 5 feet long and jaws about 4 feet long, it weighed around 9 tons, about
1.5 times as much as an African elephant. Sarcosuchus had a bulbous protuberance
on the end of its snout, probably used to locate prey by smell or sound. North Africa
and South America, Early to Late Cretaceous.
Sarmientosaurus musacchioi: a species of titanosaurian sauropod dinosaurs
found in Patagonia and described in 2016; estimated at about 40 feet long and 10
tons, it is known from an almost complete skull and several articulated vertebrae
that may indicate a drooping posture of the head; also characterized by very large
eyes; Argentina, Late Cretaceous, about 95 Ma.
Sasayamamylos: a genus of basal eutherian mammals in the order Asioryctidae;
insectivorous, known only from a skull found in Japan in 2007 and described in
2013, S. kawaii may be the earliest-known example of a modern mammalian denti-
tion; Early Cretaceous, 112 Ma.
Saurichthyidae: a speciose family of Mesozoic ray-finned chondrostean fish in
the order Saurichthyiformes. They were about 3 feet long and very slender, with
symmetrical dorsal and anal fins near the tail, a shape that probably made them fast
and efficient predators. They may have originated toward the end of the Permian but
were common throughout the Triassic and persisted at least to the Middle Jurassic
(see Saurorhynchus). North America, Europe, Asia, Australia, and Madagascar;
Early Triassic to Middle Jurassic.
Saurischia: one of the basic divisions of dinosaurs, characterized by a three-
pronged pelvic structure similar to that of most reptiles, with a pubis bone pointing
downward and slightly forward. The earliest saurischians (such as Eoraptor and
the Guaibasauridae) appeared during the Carnian Age of the Late Triassic. The
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sauropods are the most prominent group within the Saurischia, while the theropods
are variously considered as saurischians or part of some other division (see
Ornithoscelida). Global distribution, Triassic to Late Cretaceous.
Saurolophus: “lizard crest,” a genus of large hadrosaurid dinosaurs in the subfam-
ily Saurolophinae; 30–40 feet long, they were characterized by a “duck bill” and a
backward-extending cranial crest of unknown function; see also Parasaurolophus;
North America and Asia, Late Cretaceous, 71–68 Ma.
Sauropoda: an infraorder of quadrupedal saurischian dinosaurs in the suborder
Sauropodomorpha. Semi-aquatic and herbivorous, the sauropods had very long
necks, long tails, and a heavy body supported by four pillar-like legs. Estimates of
sauropod weight vary greatly, but conservative estimates range from 50 to 70 tons
for the largest. Global distribution, Late Triassic to Late Cretaceous.
Sauropodomorpha: a clade of herbivorous saurischian dinosaurs that includes the
infraorders Plateosauria and Sauropoda and their immediate ancestors; with repre-
sentatives dated between 231 and 66 Ma, the sauropodomorphs were the dominant
terrestrial herbivores worldwide from the Late Triassic to the Late Cretaceous.
Sauroposeidon: a genus of titanosaurian sauropod dinosaurs; at about 60 feet high
it was possibly the tallest dinosaur, and it had a longer neck than any other dinosaur,
conservatively estimated as 40 feet long, with some vertebrae 5 feet long; weighing
about 60 tons, it was significantly larger than Brachiosaurus; North America, Early
Sauropterygia: a superorder of aquatic diapsid reptiles characterized by pectoral
and pelvic girdle adaptations that supported strong flippers. Including the notho-
saurs, plesiosaurs, and pliosaurs, the sauropterygians arose at the beginning of the
Triassic and diversified worldwide, with the plesiosaurs surviving into the Jurassic
and the pliosaurs appearing in the Early Jurassic. The last of the sauropterygians
disappeared at the end of the Cretaceous, except that some research now indicates
that modern turtles may be distant descendants. Early Triassic to Late Cretaceous.
Saurorhynchus: a genus of chondrostean fish in the order Saurichthyiformes and
family Saurichthyidae; formerly called Acidorhynchus; gar-like in appearance, it
was 2–3 feet long and slender, with symmetrical dorsal and anal fins near the tail;
North America and Europe, Early to Middle Jurassic.
savanna: term used to refer to a wide range of environments between dense forest
and desert, usually referring to openly wooded grassland.
Savannasaurus: a monospecific genus of titanosaurian sauropod dinosaurs
described in 2016; S. elliottorum was 50 feet long and is the most complete speci-
men of a sauropod found in Australia; Cenomanian, Late Cretaceous, 100–94 Ma.
scandent: of vines and graptolites: having a climbing habit.
scansorial: of animals: able to climb trees, spending at least part of the time in an
arboreal environment.
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Scansoriopteryx: see Epidendrosaurus.

Scaphites: a genus of small, loosely coiled ammonoids in the order Ammonitida
and suborder Ancyloceratina, known worldwide. Its species are generally character-
ized by a boat-shaped shell 2–5 inches long, with an involute and compressed juve-
nile stage, then a straight segment in the middle, and an erect terminal stage bending
back over the juvenile stage like a hook. Probably a slow swimmer, mainly benthic;
several species are index fossils in the Late Cretaceous.
Scaphopoda: “boat foot,” a still-extant class of infaunal marine mollusks; called
“tusk shells,” they are characterized by a tooth-shaped, curving, tapering shell, open
at both ends; from 1 to 4 inches long, they live in the soft substrate offshore, beyond
the intertidal zone; they are probably the latest class of mollusks to appear; global
distribution, Ordovician to Recent.
Scenella: an enigmatic genus of limpetlike animals variously considered monopla-
cophorans, gastropods, helcionellids, and even hydrozoans; about half an inch
across, it is characterized by a high, curving, cap-shaped shell with radial and con-
centric striations; global distribution, Cambrian to Ordovician.
schizochroal: a type of compound eye found only in some phacopid trilobites; it
has up to 700 lenses with separate corneas, with larger lenses than in the holochroal
eyes of most trilobites.
Schizoneura: a genus of small, slight sphenophyte trees in the order Equisetales,
related to the Calamitaceae; global distribution, early Permian to Late Jurassic.
Schloenbachiidae: a family of ammonoids in the order Ammonitida and super-
family Hoplitoidea; characterized by a complex ammonitic suture and a compressed
form; irregular, some genera smooth and involute and others strongly ribbed and
evolute; North America, Europe, and Asia; Cretaceous, 113–94 Ma.
Schubertellidae: a family of foraminiferans in the superfamily Fusulinoidea; they
diversified greatly toward the end of the Permian but became extinct at the end of
the period; North America, Europe, and Asia; Pennsylvanian to late Permian.
Schwagerinidae: a family of large, complex foraminiferans in the superfamily
Fusulinoidea; the genus Schwagerina is important in petroleum geology, providing
several index fossils in lower Permian strata; North and South America, Europe, and
Asia; Late Carboniferous to late Permian.
Sciuridae: the still-extant family of rodents that includes modern squirrels, so
similar to their Miocene ancestors that they can be considered “living fossils”;
global distribution, Oligocene to Recent.
Sciurumimus albersdoerferi: “squirrel mimic,” a species of feathered theropod
dinosaurs in the suborder Coelurosauria; reported in 2012 and known only from a
juvenile specimen, 28 inches long; the tail feathers resemble a squirrel’s tail fur;
Germany, Late Jurassic, 150 Ma.
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Scleractinia: “hard-rayed,” a still-extant order of anthozoan cnidarians, corals

placed in the subclass Hexacorallia but with obscure origins. Often called stony
corals, they are mostly colonial in relatively shallow water, with some solitary g enera
found in deeper water, some even abyssal. When the mainly calcitic tabulate and
rugose corals disappeared at the end of the Permian, there were no important reef-
building animals until the aragonitic scleractinians appeared in the Middle Triassic
and became a dominant reef-builder late in the period; barely escaping extinction,
they began to spread globally in the Cretaceous. Only one purely calcitic genus,
Coelosimilia (Late Cretaceous, about 70 Ma), is known. Middle Triassic to Recent.
scolecodonts: the chitinous jaws of marine polychaete annelid worms; as large as
a few millimeters, scolecodonts appeared in the Cambrian, were common in
Ordovician and middle Paleozoic strata, became rare at the end of the Permian,
recovered in the Mesozoic, and again became rare at the end of the Cretaceous; at
least seven families of modern polychaetes have chitinous teeth that would be con-
sidered scolecodonts if fossilized; Cambrian to Recent.
Scolosaurus: a monospecific genus of armored ornithischian dinosaurs in the sub-
family Ankylosaurinae; called “horned toad dinosaur” for the many spines of vari-
ous sizes that protrude from its head, body, and tail, S. cutleri grew as long as 20 feet
and weighed 2 tons; it is the oldest known North American ankylosaur, known only
from a single skeleton dated at about 76.5 Ma; Alberta, Late Cretaceous.
scute: an external ossicle, a plate of bone embedded in the skin and covered with
horn, common in many dinosaurs and some mammals.
Scutosaurus: a genus of large pareiasaurian reptiles, the largest herbivore of the
late Permian; about 10 feet long, it was heavily armored with scutes, osteoderms,
and small skull spikes; Russia, late Permian.
Scyphozoa: the jellyfish, a class of cnidarians; they probably date back to the
Precambrian, but fossils are rare because of the lack of hard parts; one of the earliest
well-identified is Essexella, 3–4 inches wide, which is known from many specimens
in the Mazon Creek fauna of the Late Carboniferous.
sea lilies: common name for crinoids that are attached to the substrate by a stalk.
sea pens: see Pennatulacea.
sea squirts: see Tunicata.
sectorial tooth: a single-cusped, pointed tooth, especially of non-human primates,
often with a cutting edge capable of a scissors-like action in occlusion.
seed fern: see Pteridospermatophyta.
Segnosaurus: a genus of herbivorous theropod dinosaurs in the family
Therizinosauridae; about 20 feet long and weighing a little more than a ton, they are
characterized by long, flattened claws on the large forelimbs, strong hindlimbs, and
cutting denticles on the edges of some teeth; Asia, Late Cretaceous, 90 Ma.
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Seismosaurus: see Diplodocus.

Seitaad: a monospecific genus of sauropodomorph dinosaurs found in Southern
Utah in 2004 and described in 2010, probably an early sauropod; S. ruessi, known
only from a headless but articulated skeleton, was 10–25 feet long and was probably
herbivorous; United States, Early Jurassic, 185 Ma.
Selachii: the sharks, a superorder of chondrichthyan fish in the subclass
Elasmobranchii. The only Paleozoic elasmobranchs, the first sharks appeared about
420 Ma and diversified rapidly, becoming the dominant predators in the oceans in
the Devonian. The best-known early shark is the North American Cladoselache, and
another genus similar in appearance to modern sharks is Denaea. Two with very
unusual characters were Stethacanthus and Falcatus, q.v. The selachians declined
in the Carboniferous and Permian but rebounded in the Jurassic. Most modern
sharks have their roots in the Cretaceous, along with the other elasmobranchs.
Global distribution, Silurian to Recent.
Selaginella: a still-extant genus of spikemosses in the division Lycopodiophyta
and class Isoetopsida, related to the extinct giant club mosses of the Mesozoic;
global distribution, Silurian to Recent.
selenodont dentition: “crescent tooth,” low-crowned teeth with a hard chewing
surface and cusps forming a crescent shape if viewed from above; common in rumi-
nant mammals.
Selenopeltis: an Ordovician genus of trilobites in the order Odontopleurida and
family Odontopleuridae; from 4 to 6 inches long, characterized by many long pleural
spines and a rectangular carapace; Europe and North Africa, Early to Late Ordovician.
Selkirkia: a genus of predatory priapulid worms in the family Selkirkiidae, known
primarily from the Burgess Shale; about 3 inches long, it lived in a tube, oriented
vertically in the sediment, extending its proboscis to capture prey; North America
(possibly also South America and Antarctica), middle Cambrian.
Semionotus: the type genus of a family of early neopterygian fish in the order
Semionotiformes; only 5 inches long, it was mainly freshwater but is also found in
marine deposits; it is notable for its peg-like teeth and an almost-square dorsal fin
pointing backward; cosmopolitan, Middle Triassic to Early Cretaceous.
septum: a partition inside the skeleton of an animal, especially in corallites and the
shells of ammonoids, belemnites, and nautiloids.
Sequoia jeholensis: the earliest-known sequoia; North America, Europe, and Asia;
Jurassic to Oligocene.
Serridentinus: see Gomphotherium.
sessile: stationary on the bottom sediment and often anchored directly to the
substrate.
setae: slender, rigid, bristle-like extensions on living organisms, usually referring
to bristles or hairs on animals; see also chaetae.
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sexually dimorphic: having distinguishing features between male and female

individuals; dimorphism has often resulted in male and female forms being inter-
preted as different species, a mistake usually corrected by examination of a greater
number of specimens.
Seymouria: a genus of semi-aquatic stem-amniotes in the family Seymouriidae; it
was about 3 feet long and had several reptilian adaptations; related to Discosauriscus,
q.v.; known only from Central North America and Germany, early Permian,
280–270 Ma.
Seymouriamorpha: an order of reptile-like stem-tetrapods, generally placed in the
clade Batrachosauria; North America, Europe, and Asia; early to late Permian.
Shansisuchus: a genus of crocodilians in the family Erythrosuchidae; well-known
from several specimens, Shansisuchus was about 6 feet long; China, Middle Triassic.
Shantungosaurus: a genus of hadrosaurid dinosaurs in the subfamily
Saurolophinae; as long as 54 feet and weighing 16 tons, it is the largest known had-
rosaurid and one of the largest ornithischians; China, Late Cretaceous, 70 Ma.
Sharktooth Hill: one of the largest bone-beds, a densely packed layer of 15-
million-year-old fossils. The site has yielded 40-foot Carcharodon megalodon
remains, mainly teeth, and thousands of other marine species, as well as some ter-
restrial tapirs and ancestral horses. The remains were deposited over a period of
about 700,000 years along the southeastern shore of the Temblor Sea, which cov-
ered California’s long inland valley. United States, middle Miocene.
Sharovipteryx: a genus of small gliding reptiles in the order Protorosauria; known
from one specimen found in the Madygen Formation of Kyrgyzstan in 1965 and
dated at 225 Ma; notable for the membrane attached to its elongated hindlimbs;
Kyrgyzstan, Late Triassic.
Shastasaurus: a genus of ichthyosaurs in the family Shastasauridae. Known from
specimens in California, Oregon, British Columbia, and China, the type species is
S. pacificus. Since Shastasaurus is very different from other ichthyosaurs found in
North America, it is one of the many pieces of evidence of the degree to which ter-
ranes of Western North America have originated in the Pacific Ocean and been
accreted by tectonic forces to the North American continent. North America and
Asia, Late Triassic, 236–204 Ma.
shipworm: see Teredinidae
Shonisaurus: a genus of ichthyosaurs in the family Shastasauridae; about 50 feet
long, they were bulky and dolphin-shaped, with a long, thin snout; known primarily
from a large bone-bed in Nevada that contains 30–40 relatively complete speci-
mens, all lying in the same position; the causes of their deaths are uncertain; North
America and Europe, Late Triassic, 225–210 Ma.
Shumardiidae: a family of minute, blind trilobites in the order Ptychopariida;
about 5 mm long, characterized by a semicircular glabella with deep furrows and
conspicuous lateral lobes; global distribution, late Cambrian to Late Ordovician.
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Shuotherium: a genus of Mesozoic mammals in the family Shuotheriidae; known

from several dental fragments, they share characteristics with monotremes and
eutherians; England and China, Middle to Late Jurassic.
Shuvosaurus: see Effigia.
Siamopithecus eocaenus: a genus of primitive anthropoid primates in the family
Amphipithecidae, found in the 1990s in Thailand; reconstructions based on skull,
jaw, and dental fragments indicate similarities to Aegyptopithecus and Pondaungia;
late Eocene.
Siberian Traps: an area of lava flows, about 870 miles in diameter, northeast of
Lake Baikal in Siberia; the eruptions occurred over a period of less than 1 million
years, beginning about 248 Ma, and have been intensively studied because their date
is so close to the end of the Permian; some researchers believe the eruptions may be
related to the causes of the great extinction that occurred then.
Sidneyia: a monospecific genus of flattened stem-arthropods of the Burgess Shale
fauna; 2–5 inches long, S. inexpectans is one of the larger arthropods at the site; its
glabella was wide and shaped like a ram, and it probably preyed on trilobites, bra-
chiopods, and hyoliths; Canada, middle Cambrian, 508 Ma.
Sigillaria: “seal-marked,” a genus of large lycopodiophyte trees in the order
Lepidodendrales; ranging from 50 to 100 feet high, they are characterized by leaves
growing directly from the trunk and branches; the leaves are indistinguishable from
those of the related genus Lepidodendron, but the leaf-scars are arranged in parallel
vertical rows instead of the spiral arrangement of Lepidodendron; global distribu-
tion, Devonian to Permian.
Silesauridae: a family of dinosauromorphs in the clade Dinosauriformes; the fam-
ily includes several genera of archosaurian reptiles that seem close to the line of
descent to dinosaurs, such as Silesaurus, Asilosaurus, Diodorus, Lewisuchus, and
probably Agnosphitys; South and North America, Africa, and Europe; Middle to
Late Triassic, 245–203 Ma.
Silurian: the third period of the Paleozoic Era, from 444 to 416 Ma. Agnathans
and placoderms were the most abundant vertebrates, but by the end of the period all
principal groups of vertebrates had evolved. Stromatolite and coral reefs covered
large areas of the shallow seas; Silurian strata are mostly calcitic, with some sand-
stones and graptolite shales. The first vascular plants appeared in the middle Silurian,
and plants such as rhyniophytes and clubmosses increased their colonization of the
land, creating a great diversity of new terrestrial ecosystems.
Silvacola acares: a species of very small hedgehogs in the still-extant family
Erinaceidae, found in 2014 in northern British Columbia. It is only 2–3 inches long
but is otherwise similar to modern hedgehogs; whether Silvacola had spines cannot
be determined from the specimens. Canada, early Eocene, 50 Ma.
Simoedosauridae: see Choristodera.
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Sinoceratops: an Asian genus of short-frilled ornithischian dinosaurs in the family

Ceratopsidae; they were herbivorous, medium-sized, and quadrupedal, and are still
the only ceratopsians known from Asia; two species found in 2008 in China have
been identified as S. zhuchengensis and S. formosensis; China, Late Cretaceous,
72–66 Ma.
Sinoconodon: a genus of early mammals in the suborder Cynodontia; one of the
earliest cynodonts, it is probably ancestral to Morganucodon; although similar in
many ways to the non-mammalian synapsids (such as polyphyodonty), it also has
several characters that mark it as a basal mammal (especially in the jaw joint);
China, Early Jurassic, 193 Ma.
Sinodelphys szalayi: a species of metatherians, the earliest-known mammal to dis-
play marsupial traits; about 6 inches long, fur-bearing, and weighing about one
ounce; known from only one specimen found in the Yixian Formation; China, Early
Cretaceous, 125 Ma.
Sinopterus: a genus of small pterodactyloid pterosaurs in the family Tapejaridae;
dated at 120 Ma, it had a wingspan of almost 4 feet; the first tapejarid found outside
Brazil, Sinopterus is known from three specimens found in the Jiufotang Formation;
China, Early Cretaceous.
Sinornithosaurus millennii: a species of theropod dinosaurs in the family
Dromaeosauridae; they were turkey-sized and had bird-like features and short,
downy proto-feathers, but probably did not fly; characterized by single large claws
on hind feet and long claws on its arms, possibly poisonous; one of several feathered
dinosaurs in the Jehol biota; China, Early Cretaceous, 120 Ma.
Sinosauropteryx prima: a species of small theropod dinosaurs that seems to have
had rings of different-colored feathers on its tail and a colored stripe across its eyes.
Researchers base this conclusion on melanosomes, cells that contain the pigment
melanin; the color itself is not visible in the fossil, but electron-microscope analysis
of melanosomes in the follicles gives indirect evidence of a russet color. The feath-
ers were as long as 1.5 inches and were evidently bristly like those of the modern
kiwi. Sinosauropteryx was about 3 feet long but weighed only about 6 pounds; it
had 64 vertebrae in its tail, more than any other theropod. One of several feathered
dinosaurs in the Jehol biota, Early Cretaceous, 127–125 Ma.
Siphonia: a genus of demosponges in the polyphyletic order Lithistida and extinct
family Hallirhoidae; globular, cylindrical, or pear-shaped, they were anchored to the
sea floor by a stalk and a root system or holdfast; the name refers to the main feeding
tube, which runs through almost the whole length of the body; Europe, Early
Cretaceous to late Eocene.
Siphonodella sulcata: a species of conodonts whose first appearance marks the
beginning of the Early Carboniferous (Mississippian) at 358.9 Ma; France.
Siphonotretoidea: a small superfamily of inarticulate brachiopods in the order
Siphonotretida; similar to the Obolellida and still-extant Acrotretida, they are char-
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acterized by a teardrop-shaped, chitinophosphatic shell and an elongate or apical

pedicle valve; global distribution, middle Cambrian to middle Permian.
Sirenia: an order of fully aquatic mammals, some freshwater, placed in the clade
Tethytheria. In addition to several extinct families such as the Prorastomidae and
Protosirenidae, extinct genera of the still-extant Dugongidae and Trichechidae are
known from the Miocene. Global distribution, early Eocene to Recent.
Sirius Passet: a locality in Greenland that has produced soft-bodied fossils similar
to those of the coeval Chengjiang biota; known especially for Halkieria, q.v.; early
Cambrian.
sister group: a phylogenetic term referring to the most closely related taxa in an
evolutionary lineage, those more closely related to each other than either is to other
taxa; the term is used most often in relation to species and genera, but it may also
designate most closely related clades as sister clades.
Sivapithecus: a genus of hominoid primates in the subfamily Ponginae, known
from fossils found at several sites in Asia; formerly called Ramapithecus and
Brahmapithecus. About 5 feet long, they probably spent some time in trees but were
mainly ground-dwelling. There is still disagreement as to their place in hominoid
evolution, but they may be close to the ancestry of modern orangutans. India, mid-
dle to late Miocene, 12.5–8.5 Ma.
Sivatherium: a genus of artiodactyl mammals in the family Giraffidae; similar to
the modern okapi but larger, about 10 feet high and 7 feet at the shoulder, weighing
at least half a ton; it had a pair of wide, antler-like ossicones at the back of its skull
and a smaller pair above its eyes; Africa and India, Pliocene to about 8000 years ago.
Skolithos: a trace fossil ichnogenus, burrows of diameters up to 1.6 inches in
mainly marine and sometimes freshwater strata; the often-vertical burrows are
straight, helical, or slightly curved; they are made by phoronids, worms, insect lar-
vae, other arthropods, and even small vertebrates; Ediacaran to Recent.
“small, shelly fauna”: a “catch-all name that spells frustration” according to
Stephen Jay Gould. The phrase refers to an enigmatic assemblage of spines, plates,
caps, and cups that represent the first steps toward hard skeletons in animals.
Occurring globally from the end of the Ediacaran into the early Cambrian, these
fragmentary bits and pieces have long been a puzzle, but whole specimens have now
been discovered of some of the animals that bore the fragments in life; see for
example Microdictyon sinicum.
Smilodectes: a genus of small adapoid primates in the order Strepsirrhini and sub-
family Notharctinae; it weighed 4–6 pounds and was scansorial; North America,
early Eocene.
Smilodon: “knife-tooth,” a genus of cats in the subfamily Machairodontinae; one of
several genera of large sabertooth cats that flourished in the Pliocene and Pleistocene
but disappeared about 10,000 years ago; global distribution except Australia and
Antarctica; see also Megantereon, Homotherium, and Machairodontinae.
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Smilodon fatalis: the species of sabertooth cats found in the La Brea Tar Pits in
Los Angeles; middle Pleistocene to about 11,000 years ago.
Smilodon gracilis: an early California sabertooth, Pliocene to about 1 Ma.
Smilodon populator: a South American sabertooth cat, the largest known felid at
500–800 pounds; late Pleistocene.
Smith, William: an English geologist, Smith is considered the founder of biostra-
tigraphy for his ground-breaking geologic map of England of 1815, which revolu-
tionized the study of layers of rock, brilliantly combining paleontology, petrology,
and geology. By establishing the principle of faunal succession, Smith’s work imme-
diately inspired widespread use of his techniques in comparing fossils in widely
separated rock outcrops to understand the strata underlying the earth’s surface.
Solanaceae: the nightshades, a large family of still-extant plants in the order
Solanales; a 2017 description of early Eocene (52 Ma) lantern fruits in the genus
Physalis reports the earliest-known occurrence of the family, in Patagonia before
the breakup of South America from Antarctica; Argentina, Eocene.
Solenopora: a disputed genus of calcite-secreting organisms, some seeming to be
algae in the division Rhodophyta and extinct family Solenoporaceae; however, most
specimens are now recognized as chaetetid sponges; most were nodular masses,
while some late Paleozoic and Mesozoic forms occur as encrusting mats; they
appeared in the Cambrian, and some forms may still be extant.
Solitaire: see Raphinae.
Solite Quarry: an important Triassic Konservat-Lagerstätte in Virginia, noted
especially for insects and small reptiles; its lakebed deposits were laid down over a
period of less than 50,000 years; North America, Late Triassic, late Carnian to
Norian.
Solnhofen Limestone: a Late Jurassic Konservat-Lagerstätte in Bavaria, where
several quarries produce lithographic limestone, with fossils of the best quality
coming from those near the towns of Solnhofen and Eichstätt. The extremely
fine-grained limestone layers of the Solnhofen Formation were laid down in a pro-
tected lagoon on the northern edge of the Tethys Sea over a period of half a million
years. The stagnant lagoon contained anoxic and hypersaline areas with few benthic
organisms, and most of the 600 species of fossils are allochthonous, probably
washed into the lagoon by storms. Several crustacean fossils lie at the end of “death
trail” trackways, one 30 feet long (see Mesolimulus and Mecochrinus). Only one
dinosaur, Compsognathus, is known here; it has several skeletal similarities to
Archaeopteryx, the best-known of the Solnhofen fossils. Marine vertebrates such as
ichthyosaurs are known only from poorly preserved fragments, having reached the
lagoon through the coral reefs at its edge during severe storms. Other well-preserved
fossils include pterosaurs, some with toe-webbing and hair-like covering of wing
membranes intact; fish; amphibians; insects; and many arthropods. Germany, Late
Jurassic (Kimmeridgian, 150 Ma).
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Solo man: see Ngandong fauna.

Sonorasaurus: a mid-Cretaceous sauropod of North America, a monospecific
genus of large sauropod dinosaurs in the family Brachisauridae; notable for its
extremely long neck, S. thompsoni grew as long as 55 feet and 30 feet high, its neck
evidently almost as long as the rest of its body and tail; Sonora Desert, Albian to
Cenomanian.
Soomaspis: a monospecific genus of trilobite-like arthropods in the order
Nektaspida; S. splendida resembles a soft-bodied agnostid trilobite, isopygous and
eyeless, and with only three thoracic segments; known only from the Soom Shale of
South Africa, Late Ordovician.
Soom Shale: a Konservat-Lagerstätte near Cape Town, South Africa; several sites
contain exceptionally well-preserved fossils in cold-water deposits of the Upper
Ordovician, 445–444 Ma. Although the formation is unusual for the almost com-
plete absence of benthic organisms, its biota includes several organisms familiar
from the Cambrian Lagerstätten as well as some that became more prominent later
in the Paleozoic, such as the eurypterid Onychopterella. Among other important
specimens are the conodont Promissum pulchrum and the nektaspid Soomaspis, as
well as brachiopods, lobopods, orthocone cephalopods, and chitinozoans.
Soricidae: the shrews, a still-extant large family of small, mouse-like mammals in
the order Eulipotyphla, mainly insectivorous; they first appeared in the late Eocene
and have diversified worldwide.
Spatangoida: the heart urchins, a still-extant order of atelostomate echinoids,
characterized by an oval shape and a deep groove into the mouth, and by the absence
of a lantern; global distribution, Early Cretaceous to Recent.
speciation: development of new species through lineage splitting, resulting in
reproductively isolated branches.
species: 1. biological species concept: a group of actually or potentially interbreed-
ing organisms that are almost completely isolated reproductively from other
organisms, exchanging only a few genes. 2. evolutionary species concept: a single
lineage of ancestor-descendant populations that maintains its own identity distinct
from other lineages.
species duration: The length of time that a species persists varies widely, with
some long-lived species lasting many millions of years, but the average duration is
estimated to be about 4 million years.
speciose: species-rich, containing many species.
Sphaeractinoidea: Mesozoic marine hydrozoans possibly related to the stromato-
poroids and the spongiomorphs; they have long been called “Mesozoic stromatopo-
roids,” but the discovery of spicules in some species has resulted in those species
being reclassified as demosponges and raised the possibility that all sphaeractinoids
may eventually be reclassified.
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Sphaerirhynchia: a genus of articulate brachiopods in the still-extant order

Rhynchonellida and extinct superfamily Rhynchonelloidea; strongly biconvex and
almost spherical, less than an inch across, they attached to the soft substrate with a
functional pedicle; global distribution, Silurian and Devonian.
Sphenacodon: “wedge-point tooth,” the type genus of the synapsid family
Sphenacodontidae; 6–10 feet long, Sphenacodon was slightly smaller than its
cousin Dimetrodon and had only a low sail about a foot high; North America, Late
Pennsylvanian to middle Permian.
Sphenacodontidae: a family of large, advanced pelycosaurian synapsids; they
were characterized by relatively long and narrow snouts and skulls; most genera
have tall, sometimes greatly elongated neural spines; the most familiar sphena-
codontid is Dimetrodon, which is well-known for its sail more than 3 feet high;
North America and Europe, Late Carboniferous to middle Permian.
Spheniscidae: the penguins, a family of large flightless birds of the Southern
Hemisphere; they first appeared in the middle Eocene and diversified rapidly; nota-
ble for having retained the bone structure and musculature of flying birds despite
being flightless, penguins adapted these flight structures to swimming, essentially
flying underwater; middle Eocene to Recent.
Sphenodon: “wedge tooth,” the tuatara, a still-extant genus of lizard-like reptiles
in the family Sphenodontidae, represented by two species in New Zealand, S. punc-
tatus and S. guntheri; they are as long as 3 feet and weigh up to 3 pounds; New
Zealand, Miocene to Recent.
Sphenodontidae: a still-extant family of lizard-like reptiles in the order
Rhynchocephalia. Sphenodonts were numerous in the Jurassic but are represented
today by only one genus, the tuatara of New Zealand (Sphenodon); see also
Planocephalosaurus. Global distribution, Triassic to Recent.
Sphenophyllales: “wedge leaf,” an extinct order of articulate vascular plants in the
class Equisetopsida, related to the modern horsetails; small and slender, they grew
only up to about 3 feet, with jointed stems and linear, forked, or wedge-shaped
leaves; common as fossils in the Carboniferous and early Permian; global distribu-
tion, Early Devonian to Late Triassic.
Sphenopteris: a form genus of leaves from late Paleozoic and Mesozoic plants,
generally of seed plants such as Lyginopteris but also of some ferns, in which the
leaves produced spores; global distribution, Devonian to Late Cretaceous.
Sphenosuchia: an order of basal crocodylomorph reptiles; between 1 and 2 feet long
and characterized by an upright posture and long, slender legs, they were fast runners;
the monophyly of the order is in question; global distribution, Middle Triassic.
sphinctozoans: a polyphyletic group of hypercalcified sponges that grow as a
series of chambers, one on top of the next; they originated in the Cambrian and
flourished in the late Paleozoic and the early Mesozoic; absent from the fossil record
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since the Late Cretaceous, but a single surviving species, Vaceletia crypta, was dis-
covered in Australia in 1977; global distribution, Cambrian to Recent.
Spiclypeus shipporum: a species of chasmosaurine dinosaurs in the family
Ceratopsidae, found in Montana in 2005 and described in 2016; known from a par-
tial skull and bones of the spine, legs, and hips, it was about 15 feet long and weighed
up to 4 tons, a medium-sized ceratopsian; like other chasmosaurs, it had a very large
and highly ornamented neck frill; North America, Late Cretaceous, 76–75 Ma.
spicule: a pointed mineral structure that is part of an animal’s skeleton, as in some
sponges and brachiopods; most often of silica or calcium carbonate.
Spinosaurus aegyptiacus: the type species of a genus of very large carnivorous
dinosaurs characterized by sturdy 6-foot-long vertebral spines and by crocodile-like
teeth as long as 7 inches. It was possibly the largest known carnivore at 36–50 feet
long, with its weight centered on its neck. Study of various fossils, including a par-
tial skeleton from the Kem Kem beds of Morocco — described in 2013 — seems to
prove that Spinosaurus spent a large part of its life in North African rivers, preying
on the numerous large species of fish and turtles. It was discovered about 1913 by
Ernst Strömer, who may have suspected its riverine habit but could not prove it. See
also Bahariasaurus, Baryonyx, Carcharodontosaurus, Suchomimus, and
Strömer’s Riddle. North Africa, Late Cretaceous, 112–94 Ma.
Spiriferida: an order of articulate brachiopods in the class Rhynchonellata.
Strongly ribbed and either punctate or impunctate, spiriferides are characterized
externally by a long, straight hinge line that is the widest part of the body, so that the
animal appears to be winged. Their name refers to the laterally directed spiral bra-
chidia, which support the lophophore. The Spiriferida appeared in the Ordovician,
were still rare in the Silurian, widespread in the Devonian and Carboniferous, sur-
vived the end-Permian extinction, and persisted to the Early Jurassic.
Spiriferidina: the larger of the two suborders of brachiopods in the order
Spiriferida (the other is Delthyridina, early Silurian to Permian); characterized by
strong ribbing; global distribution, Late Ordovician to late Permian.
Spiriferina walcotti: a species of articulate brachiopods in the order Spiriferinida;
characterized by a broad, ribbed shell with a wide hinge line and folds at the edge;
from 1 to 2 inches wide with faint concentric growth lines; Europe, Early Jurassic.
Spiriferinida: an order of later articulate brachiopods in the class Rhynchonellata,
similar to Spiriferida but generally smooth-shelled; global distribution, Early
Devonian to Early Jurassic.
Spondylus: a still-extant, large, and diverse genus of spinose bivalve mollusks in
the family Spondylidae, and in the same superfamily as scallops, the Pectinoidea;
they cement themselves to rocks and are often called spiny oysters, although unre-
lated; up to 4 inches wide, they have several eyes around the edges of the shell;
global distribution, Middle Triassic to Recent.
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sponges: see Porifera.

Spongiomorphidae: the single family of colonial hydrozoans in the extinct order
Spongiomorphida; their massive colonies are characterized by longitudinal rods
perpendicular to the growth surface and connected by horizontal bars parallel to the
surface; global distribution, Late Permian to Eocene.
Spriggina: an enigmatic genus of soft-bodied organisms known definitively only
from the Ediacara Hills; from 1 to 2 inches long, it is segmented, with a small head;
it superficially resembles an arthropod, but several characters seem to mark it as un-
arthropod; it also resembles sea pens and some rangeomorphs; Australia, 560 Ma.
Squalodontidae: a possibly polyphyletic family of large toothed whales in the
superfamily Platanistoidea; although poorly understood, they seem to be intermediate
between the Archaeoceti and the Odontoceti; the type genus Squalodon is the only
unquestioned member of the family; global distribution, Oligocene to Pleistocene.
Squaloraja: a genus of chimaerid fish in the family Squalorajidae that seems to be
transitional between sharks and rays; known from a specimen collected by Mary
Anning and one from Italy; Europe, Early Jurassic.
Squamata: the order of reptiles that contains the snakes (Serpentes), lizards
(Lacertilia), and worm lizards (Amphisbaenia); the second-largest order of verte-
brates, after the perciform fish, its sister group is the Rhynchocephalia; global dis-
tribution, Early Jurassic to Recent.
Staffellidae: a family of benthic, semi-infaunal foraminiferans in the superfamily
Fusulinoidea, related to the Ozawainellidae; the test is generally circular; global
distribution, Late Carboniferous to late Permian.
stage: see Geologic Time Scale.
Stagonolepis: a genus of herbivorous aetosaurian reptiles in the family
Stagonolepididae; quadrupedal and slow-moving, it was about 10 feet long and
well-armored, with osteoderms covering most of its body; Stagonolepis had no
teeth at the front of its mouth but cheek teeth adapted for chewing vegetation;
Europe and possibly North and South America, Late Triassic, 237–227 Ma.
Staphylinidae: the rove beetles, the largest family of beetles, in the order
Polyphaga; staphylinids are carnivorous or omnivorous scavengers; probably the
oldest is Leehermania prorova, found in the Solite Quarry of Virginia and described
in 2012; global distribution, Late Triassic to Recent.
Staurikosaurus: a genus of early theropod dinosaurs in the family Herrerasauridae;
dated at 225 Ma, it is one of the earliest-known dinosaurs; it was 7 feet long and
weighed about 66 pounds; Brazil, Late Triassic.
Stauropteridae: a group of enigmatic Carboniferous plants usually placed in the
order Cladoxylopsida and considered ancestral ferns; poorly known but character-
ized by forked, spore-bearing branches, similar to the ancestral Cooksonia of the
Silurian; cosmopolitan, Late Devonian to Late Carboniferous.
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stegocephalians: a paraphyletic group of choanatan lobe-fins, formerly considered

rhipidistian fish but now identified as tetrapods (such as Tiktaalik and Ventastega),
which are transitional forms leading to the first terrestrial animals; cosmopolitan,
Middle to Late Devonian.
Stegodontidae: a family of proboscideans in the extinct suborder Mammutoidea,
considered by some to be a subfamily in the Elephantidae; they had low-crowned
teeth with thick enamel but like modern elephants had lost their lower tusks; Africa
and Asia, middle Miocene to late Pleistocene.
Stegosauria: “plate lizard,” a suborder of herbivorous ornithischian dinosaurs;
they were well-armored, with a row of plates along the spine and scutes elsewhere;
the head was small and narrow, and they had long hindlimbs but were quadrupedal;
global distribution, Jurassic to Cretaceous.
Stegosauridae: a family of large ornithischian dinosaurs in the suborder
Stegosauria; this family was larger than the earlier stegosaurs, the type genus
Stegosaurus weighing more than 5 tons; North America, Europe, Asia, and Africa;
Middle Jurassic to Early Cretaceous.
Steinheim skull: a fairly well-preserved hominin skull without the lower jaw,
found at Steinheim, Germany; it has characteristics of Neanderthals and of Homo
heidelbergensis, and is estimated to be between 250,000 and 350,000 years old.
stem group: a paraphyletic grouping of organisms that consists only of extinct
species, excluding all members of the crown group.
Steneosaurus: a genus of thalattosuchians in the family Teleosauridae; the species
in the genus provide an example of niche partitioning, being divided into two
groups, one with short, broad jaws, and the other with long, very narrow jaws;
especially good specimens of the latter species are known from the Holzmaden
Shale; Europe and Madagascar, Early Jurassic to Early Cretaceous.
stenohaline: not tolerant of a wide range of salinity; trilobites and brachiopods, for
example, are stenohaline and less able to tolerate changes in salinity than bivalves,
which are generally more euryhaline.
Stenolaemata: the oldest class of marine bryozoans. Characterized by sac-shaped,
tubular, or conical exoskeletons, stenolaemates were the dominant bryozoans
throughout the Paleozoic, contributing to reefs in many places. One order, the
Cyclostomatida, is still extant; the Cystoporata and Cryptostomata became extinct
at the end of the Permian and the Trepostomata in the Triassic. See also
Gymnolaemata. Global distribution, Early Ordovician to Recent.
Stenophlebiidae: a family of damsel-dragonflies in the order Odonata; medium-
sized to large, distinguished by their wing venation pattern and long, slender wings;
a 2015 discovery in northwestern China represents the earliest specimen of the fam-
ily; Europe, Asia, and South America; Early Jurassic to Early Cretaceous.
Stenopterygius: “narrow wing,” a genus of thunnosaurian ichthyosaurs known
from well-preserved specimens from Holzmaden and other Lower Jurassic sites;
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up to 12 feet long, with a long, narrow beak and long, thin pectoral fins; a famous
Holzmaden specimen is a mother and fetus which died before childbirth, with the
fetus in a position to be born tail-first; Europe, Early Jurassic.
Stensioellida: a monospecific order of primitive fish, generally considered the
most primitive placoderms. The order consists of one species, Stensioella heintzi,
known only from the Hunsrück Slate. It has a whip-like tail and large pectoral fins,
and its elongate body has no large bony plates but is armored by a covering of small
dermal tubercles. Germany, Early Devonian.
Stenurida: a small order of primitive asterozoan echinoderms in the class
Ophiuroidea; this group of brittle stars includes widely disparate families and may
be polyphyletic; cosmopolitan, Early Ordovician to Late Devonian.
Stephanodiscus yellowstonensis: a species of diatoms in the order Thalassiosirales
with a unique history. Recent research shows that it evolved within a short span of
time in the Yellowstone area of Wyoming, between 12,000 and 8000 years ago. Once
found in many lakes of the area, it is now restricted to Yellowstone Lake. It has not
evolved further in the last 8000 years and now seems to be becoming less abundant.
stereom: the sponge-like network of calcium carbonate crystals that composes the
body tissue of all echinoderms.
Stereospondyli: a large suborder of late temnospondyl reptiles, the largest about
7 feet long; characterized by a broad, flat head, they were among the last of the
temnospondyls; known only from Gondwana up to the end of the Permian, they
radiated worldwide in the Triassic; late Permian to Early Cretaceous.
Steropodon: one of several extinct genera of monotremes in the family
Steropodontidae; closely related to the modern platypus (Ornithorhynchus
anatinus), it had teeth and grew as long as 2 feet, large for a Mesozoic mammal;
New South Wales, Australia, Early Cretaceous, 110–105 Ma.
Stethacanthus: “chest spine,” a genus of Devonian sharks in the order Symmoriida,
notable for a mysterious structure on a male’s back in place of the first dorsal fin; it
is a flattened area covered with dermal denticles, looking like a narrow, toothed
anvil, and its purpose is unclear; North America and Europe, Late Devonian to Early
Carboniferous.
Sthenurus: “strong tail,” a genus of kangaroos in the family Macropodidae and
extinct subfamily Sthenurinae; short-faced and with a tail shorter but stronger than
modern kangaroos, they grew as long as 10 feet and weighed more than 500 pounds;
Australia, Pliocene to Pleistocene, becoming extinct later than 19,000 years ago.
Stigmaria: a form genus of the rooting structures of lycopsid trees such as
Lepidodendron and Sigillaria; global distribution, Carboniferous.
stipe: the branch of a graptolite, to which the individual thecae are attached; also,
the stem of algae connecting the holdfast to the upper parts.
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Stolonoidea: a group of encrusting or sessile animals generally considered grapto-

lites, but possibly belonging to the pterobranch family Rhabdopleuridae; global dis-
tribution, Ordovician.
stratigraphy: the descriptive study of rock strata and interpretation of geological
history, including deposition, relative age, and correlation between outcrops that
may be separated by great distances; see biostratigraphy.
Strawberry Bank: a Lower Jurassic Lagerstätte in Somerset, England. It is one of
four Lagerstätten dated near the boundary of the Pliensbachian and Toarcian Stages
of the Early Jurassic, along with the well-known Holzmaden deposits in Germany,
the Osteno deposits of Italy, and the recently explored Ya Ha Tinda site in Canada.
Strawberry Bank is especially notable for its nearshore deposits containing 20 spe-
cies of insects as well as its specimens of nektic organisms (ichthyosaurs, fish, and
cephalopods). It differs from other sites also in having predominantly three-
dimensional specimens inside calcareous nodules. England, Early Jurassic, 183 Ma.
Strepsirrhini: a suborder of primates that includes the lemuroids, galagos, and
lorises, and their extinct adapiform primate relatives; formerly called Prosimii; dis-
tinguished from haplorrhines by the moist area around their nostrils; see also
Haplorrhini; widespread in Laurasia from the Eocene to the Miocene but now
found only in tropical Asia and Africa; early Eocene to Recent.
Strepsodiscus: one of the earliest mollusks, a genus of primitive snail-like mol-
lusks generally considered bellerophontid gastropods; two species are known, with
an asymmetrical shell about an inch high; see also Chippewaella; North and South
America and Antarctica, late Cambrian to Ordovician.
Streptognathodus isolatus: a species of conodonts in the family Polygnathidae; its
first appearance, in Kazakhstan, marks the beginning of the Permian at 298.9 Ma;
several other species of Streptognathodus are important for stages of the Late
Carboniferous and early Permian.
Streptospondylus: a genus of medium-sized theropod dinosaurs in the family
Megalosauridae; one of the first dinosaurs to be discovered, its fossils were found in
the late eighteenth century in France and described variously for several decades
until finally recognized as dinosaurs; France, Middle Jurassic.
stridulation: see Orthoptera.
strobilus: a cone; a common form genus of many cone-bearing plants.
stromatolites: layered, often bulbous, accretionary structures formed by micro-
organisms in a process of microbial growth, decomposition, and lithification. They
accumulate grains of sediment of many kinds in bio-chemical films called laminae,
whose morphology varies according to water depth and wave action. Some as much
as 3.5 billion years old, they occur in varied shapes from conical to horizontal, often
forming reefs. Global distribution, Archaean to Recent.
Stromatoporoidea: a group of primitive reef-building organisms now generally
considered poriferans, either demosponges or sclerosponges, but their precise affin-
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ities are still in dispute; see also Sphaeractinoidea; cosmopolitan, Precambrian to

Cretaceous, but especially important reef-builders in the Silurian and Devonian.
Strömer’s Riddle: the question why so many large predatory dinosaurs inhabited
the riverine environments of North Africa in the middle of the Cretaceous although
relatively few herbivores (such as Aegyptosaurus) were present in the fossil record.
Ernst Strömer, of the University of Munich, raised the question in the course of his
extensive excavations in North Africa in the early twentieth century; most of his
specimens were lost in a bombing raid in Munich in 1944. Some research between
2010 and 2014 indicates that the large carnivores were supported by a large popula-
tion of gigantic aquatic animals, such as a variety of very large turtles, 8-foot lung-
fish, 13-foot coelacanths, and 25-foot sawfish. If further research confirms that
Spinosaurus and other large predators did in fact spend most of their time in the
water, the riddle may have been solved.
strophic: of brachiopods, having a well-defined and elongated hinge line.
Strophomenida: the largest and most diverse brachiopod order, articulate bra-
chiopods in the class Strophomenata; the type genus Strophomena appeared in
the Ordovician but became abundant only in the Triassic; the order is generally
characterized by one convex valve and the other flat or concave; they are usually
wider than they are long, with a strophic hinge line; but see also the aberrant horn-
shaped Prorichthofenia and conical Richthofenia; global distribution, Ordovician
to Early Jurassic.
Strunius: a genus of unusual lobe-finned fish in the order Onychodontida; only 4
inches long, Strunius had the articulated skull characteristic of the rhipidistians and
the coelacanths; its fins were not so muscular as those of other lobe-fins, being stiff-
ened by bony rays like the fins of actinopterygians; Europe, Late Devonian.
Struthiomimus: “ostrich mimic,” a genus of medium-sized saurischian dinosaurs
in the family Ornithomimidae; they were bipedal and bird-like, with a small, slender
head and long neck; the type species, S. altus, was about 14 feet long and weighed
about 330 pounds; Canada, Late Cretaceous, 78–68 Ma.
Stylasterina: an order of delicate, lacy cnidarians in the class Hydrozoa; like the
milleporines, they are colonies of soft-bodied polyps but different from other
Hydrozoa in having calcareous parts that fossilize; New Zealand, Asia, Australia,
and North America; Cretaceous to Recent.
Stylemys: a genus of small terrestrial tortoises in the family Testudinidae; the
earliest-known land turtle in North America, up to 3 feet long; forelimbs too weak
for burrowing; North America, Europe, and Asia; late Eocene to Miocene.
Stylemys nebrascensis: a species of North American tortoise in the family
Testudinidae; about 2 feet long, with an unusually high-domed shell; Nebraska, late
Eocene to early Oligocene.
Styracosaurus: a genus of North American ornithischian dinosaurs in the family
Ceratopsidae; characterized by large armor plates on the head, a robust nasal horn,
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and a large neck frill crested with long spikes; rhinoceros-sized, about 18 feet long
and weighing more than 3 tons; Alberta and Montana, Late Cretaceous, 76–74 Ma.
Subulitidae: a family of marine siphonate gastropods in the clade Caenogastropoda
and superfamily Subulitoidea; from 1 to 2 inches long, generally awl-shaped; global
distribution, Ordovician to Carboniferous.
Suchomimus: a monospecific genus of primitive, crocodile-like dinosaurs in the fam-
ily Spinosauridae and subfamily Baryonychinae, closely related to Baryonyx. A strik-
ing example of convergent evolution, S. tenerensis was bipedal and had a long tail and
strong forearms. Its long, flattened skull was very similar to that of a crocodile and was
well-adapted to catching fish. Suchomimus had 120 cone-shaped teeth and was 40 feet
long and 10–12 feet high at the hip. North Africa, Early Cretaceous, 121–113 Ma.
sulcus: a shallow groove or furrow on the surface of some part of an organism’s
body; in brachiopods, a characteristic furrow in the pedicle valve, usually corre-
sponding to a fold or ridge in the brachial valve; in vertebrates, usually referring to
a fold separating parts of the brain.
Sundance Formation: a mostly sandstone Middle Jurassic formation in Western
North America; composed of mainly marine deposits laid down in the inland water-
way called the Sundance Sea, it contains few but important fossils of the period.
Supersaurus: a genus of very large sauropodomorph dinosaurs in the family
Diplodocidae; on the basis of specimens from Colorado and Wyoming, which
account for about 35 percent of the skeleton, its length is estimated at 110–112 feet
and its weight at up to 40 tons; see also Sauropoda; United States and Portugal,
Late Jurassic, 152–150 Ma.
suture line: of ammonoids, the pattern of lobes and saddles on the edge of a sep-
tum, often used in classification of genera.
Swanscombe Man: three skull fragments found at Swanscombe, England, ana-
tomically similar to Homo sapiens with a cranial capacity of around 1300 cc; dated
at about 450,000 years ago.
Sycidiales: an extinct order of charophytic algae in the class Charophyceae and
division Charophyta; notable for its uncommon gyrogonites, which have porous
vertical walls, as opposed to the characteristically spiral walls of later charophytes;
the oldest known species, Sycidium siluricum, is considered the oldest known bisex-
ual plant; global distribution, Silurian to Early Carboniferous.
Sylvian sulcus: the lateral sulcus of the human brain, a shallow furrow between
three lobes and the most prominent human sulcus; deeper in most primate brains, it
is a deep cleft in the most primitive monkeys.
Symbos cavifrons: a species of North American woodland musk ox, an artiodactyl
ruminant mammal in the family Bovidae; it is known from the early Pleistocene and
was one of many large ruminants whose extinction at the end of the Pleistocene was
probably human-related.
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symmetrodonts: formerly considered an order of primitive mammals, but the term

is now used in an informal sense to refer broadly to a large number of early mam-
mals close to the line of descent from cynodonts to mammals; the name refers to the
symmetrical arrangement of the upper and lower molar teeth; see also
Kuehniotherium and Morganucodon; North America, Europe, and Asia; Late
Triassic to Late Cretaceous.
Symphysurus: a genus of asaphid trilobites in the family Nileidae; medium-sized,
about 3 inches long; global distribution, Early to Late Ordovician.
synapomorphy: an apomorphy shared by a group of lineages to the exclusion of
other groups, a key to inferring evolutionary relationships by identifying common
ancestors.
Synapsida: the advanced amniotes formerly called mammal-like reptiles. They are
a sister group of reptiles but are now considered part of the clade Reptiliomorpha.
Synapsids are identified primarily by a lateral temporal fenestra (a single opening in
the skull behind each eye). The Synapsida include the pelycosaurs (Permian) and
the more advanced therapsids (chiefly Triassic, descendants of pelycosaurs and
ancestors of mammals). Middle Pennsylvanian to Recent .
synonyms: two or more different names referring to the same taxon.
Syntrophiidina: one of two suborders of impunctate articulate brachiopods in the
extinct order Pentamerida; more primitive than the second suborder, the
Pentameridina; characterized by an almost circular shell with a pronounced dorsal
fold; global distribution, early Cambrian to Early Devonian.
syntypes: a set of several specimens that serve together as name-bearers for a spe-
cies if no holotype has been designated.
Synziphosurina: a paraphyletic group of chelicerate arthropods in the still-extant
class Xiphosura, related to the modern Limulus; characterized by free abdominal
segments; cosmopolitan, Ordovician to Devonian.
Syringophyllidae: a family of tabulate corals in the extinct order Sarcinulida, with
a horizontal and tubulate structure; global distribution, Ordovician and Silurian.
Syringopora: a genus of tabulate corals in the order Auloporida and family
Syringoporidae. Syringopora is phaceloid, with a sub-colonial structure of long and
vertical cylindrical corallites, interconnected irregularly by horizontal tubuli, so
that there was no contact in life between the polyps living at the top of the column
of corallites. As the colony grew, it expanded horizontally. Because of this struc-
ture, Syringopora is often called organ-pipe coral. Global distribution, Ordovician
to Permian.
Syringothyris: a genus of articulate brachiopods in the superfamily Spiriferinida
and family Syringothyrididae; characterized by an internal tube around the pedicle
valve, a punctate shell, and a pyramidal shape; about 2 inches across and semi-
infaunal in lime substrates; global distribution, Silurian to Permian.
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tabula: (plural tabulae) a transverse septum between the walls of tabulate corals
and some archaeocyathans.
Tabulata: one of the two dominant Paleozoic corals, an order of wholly colonial
cnidarians in the subclass Hexacorallia. The Tabulata are distinguished by tabulae
within each of the corallites, which are almost always hexagonal. Appearing along
with the rugose corals and stromatoporoids at the beginning of the Ordovician, they
are characteristic of Silurian and Devonian shallow seas. Less common after the
Devonian, they became extinct at the end of the Permian. Global distribution, Early
Ordovician to Permian.
Tachypleus: one of two surviving genera of horseshoe crabs, chelicerates in the
order Xiphosurida and family Limulidae; see also Limulus; China and Southeast
Asia, Late Jurassic to Recent.
Taeniodonta: a suborder of non-placental eutherian mammals in the order
Cimolesta; the taeniodonts varied greatly in size, from rat-sized to bear-sized
(Stylinodon, of the Eocene); some became highly specialized for digging, but efforts
to categorize them on that basis are still tentative because of a sparse fossil record;
Western North America, Late Cretaceous to Eocene.
Taeniolabidoidea: a suborder of primitive mammals in the order Multituberculata;
although mostly small and rodent-like, some were the largest of the multituberculates;
Taeniolabis taoensis weighed more than 200 pounds, similar to Castoroides, the largest
beaver of the Pleistocene; North America and Asia, Late Cretaceous to early Eocene.
Taeniopteris: a genus of Mesozoic tree-like plants in the Pentoxylales with leaves
as long as 16 inches; global distribution, Carboniferous to Late Cretaceous.
Taimyr wolf: According to a DNA analysis in 2015, a 35,000-year-old fossil of a
wolf on the Taimyr Peninsula is a previously unknown species that indicates a date
between 27,000 and 40,000 years ago for the splitting off of dogs from wolves,
earlier than previous estimates; Siberia.

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Tanystropheus: a genus of large archosauromorph reptiles in the extinct order

Protorosauria and family Tanystropheidae; notable for a very long neck (10 feet) com-
posed of only 13 elongated vertebrae, while its body and tail together were only about
10 feet long; probably a semi-aquatic piscivore; Europe and Asia, Middle Triassic.
Tanytrachelos: a genus of small amphibious reptiles in the extinct order
Protorosauria and family Tanystropheidae; the type species T. ahynis is a long-necked
archosauromorph only 1–2 feet long; United States, Late Triassic, 225–220 Ma.
Tapejaridae: a family of pterodactyloid pterosaurs, probably related to the
Azhdarchidae; characterized by a large bony crest and better vision than most ptero-
saurs; many possessed a distinctive rounded crest, sometimes as wide as 3 feet;
global distribution, Early to Late Cretaceous.
taphocoenosis: a taphonomic association of fossils, either autochthonous or
allochthonous. The term is sometimes used to refer specifically to an allochthonous
assemblage of fossils transported after their death and deposited together, so that the
assemblage does not represent a functional community in life. The term thanatocoe-
nosis, on the other hand, specifically refers to an autochthonous death assemblage
of organisms that died together.
taphonomic artifact: a product of artificial character resulting from some acci-
dent during the process of fossilization and thus not indicative of the actual struc-
tural relationships of the parts of the organism.
taphonomy: the study of an organism’s history between its death and the time of
its discovery. Three taphonomic stages are recognized: necrosis (death), biostratin-
omy, and diagenesis. Biostratinomy refers to what happens to the organism between
necrosis and burial, while diagenesis refers to what happens to it after burial, prin-
cipally the process of mineralization. Taphonomy is especially concerned with
transported allochthonous deposits and with biases present in deposition and fos-
silization, such as scavenger activity, different rates of disintegration and decompo-
sition, displacement of the fossil by soil movement, and dissolution or mineral
replacement of bones.
Tapiroidea: a still-extant superfamily of perissodactyl mammals in the suborder
Ceratomorpha. Probably related to the early Eocene Homogalax and Hyracotherium,
the tapiroids spread throughout the northern continents and diversified slightly in
the middle Eocene into four short-lived families: the Isectolophidae in North
America, the Lophiodontidae in Europe, and the Deperetellidae and Lophialetidae
in Asia. Spreading globally in the Oligocene with little evolutionary change, the
tapiroids became extinct in the northern land masses, persisting in some places to
about 10,000 years ago. Several genera are still extant elsewhere. See also
Cambaytherium and Heptodon.
Tapirus californicus: a species of tapir known from New Mexico to Southern
California in the early and middle Pleistocene; one specimen is known from the Oregon
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coast, a well-preserved jaw with teeth; it is either T. californicus or the better-known T.

haysii, which is common in lower Pleistocene deposits in Eastern North America.
Tapocyon: a genus of primitive mammals in the family Miacidae; coyote-sized, it
had retractable claws and was partly arboreal; Southwestern United States, Eocene.
Tarbosaurus bataar: a species of large carnivorous dinosaurs in the subfamily
Tyrannosaurinae. A close Asian relative of Tyrannosaurus rex, it was 33–39 feet
long and weighed 5 tons or more. Two elongated eggs discovered in 1993, purport-
edly of this species, are at 16.5 inches long the largest dinosaur eggs known;
Northern China, Late Cretaceous.
Tardigrada: The “water bears,” a still-extant minor phylum of almost microscopic
animals, called tardigrades because of their slow, lumbering gait. There are about
400 modern species, most of them living in water films on lichens and other plants,
and some living on the bottom of ponds and ocean basins. They have eight legs, most
of which are only 0.1 to 0.5 mm long. They are notable for their ability to survive
long periods of cryptobiosis, complete dormancy; a large group in a European Space
Agency experiment survived in a vacuum for a week in orbit around the Earth. No
fossils were known until their discovery in the 1990s in Cambrian deposits in Siberia,
so that the group does seem to have existed as long as most other modern phyla.
Tarsiidae: a still-extant family of haplorrhine primates in the infraorder
Tarsiiformes; tarsiers, now restricted to Southeast Asia, arose in the middle Eocene
and became widespread in North America, Europe, and Asia, with two disputed
African species from the Eocene and Oligocene (see Afrotarsius).
Tasmanian tiger: see Thylacinus.
Taxideinae: the American badgers, a subfamily of caniform omnivores in the fam-
ily Mustelidae; North America, Miocene to Recent.
Taxocrinida: an order of crinoid echinoderms in the extinct subclass Flexibilia;
characterized by a small, conical cup and crown; global distribution, Middle
Ordovician to Late Carboniferous.
taxon: a group of organisms sharing unique features that differentiate the group
from other taxa.
taxonomy: the study of the nomenclature and classification of organisms in groups
according to their descent and evolutionary relationships.
Teilhardina: a genus of marmoset-like haplorrhine primates tentatively placed in
the family Omomyidae, but probably polyphyletic and currently being reassessed;
North America, Europe, and Asia; early to middle Eocene.
Teilhardina magnoliana: the earliest-known primate in North America; mouse-
sized, weighing about an ounce, it resembled a small marmoset; discovered in 2008
and estimated to have lived at the boundary of the Paleocene and Eocene; United
States.
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Teleoceras: a genus of perissodactyl mammals in the family Rhinoceratidae;

standing 9 feet high and weighing 1 ton, it resembled the hippopotamus but bore a
small nasal horn; North America and France, early Miocene to early Pliocene.
Teleosauridae: a family of aquatic crocodylomorphs in the clade Thalattosuchia;
generally characterized by elongate snouts and forelimbs only half as long as the
hindlimbs, they resembled the modern gharial; they ranged in length from 6 to more
than 16 feet, with some estimates much higher; North and South America, Europe,
Africa, and possibly China; Early Jurassic to Early Cretaceous.
Teleostei: an infraclass of actinopterygian fishes in the superclass Osteichthyes.
The teleosts comprise 50 percent of all known vertebrate species and 96 percent of
all extant species of fish. They are characterized by movable upper-jaw bones,
which allow the jaw to protrude forward; fully homocercal (symmetrical) tail fins;
and thin, bony scales. Most teleosts reproduce by external fertilization of eggs, with
generally no paternal care; a few genera are viviparous and somewhat altricial.
Early Triassic to Recent.
Telephinidae: a family of nektic trilobites in the order Proetida; see also
Opipeuterella; global distribution, Ordovician.
telson: the terminal section of the body of many arthropods.
Temnospondyli: “cut vertebra,” a large, diverse, and long-lived order of semi-
aquatic and marine tetrapods, generally considered primitive amphibians. They
include the first vertebrates to fully adapt to terrestrial life. Ranging in size from
very large (Mastodonsaurus) to very small, they flourished in the late Paleozoic and
the Triassic, and some genera survived into the Cretaceous. Whether they are ances-
tral to modern amphibians is unclear. (See Amphibamus.) Global distribution,
Carboniferous to Cretaceous.
Tempskya: a genus of Cretaceous ferns, the only genus in its family, Tempskyaceae;
it grew to about 15 feet and appeared tree-like, but its false trunk consisted of thin
rhizomatous stems intertwined with small adventitious roots; its leaves grew directly
from the upper parts of the trunk, not on branches or in a crown; global distribution
but primarily Northern Hemisphere, Early to Late Cretaceous.
Tentaculitida: an enigmatic order of mollusk-like lophophorates, incertae sedis;
less than an inch long but abundant in some localities, their ribbed calcitic shell is
straight and conical; global distribution, Early Ordovician to Early Carboniferous.
Teratophoneus curriei: a species of carnivorous tyrannosaurid dinosaurs known
only from a partial skull and post-cranial skeleton found in 2010 in the Kaiparowits
Formation of Southern Utah; about 12 feet tall and 20 feet long; Late Cretaceous,
about 76–74 Ma.
Teratornithidae: a family of vulture-like birds that includes the largest known fly-
ing birds, known principally from the Americas. The North American genera (at
least four species in three genera identified so far) on average were about 40 percent
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larger than the modern condor. Teratornis merriami, the best-known teratorn
because of many specimens recovered from the La Brea Tar Pits, had a wingspan of
12 feet but weighed only 30–35 pounds. The largest known of the family is
Argentavis magnificens, known from several partial specimens found in South
America; it had a wingspan of 20–26 feet and may have weighed up to 170 pounds.
(For the largest wingspan of a flying bird, see Pelagornis sandersi.) Predators and
scavengers, the last of the teratorns became extinct about 10,000 years ago. North
and South America, Miocene to late Pleistocene.
Terebratellidina: a still-extant small suborder of articulate brachiopods in the
class Rhynchonellata and order Terebratulida; characterized by a long loop, a punc-
tate shell, and a median septum; global distribution, Triassic to Recent.
Terebratulidina: a still-extant suborder of large articulate brachiopods in the class
Rhynchonellata and order Terebratulida; characterized by a short loop, internal
spicules, and a punctate and generally biconvex, egg-shaped shell; the widespread
modern genus Terebratula grows as long as 4 inches; global distribution, Late
Devonian to Recent.
Teredinidae: the shipworms, a family of marine bivalve mollusks; they are worm-
like except that they use two small shells at the anterior end of the body to bore
through wood, living inside their tunnel; known worldwide, they range from 2
inches to 3 feet long and are very destructive; they live worldwide but are less active
in brackish water, such as the Baltic Sea; Early Jurassic to Recent.
Teredo: the most common genus of shipworms, marine bivalve mollusks in the
family Teredinidae; global distribution, Late Jurassic to Recent.
Ternifine man: fossil bones found in Algeria in 1954 in a stratum about
700,000 years old, originally named Atlanthropus mauritanicus but later recognized
as an example of Homo erectus.
terrane: a large fragment of continental crust differing from the surrounding
crustal material because it originated a considerable distance away and was accreted
to the continent by tectonic forces.
Tertiary: a discarded term formerly used to refer to the first part of the Cenozoic,
from the end-Cretaceous extinction to the end of the Pliocene.
Tessarolax: a genus of gastropods in the still-extant family Aporrhaidae; character-
ized by thin, curving spines 1–3 inches long, growing from the edge of the lip;
North America, Europe, Africa, and Madagascar; Cretaceous to Paleocene.
test: the external skeletal covering of echinoderms, composed of calcium carbon-
ate; the internal walls and external covering in foraminiferans, composed of either
calcium carbonate or agglutinated grains of sediment.
Testudines: the turtles, a still-extant order of mainly aquatic reptiles, characterized
by a hard dorsal carapace above the rib cage and a hard ventral plastron attached to
the rib cage. Whether their origin was terrestrial or marine is still debated. The order
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was once called Chelonii, and the term chelonian refers to turtles in general. The
earliest turtles are usually placed in a slightly different order or clade; see for exam-
ple Archelon, Desmatochelys, Odontochelys and Proganochelys. See also
Cryptodira and Pleurodira. Global distribution, Late Triassic to Recent.
Testudinidae: the modern land tortoises, a family of cryptodire tortoises in the
still-extant order Testudines; global distribution, Cretaceous to Recent.
tetanurans: the large group of dinosaurs (including tyrannosaurids, ornithomim-
ids, and others) that are more closely related to modern birds than to the
ceratosaurs.
Tethys Sea: a chiefly tropical body of water to the east of Pangaea from the late
Permian to the Miocene, with Laurasia to the north, Gondwana to the south, and the
land masses that became China and Southeast Asia to the east.
Tethytheria: a clade of ungulate mammals containing the proboscideans, the sire-
nians, and the extinct desmostylians.
Tetraceratops: an obscure genus of small synapsids represented by a single 4-inch
skull found in Texas in 1908; despite its four-horned name, T. insignis has six small
horns on its face; incertae sedis, it appears to be a transitional genus midway
between pelycosaurs and therapsids, the precursors of mammals; North America,
early Permian.
Tetragraptus: an early genus of dichograptid graptolites in the order Graptoloidea;
planktic, with four stipes, pendent or horizontal, and no bithecae; important index
fossil because of its many widely distributed species in the Early Ordovician.
Tetralophodon: a genus of elephant-like proboscideans in the extinct family
Gomphotheriidae; about 8 feet high, with four tusks and a trunk, but named for its
specialized four-cusped teeth; North America, Europe, and Asia; middle Miocene to
early Pliocene.
Tetrapoda: a superclass of vertebrate animals that includes many living and
extinct taxa. The earliest tetrapods appeared in the fossil record in the Devonian
period, at about 370 Ma. Various environmental pressures — including intense
competition among the many marine animals, the growing presence of terrestrial
plants along the shorelines, and climatic changes — led some marine organisms to
be attracted to habitats along shallow shorelines, evolving into species that could
adapt to terrestrial habitats. Anatomical changes in some Middle Devonian rhipi-
distian fish such as Eusthenopteron gave rise to lobe-finned sarcopterygians such
as Tiktaalik, some of which evolved in the Late Devonian into an amphibian life-
style. Especially notable are the bones of the lobe fins, which correspond closely
to the arm, wrist, and hand bones of later vertebrates. These bones are a homolo-
gous trait in all vertebrates descended from the lobefin fish, including humans and
even snakes (considered tetrapods because they are descended from diapsids,
which had four limbs like other vertebrates). The only modern sarcopterygians are
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the coelacanths and lungfish. See also Acanthostega, Ichthyostega, Koilops,

Latimeria, Perittodus, Romer’s Gap, Tiktaalik, Tulerpeton, and Whatcheeridae.
Tetrarhynchia: a genus of articulate brachiopods in the still-extant order
Rhynchonellida; characterized by coarse ribs, incurved beak, and a width greater
than the length; Europe, Asia, and North America; Jurassic.
Texas Red Beds: a series of lower-Permian formations in Texas and Oklahoma,
one of the most prolific sites for a range of reptiles, sharks, stem-tetrapods, temno-
spondyls, and other amphibians; North America, early Permian.
Thalassocnus: a genus of xenarthran sloths in the family Megatheriidae. It is rep-
resented by several species living off the coasts of Peru and Chile; herbivorous and
about the size of a human, it is one of the few tetrapods that have returned to the sea
and then become extinct (see also Neovison). The earlier species were probably
semi-aquatic and the later species fully aquatic. South America, late Miocene to late
Pliocene.
Thalattosauria: “ocean lizard,” an order of primitive marine reptiles, difficult to
classify but generally considered archosauromorph diapsids; growing as long as
13 feet, they had a broad body, flattened tail, and paddle-shaped limbs; North
America, Europe, and Asia; Middle to Late Triassic.
Thalattosuchia: “sea crocodile,” a clade of marine crocodylomorphs consisting of
the Metriorhynchidae and the Teleosauridae; cosmopolitan, Early Jurassic to Early
Cretaceous.
thanatocoenosis: see taphocoenosis.
Thaumaptilon: an enigmatic organism of the Burgess Shale, possibly a cnidarian;
it resembles a sea pen and may be the earliest pennatulacean, although some
Ediacaran organisms may be even earlier pennatulaceans; Canada, middle Cambrian,
508 Ma.
thecae: cases, receptacles, or sheaths in various animals such as graptolites and
echinoderms; especially the chitinous structures protruding from a graptolite’s
stipes, often resembling the teeth of a saw blade, which are the tubular skeletons of
the individual zooids of a rhabdosome, a graptolite colony.
Thecideidina: a still-extant small suborder of brachiopods in the class
Rhynchonellata, exact relationships unclear; very small, cemented to the undersides
of corals or in coral debris, they are the only extant tropical brachiopods; cosmo-
politan, Triassic to Recent.
thecodonts: “socket tooth,” a paraphyletic group of basal archosaurs that appeared
in the Permian and diversified widely in the Triassic; formerly classified as an order,
the group includes the ancestors of dinosaurs, crocodilians, and pterosaurs; similar
in appearance to a more upright crocodile or to a four-legged theropod dinosaur,
they became abruptly extinct at the end of the Triassic; global distribution, Permian
to Triassic.
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Thecosmilia: a genus of stony corals, colonial scleractinian corals in the s till-extant

suborder Faviina; the corallites are phacelo-dendroid and about half an inch in
diameter; an important reef-builder, especially in the Jurassic; global distribution,
Middle Triassic to Cretaceous.
Thectardis avalonensis: a species of triangular fossils known only from the
Mistaken Point biota; it is incertae sedis but has been suggested as the earliest
sponge; probably cone-shaped in life, it grew to 2.5 inches wide and 7 inches high;
Canada, Ediacaran, 575–560 Ma.
Thelodonti: a contentious group of jawless fish still considered a class by some
but probably polyphyletic; 6–8 inches long, they resembled the heterostracans and
anaspids but are characterized especially by their thelodont scales, tiny, oval dermal
denticles with ridges, furrows, and spikes; global distribution, Middle Ordovician to
Late Devonian.
Thelodus: an obscure genus of thelodont agnathan fish in the family Coelolepididae;
characterized by a broad, blunt head and thick, bony, pear-shaped scales covering its
body; estimates of its size vary from a few inches to 3 feet long; global distribution,
Silurian.
Theosodon: a genus of litoptern mammals in the family Macraucheniidae; similar
in appearance to the modern guanaco, they were almost 7 feet long and weighed
about 200 pounds; South America, early Miocene.
Therapsida: a group of synapsids that includes the dominant predators of the
Permian, becoming a transitional link between their reptile ancestors and the early
mammals of the Triassic, which probably evolved from a therapsid group called
cynodonts. The earliest therapsid may be the still-controversial Tetraceratops of the
early Permian, and other basal therapsids like the Biarmosuchia appeared in the
middle Permian. Developing from the sphenacodontid pelycosaurs, therapsids
became the dominant predators, replacing the pelycosaurs in the middle Permian
and diversifying globally in the late Permian and Early Triassic. Only a few genera,
including the cynodonts, persisted into the Jurassic, as the dinosaurs gradually
replaced most of the therapsids.
Theria: the principal subclass of mammals, including the Metatheria (marsupials)
and Eutheria (placentals) but excluding the Prototheria (egg-laying monotremes);
the earliest-known therians (see Juramaia and Montanalestes) appeared in the
Middle Jurassic, but there is some evidence of their existence in the Early Jurassic.
Theridomyoidea: a superfamily of rodents possibly related to the sciuromorphs;
Europe and Africa, Eocene to Miocene.
Theriodontia: an important group of advanced carnivorous therapsids, including
the therocephalians and gorgonopsians in the late Permian, and then in the Early
Triassic the more advanced cynodonts, from which the ancestors of mammals
evolved; cosmopolitan, middle Permian to Recent.
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Theriognathus: a genus of therapsids in the family Whaitsiidae; about 3 feet long

with a long, narrow snout, it exhibits mixed reptile and mammal characteristics; a
close relative may have survived the end-Permian extinction; Africa and Russia, late
Permian.
Therizinosauridae: a large family of feathered herbivorous theropod dinosaurs,
related to the sharp-clawed, carnivorous raptors; species in the group varied in
length from 3 to 33 feet and weighed up to 5 tons; they were basically bipedal, with
claws as long as 3 feet on their robust forelimbs; numerous eggs have been found
that indicate little if any care by parents; also called segnosaurs; North America and
China, Early to Late Cretaceous.
Theropithecus: a still-extant genus of large monkeys in the family Cercopithecidae;
weighing from 25 to 40 pounds, three species are known from the Pliocene; East
Africa.
Theropoda: a large group of dinosaurs that spanned a great range of types and
sizes. Traditionally considered saurischians, in a proposed reclassification in 2017
they are considered a separate suborder. The theropods were generally carnivorous
and were the principal terrestrial carnivores of the Jurassic and Cretaceous. They
were among the earliest dinosaurs, dating back to 231 Ma, and their clade includes
all modern birds. The most important theropod families are the Allosauridae,
Compsognathidae, Dromaeosauridae, Spinosauridae, Therizinosauridae, and
Tyrannosauridae. Global distribution, Late Triassic to the end of the Cretaceous.
Thescelosaurus: “marvelous lizard,” a parrot-beaked genus of bipedal ornithis-
chian dinosaurs in the herbivorous family Thescelosauridae, usually classified as
ornithopods. About 12 feet long, it appeared in North America at the very end of the
Cretaceous, about 66 Ma. In a specimen found in 1993 the heart was preserved well
enough that computer tomographic scans seemed to reveal a four-chambered heart
and an aorta, significant because close to the evolutionarily advanced heart structure
of birds and mammals, and more efficient than the three-chambered heart of snakes
and crocodilians. A more thorough examination in 2011, however, disputed these
interpretations, indicating that the “heart” may be simply a concretion.
Thlipsuridae: a family of podocopid ostracods in the suborder Platycopina and
superfamily Thlipsuroidea; North America and Europe, early Silurian to Late
Devonian.
Thrinaxodon: “trident tooth,” a genus of early cynodont therapsids in the family
Thrinaxodontidae; a fox-sized carnivore, probably in the line of descent to mam-
mals; Thrinaxodon’s features are transitional between the reptilian traits of early
synapsids and the mammalian traits of advanced cynodonts; South Africa and
Antarctica, Early Triassic, 250–245 Ma.
Thylacinus cynocephalus: a species of carnivorous marsupials in the still-extant
order Dasyuromorphia and extinct family Thylacinidae. One of the “Tasmanian
tigers,” superficially dog-like but unrelated to the Canidae, it is a prime example of
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convergent evolution. It was the size of a large dog (6 feet long), with a long muzzle
and striking posterior dorsal stripes. The family appeared in the Oligocene and
became a top predator in Meganesia. T. cynocephalus is the only species to survive
into the twentieth century, evidently becoming extinct in the 1930s. Meganesia;
early Pliocene to Recent.
Thylacoleo: “pouch lion,” a genus of Australian marsupials in the still-extant order
Diprotodontia and extinct family Thylacoleonidae; resembling a small lion, it
weighed up to 350 pounds and was about 4 feet long; it was probably carnivorous,
but ambivalent dentition has led to intense debate about its diet and habits;
Thylacoleo became extinct between 45,000 and 40,000 years ago, shortly after the
arrival of humans in the continent; late Pliocene to late Pleistocene.
Thylacosmilus: a genus of scansorial carnivores in the order Sparassodonta; not a
felid, it was more closely related to marsupials than placentals; the leopard-sized
Thylacosmilus was similar by convergent evolution to the sabertooth cats of the
Oligocene and Miocene on other continents; Argentina, late Miocene.
Tianyulong: a monospecific genus of ornithischian dinosaurs in the family
Heterodontosauridae; the specimen of T. confuciusi was only about 28 inches long
but probably a juvenile; it was bipedal and had tufts of fuzz that seem to be primitive
feathers; China, Late Jurassic, 158 Ma.
Tiaojishan Formation: a series of interspersed volcanic and sedimentary rocks in
northeastern China, dated between 165 and 153 Ma; an important source of fossil
fauna and flora well-preserved by volcanic eruptions; especially notable for speci-
mens of the stem-mammaliaforms Juramaia, Castorocauda, Agilodocodon, and
Docofossor; China, Late Jurassic.
Tiktaalik roseae: a species of stegocephalians known from several skeletons found
on Ellesmere Island. The predatory Tiktaalik is a significant link between earlier
sarcopterygians and the first four-legged land animals. It was 6–8 feet long and pos-
sessed a suite of characters adapted to moving on land as well as in water, especially
its strong shoulders, bendable wrists and neck, and weight-bearing fins. It lived in
shallow coastal waters about 375 Ma, possibly making brief forays onto land. See
also Ventastega and Tetrapoda. Canada, Late Devonian.
Tillodontia: a suborder of eutherian mammals in the order Cimolesta, probably
related to the pantodonts; they were quadrupedal, and the largest was about 3 feet
long; they combine ungulate, rodent, and carnivore features, such as gnawing teeth
and clawed feet; North America, Europe, and Asia; early Paleocene to late Eocene.
Timurlengia: a genus of early tyrannosaurs intermediate between the first tyran-
nosaurs and the larger ones of the Late Cretaceous; discovered in 2004 and described
in 2016, T. euotica was still only horse-sized but was similar in several ways to its
large cousin Tyrannosaurus rex, notably in having similar brain and ear features; it
is dated at 90 Ma in the early Late Cretaceous, in the middle of a 20-million-year
gap in tyrannosaurid fossils; Uzbekistan.
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Titanis walleri: a North American genus of large, flightless phorusrhacid birds in

the order Cariamiformes; like the other Phorusrhacidae, it probably attacked smaller
animals from ambush; known from fragments but no skull, it is estimated to have
been 8 feet tall and up to 350 pounds; Florida, early Pliocene to early Pleistocene.
Titanites: a genus of ammonoids in the order Ammonitida and family
Dorsoplanitidae; strongly ribbed, it is one of the larger ammonoids, the largest spe-
cies more than 4 feet in diameter; England, Late Jurassic (end Tithonian).
Titanoides: a genus of herbivorous mammals in the extinct order Pantodonta;
about 3 feet long and 350 pounds, these were unusual herbivores in having large
canine teeth and five-clawed feet, although otherwise resembling a small modern
rhinoceros; North America, late Paleocene.
Titanophoneus: a genus of large carnivorous therapsids in the order Dinocephalia
and family Anteosauridae; the type species, T. potens, was as much as 10 feet long,
with the typical long, heavy snout and interlocking incisors typical of dinocepha-
lians; Russia, middle Permian.
Titanoptera: an order of neopteran insects in the subclass Pterygota; large, some
species having wingspans up to 16 inches; although related to grasshoppers, they
could not jump; Europe, Early to Late Triassic.
Titanosauria: a clade of large, herbivorous saurischian dinosaurs in the suborder
Sauropodomorpha. They had a long tail, a fairly long neck, spoon-like or peg-like
teeth, and solid vertebrae. The largest specimen, found in 2012 in Patagonia, was
among the largest of all dinosaurs at 130 feet long and about 90 tons, and several
juvenile specimens found there appear to be longer than 120 feet. A very large colo-
nial nesting ground has been found in Patagonia, consisting of hundreds of titano-
saur nests containing an average of 25 eggs each; other similar nesting grounds have
been found in Madagascar, India, and Spain. Global distribution but principally in
southern continents, Early to Late Cretaceous.
Titanotheriidae: see Brontotheriidae.
Titanotylopus: a genus of even-toed ungulates in the family Camelidae; the largest
camelid, about 13 feet high at the shoulder; North America, Miocene to Pleistocene.
Tomarctus: a genus of canids in the extinct subfamily Borophaginae; 3 feet long
and about 40 pounds, it had very powerful jaw muscles, indicating bone-crushing
ability, a hallmark of scavengers; probably close to the ancestry of modern dogs;
North America, early to middle Miocene.
Tommotian biota: an assemblage of invertebrates of the early Cambrian, named
for the area where they were first discovered in Siberia but probably occurring
worldwide; there are a few entire skeletons, but the assemblage consists mainly of
fragments known collectively as a “small, shelly fauna,” q.v.; 535–520 Ma.
Tongtianlong limosus: a species of maniraptoran oviraptorosaurs, one of
many similar recent Chinese discoveries; the specimen named in 2016 died in a
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s played-limb and raised neck posture; feathered and bipedal, it has a sharp beak and
a dome-like skull roof with a low crest; China, Late Cretaceous, 71–66 Ma.
toothplate: a specialized biting or crushing structure in the occlusal areas of the
mouth of several types of fish, including placoderms and several genera of rays,
lungfishes, and chimaeroids; even some rhynchosaurian reptiles have a type of
toothplate.
Torosaurus: a genus of chasmosaurid ornithischian dinosaurs in the family
Ceratopsidae; very similar to Triceratops, Torosaurus is characterized by a skull
9 feet long and by two large openings in its 5-foot-wide neck frill which give it its
name; it had two large horns projecting forward above its eyes and other smaller
horns behind its frill; the herbivorous Torosaurus weighed about 5 tons and was
about 30 feet long; North America, Late Cretaceous, 68–66 Ma.
Torrejonia: a genus of plesiadapiform primates generally considered in the family
Paromomyid but proposed as a member of a new family Palaecthonidae; one of the
oldest known primates, Torrejonia had a body length of 15 inches; it was arboreal,
with long legs, fingers, and toes; see also Plesiadapiformes; North America,
Paleocene.
Torvosaurus: a genus of large carnivorous saurischian dinosaurs in the family
Megalosauridae. Found in Utah, Colorado, Portugal, and possibly China, the genus
appeared at the time that North America and Europe were drifting apart. The largest
species, T. gurneyi, weighed almost 5 tons and was about 33 feet long; it is the larg-
est theropod known from Europe. A nest of eggs found in Portugal contains the
most primitive dinosaur embryos known. United States, Europe, and Asia; Late
Jurassic, 153–148 Ma.
Toxasteridae: a family of sea urchins, irregular euechinoids in the order
Spatangoida; they were atelostomate and had an elongated oval shape; North
America, Europe, and Asia; Early to Late Cretaceous.
Toxodon platensis: a South American ungulate in the suborder Toxodonta, the type
species of the family Toxodontidae; 8 feet long and 5 feet high, it was similar to the
modern rhinoceros; southern South America, Pliocene to Pleistocene.
Toxodonta: a suborder of mostly South American ungulates in the order
Notoungulata; the toxodonts were between 4 and 10 feet long, the larger genera
being similar to a modern rhinoceros in appearance and size; some species have
nasal openings on the top of the skull; research in 2014 suggests that the toxodonts
may be related to the perissodactyls (odd-toed ungulates); South and North America,
late Paleocene to Pleistocene.
Toxodontidae: a family of large notoungulates in the suborder Toxodonta; princi-
pally South American but known also from the Pleistocene in Central America and
the Southwestern United States; Oligocene to Pleistocene.
trace fossil: see paleoichnology and fossil.
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Trachodon: a dubious genus of dinosaurs, known only from teeth and dental
f ragments, some of which are now known to be from a ceratopsid and some from a
hadrosaur; see also Edmontosaurus; North America, Late Cretaceous, 77 Ma.
Tragulidae: a still-extant family of primitive hornless artiodactyls in the suborder
Ruminantia; commonly called chevrotains or mouse deer, they appeared in the
Eocene, descended from the Hypertragulidae; currently represented by three genera
in Asia and Africa; Europe, Asia, and Africa, late Eocene to Recent.
transitional species: a species that bears traits clearly indicating the sequential
phases in the evolution of a new species, specifically traits common to both its
ancestors and its descendants; see for example Amphistium, flatfish, Heteronectes,
Tetraceratops, and Theropoda.
Traversodontidae: a family of sheep-sized herbivorous therapsids in the suborder
Cynodontia; the traversodonts appeared in the Early Triassic, diversified widely
throughout the period, and became extinct in the Early Jurassic; mainly Gondwana
but also Laurasia.
Tremarctinae: a subfamily of carnivorous short-faced bears, still extant only in
one species, Tremarctos ornatus, found in limited places in the Andes; South and
North America, late Miocene to Recent.
Trematosauroidea: a superfamily of labyrinthodont amphibians in the suborder
Stereospondyli; medium-sized fish-eaters, some almost completely marine; one of
only two or three groups of temnospondyls that survived the Triassic extinction;
global distribution, Early Triassic to Middle Jurassic.
Trepostomata: an order of colonial bryozoans in the still-extant class Stenolaemata;
characterized by long, curving, impunctate, calcareous tubes, filled with closely
spaced diaphragms; tree-like colonies, seldom preserved intact, the fragmented
branches forming a major part of some Ordovician limestones; global distribution,
Ordovician to Triassic.
Treptichnus pedum: a trace fossil that marks the GSSP for the beginning of the
Cambrian Period and the Paleozoic Era; one of the earliest complex trace fossils, it
is the burrow of an animal similar to modern priapulid worms; Newfoundland,
Canada, 541 Ma.
Triadobatrachus: a genus of frog-like amphibians in the order Proanura and fam-
ily Triadobatrachidae; the earliest-known frog, Triadobatrachus was 4 inches long
and had 14 trunk vertebrae, a transitional feature since the typical amphibian had 24
and modern frogs only 4–9; it also had frog-like pelvic characteristics; Madagascar,
Early Triassic.
Triarthrus: “three-jointed,” a genus of ptychopariid trilobites in the family
Olenidae, it was the last of its family. One of the few trilobites found with intact
appendages, it was about 2 inches long and characterized by jointed, inward-facing
spines on the legs and a smoothly rounded pygidium. Good examples are known
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from Beecher’s Trilobite Bed in New York, especially notable for the spine details
and for information about the protaspis and meraspis phases. Researchers in 2017
also reported the first known direct evidence of trilobite reproduction in a cluster of
very small pyritized eggs near the head of a Triarthrus specimen from Beecher’s.
North America, Europe, and Asia; Late Ordovician.
Triassaraneus: a genus of true spiders (Araneae), the first found in South Africa;
known from several specimens of varying completeness; South Africa, Late Triassic
(Carnian), 235 Ma.
Triassic: the first period of the Mesozoic Era, from 252.2 to 201.3 Ma, divided into
Early, Middle, and Late Epochs. It was overall a relatively hot and dry period, domi-
nated by a great diversity of reptiles on land, especially the pseudosuchian archo-
saurs. Mammaliaforms and dinosaurs appeared in the middle of the period, and the
pterosaurs — which were the first flying vertebrates — at the end of the Middle
Triassic. Sauropterygians and ichthyosaurs dominated the seas, and scleractinian
corals appeared in the Middle Triassic, becoming important reef-builders late in the
period. Only one lineage of ammonoids survived the end-Permian extinction, but
from it the ceratitid ammonoids diversified rapidly and became very numerous
throughout the Triassic. Evolving by the end of the Triassic were also the ammon-
itids, which were to become the dominant ammonoid group of the Jurassic and
Cretaceous. The great coal swamps of the late Paleozoic almost disappeared during
the dry Triassic, but conifers, cycads, and corystosperms (see Dicroidium) flour-
ished throughout the period.
Triassic extinction: the fourth of the five great extinctions, at the end of the
Triassic Period. It affected land and sea, causing the extinction of one class of
marine animals, the conodonts; one-third of all marine genera; and many families of
terrestrial animals. One-third of all known animal species of the Triassic did not
survive, creating vacant ecological niches that the dinosaurs could occupy in the
Jurassic. However, plant fossils and palynomorphs show no appreciable effects. As
with all five mass extinctions, the causes are unclear; paleoclimatological research
has found a marked increase in the level of carbon dioxide in the atmosphere near
the end of the period, and other possible causes have been suggested. Some research-
ers have argued that at least two extinction pulses occurred. But the second, between
201.5 and 201.3 Ma at the end of the Rhaetian Stage, is the one usually referred to
as the Triassic extinction.
Triassochelys: see Proganochelys.
Tribrachidium heraldicum: a unique species of soft-bodied benthic organisms in
the monospecific phylum Trilobozoa, given its own phylum since it could not be
classified in any known phylum; it was limpet-shaped, circular and about an inch in
diameter, but its upper surface was a shallow cone with 3 lobes in a triskelion pat-
tern; triradially symmetrical and known only from negative impressions in overly-
ing strata; Australia and Russia, Ediacaran, 558–555 Ma.
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Triceratops: “three-horned face,” a genus of ornithischian dinosaurs in the family

Ceratopsidae. The largest weighed as much as 12 tons and were almost 30 feet long
and 10 feet tall. They were characterized by a very large, 10-foot skull with a parrot-
like beak; a large, bony neck frill; and three horns. Herbivorous and eating a highly
fibrous diet, they had replaceable grinding teeth. Known only from the Late
Cretaceous of North America, 68–66 Ma.
Triconodonta: see Eutriconodonta.
Trigoniidae: a still-extant family of articulate bivalve mollusks in the order
Trigoniida. They are large and roughly triangular, with striking patterns of ribbing
and a very complex articulation with elaborate hinge teeth. Although trigoniids
were widespread in the Jurassic and Cretaceous, they were thought to have become
extinct at the end of the Cretaceous until a living member of the family (now named
Neotrigonia) was discovered in 1802 in Australia. Global distribution, Late
Carboniferous to Pliocene.
Trigonostylopidae: a family of South American hoofed mammals in the order
Astrapotheria, q.v.
Trigonotarbida: a large group of primitive arthropods in the order Arachnida,
superficially resembling spiders. They were probably predators on other arthropods,
and some Devonian genera developed armor, spines, and tubercles. They did not
have spinnerets and probably did not produce venom. Trigonotarbids are well-
known, especially from the European Coal Measures of the Late Carboniferous.
Europe and North and South America, early Silurian to early Permian.
trilobites: a very large group of mostly small marine arthropods, generally referred
to as a class called Trilobita. The earliest definitive arthropods (see also Spriggina
and Parvancorina), the trilobites were characterized by a hard exoskeleton, a longi-
tudinally three-lobed body, and a series of biramous limbs. The body is also divided
in three parts from head (cephalon) to thorax to tail (pygidium), generally also hav-
ing a post-segmental part called the telson. They are sometimes found enrolled,
curled into a ball. Although trilobites averaged around an inch in length, a few spe-
cies grew as long as 28 inches (see Isotelus rex). Mostly benthic and sometimes
burrowing in the top layers of sediment, but some small species were evidently
planktic.
Trilobites arose in the early Cambrian and flourished in the late Cambrian and
the Ordovician. Since the hard exoskeleton fossilized relatively easily, they are
among the most abundant fossils of the Paleozoic. Their abundance and their world-
wide distribution make them stratigraphically important through the middle of the
Paleozoic; they dwindled in numbers thereafter up to their extinction at the end of
the Permian.
Trimerellida: a superfamily of inarticulate brachiopods, formerly called
Trimerellacea; they have a strong calcite shell and a pedicle, sometimes attached to
another brachiopod; global distribution, Middle Ordovician to late Silurian.
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Trimerocephalus: a genus of eyeless trilobites in the order Phacopida and family

Phacopidae; noted for a 2016 report of a single-file line of individuals in close physi-
cal contact with each other, evidently dying suddenly while migrating; Europe, Asia,
and Australia; restricted to the Famennian Stage of the Late Devonian, 371–365 Ma.
Trimerophytopsida: an extinct class of early vascular plants in the division
Tracheophyta; arising in the Early Devonian from the rhyniophytes, the trimero-
phytes are considered ancestral to the progymnosperms and ferns, and thus to
almost all higher plant life today; also called Psilophytopsida, Trimerophytales, and
Trimerophyta; global distribution, Devonian to Carboniferous.
Trimerus: a genus of phacopid trilobites in the family Homalonotidae and related
to the calymenids; characterized by a wide, shovel-shaped cephalon and small,
raised eyes; global distribution, Silurian to Devonian.
Trinil fauna: a biostratigraphic faunal assemblage at Trinil, Java, that contains the
hominin remains commonly referred to as “Java Man”; the fauna is dated at about
1.8 Ma; the Trinil site is about the same age as the Jetis fauna and considerably older
than the “Solo Man” remains at the Ngandong site; all three sites are within 60 miles
of each other.
Trinucleidae: a family of small, blind trilobites in the order Asaphida and super-
family Trinucleoidea, important as index fossils in the Ordovician; characterized by
an almost circular shape up to an inch across, long genal spines, a vaulted glabella,
and a ribbed fringe around the large cephalon, which accounts for about half of the
body length; the trinucleids went extinct in the Ashgill event, 444 Ma, at the end of
the Ordovician; the type genus is Trinucleus, but see also Cryptolithus; global dis-
tribution, Middle to Late Ordovician.
Trinucleoidea: a superfamily of trilobites in the order Asaphida; the only asaphids
to survive the Ordovician extinction into the Silurian; Ordovician to late Silurian.
Trionyx: a still-extant genus of softshell freshwater turtles in the order Cryptodira;
3 feet long, with a long, narrow head; global distribution, Early Cretaceous to Recent.
Triops cancriformis: a still-extant species of tadpole shrimps, notostracans in the
family Triopsidae; 2–4 inches long and resembling a small Limulus, it is known
from the Grès à Voltzia Lagerstätte and remains almost unchanged today; Europe,
Early Triassic to Recent.
Tritheledontidae: a family of advanced cynodonts in the infraorder Eucynodontia;
formerly called Ictidosauria; among the longest-lived non-mammalian therapsids,
they were probably replaced in the Jurassic by early mammals; mainly carnivorous
or insectivorous, the largest species were less than 10 inches long; South America
and Africa, Late Triassic to Middle Jurassic.
Tritylodontidae: a family of mammaliamorph therapsids in the suborder Cynodontia;
tritylodonts were generally herbivorous, characterized by protruding incisors and
three rows of grinding cheek teeth; they had a prominent sagittal crest and are esti-
mated to have been about 2 feet long; global distribution, Triassic to Cretaceous.
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Trochiliscales: an early order of charophytic algae characterized by dextrally

coiled gyrogonites; cosmopolitan, Silurian to Carboniferous.
Troodon: a genus of small, agile theropod dinosaurs in the family Troodontidae;
up to 8 feet long and 150 pounds, it is notable for evidence of bird-like nesting and
egg-laying behavior, including possible brooding behavior by males; China, Russia,
and North America, found recently even in Northern Alaska; Late Cretaceous,
78–76 Ma.
Trucherognathidae: “ragged jaw,” a family of conodonts characterized by fibrous,
platelike teeth resting in a depression on the jaw ramus rather than grasping the
ramus or the jaw tip; North America, Europe, and Asia; Ordovician.
Tryblidium: a large genus of monoplacophorans in the family Tryblidiidae; from 1
to 2 inches long with a pear-shaped shell, in some species composed of an outer
calcitic and an inner aragonitic layer; North America and Europe, Ordovician to
Silurian.
Tsidiiyazhi abini: a species of sparrow-sized birds in the still-extant family
Coliidae (the mousebirds), described in 2017 as the oldest known tree-dwelling
bird; found in New Mexico, it is dated at 62.5 Ma and is evidence of a rapid specia-
tion in birds following the end-Cretaceous extinction; United States, early Paleocene.
tuatara: see Sphenodontidae.
Tubuliporina: still-extant, the largest suborder of bryozoans in the order
Cyclostomatida; global distribution, Late Triassic to Recent.
Tulerpeton curtum: a species of early tetrapods in the order Ichthyostegalia,
closely related to Acanthostega and Ichthyostega; Tulerpeton lacked gills and had
six digits on each limb; one of the few well-known labyrinthodonts from the period
of time called Romer’s Gap; Russia, Late Devonian, 365–361 Ma.
Tullimonstrum gregarium: the famous “Tully monster,” a soft-bodied chordate in
the class Hyperoartia; possibly a stem lamprey, it is known only from the Mazon
Creek site southwest of Chicago and is the Illinois state fossil; 6–8 inches long, it
has stalked eyes and a notochord; United States, Late Pennsylvanian.
Tunicata: a large, still-extant subphylum of basal marine chordates, also called
Urochordata; tunicates have a notochord only in their free-swimming larval phase;
as adults they are benthic and sessile; the main group is the Ascidiacea, commonly
called sea squirts; global distribution, Cambrian to Recent.
Tuojiangosaurus: a monospecific genus of ornithischian dinosaurs in the family
Stegosauridae; similar to the better-known Stegosaurus, T. multispinus was 24 feet
long and weighed about 3 tons; it is notable for 17 pairs of spikes and plates along
its backbone; China, Late Jurassic, 161–155 Ma.
Turkana Boy: the name given to a nearly complete hominin skeleton found in
1984 near Lake Turkana, tentatively assigned to H. ergaster; the most complete
early hominin skeleton known; Kenya, 1.6 Ma.
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Turrilites: an unusual genus of ammonoids in the order Ammonitida, suborder

Ancyloceratina, and family Turrilitidae; characterized by its resemblance to gastro-
pods, a strongly ribbed surface, and a high-spired, tightly coiled helical shell; global
distribution, Late Cretaceous, 100–90 Ma.
turtles: see Testudines.
Tuzoia: a genus of bivalved organisms tentatively classified as stem-arthropods;
probably nektobenthic, it had a circular carapace from 6 to 8 inches in diameter and
a pair of large stalked eyes; North America, Europe, Asia, and Australia; early to
middle Cambrian.
Two Medicine Formation: an important geologic formation of the Campanian,
encompassing a large area of Northwestern Montana and Southern Alberta; laid
down along the western shore of the Western Interior Seaway between 80 and
74 Ma, the Two Medicine strata contain many fossils of Late Cretaceous dinosaurs
as well as many freshwater and terrestrial animals; North America, Late Cretaceous.
Tylocidaris: a genus of sea urchins, echinoids in the order Cidaroida and family
Psychocidaridae; notable for club-shaped spines, narrow at the proximal end and
bulbous at the distal end; Europe and North America; Early Cretaceous to late
Eocene.
Tylopoda: “swollen foot,” a still-extant suborder of artiodactyl ungulates, origi-
nating in North America in the middle Eocene and found only there until the early
Pliocene; although once placed in the Ruminantia, they are not true ruminants; the
modern representatives are the Camelidae, the camels and camelids of Asia and
South America.
Tylosaurus proriger: a large species of mosasaurs in the subfamily Tylosaurinae;
the type species of the genus, T. proriger was almost 50 feet long and weighed up to
11 tons; characterized by an elongated, cylindrical snout, it inhabited the shallow
Western Interior Seaway in the middle of North America in the Late Cretaceous,
with a diet including fish, shellfish, and small aquatic birds; although most
Tylosaurus specimens have been found in North America, they are known also from
Spain, New Zealand, and Antarctica; 85–70 Ma.
type species: as defined by the ICZN, the name-bearing type of a genus or
subgenus, considered the typical representative of a genus with which the name is
permanently taxonomically associated; a type genus has the same relationship to a
family.
type specimen: see holotype.
typostrophism: a saltationist theory of evolution espoused by the German paleon-
tologist Otto Schindewolf in the middle twentieth century; stressing catastrophi-
cally destructive events as the principal cause of mass extinctions and of the
subsequent emergence of new taxa, typostrophism downplays natural selection as
the basic mechanism of evolution. See also Unspecialized Descent.
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Typotheria: a possibly paraphyletic suborder of South American ungulates in the

extinct order Notoungulata, possibly including the hegetotheres; ranging from
rabbit-like to bear-sized, the typotheres shared several characters with rodents; the
Interatheriidae and Mesotheriidae are the largest of the five families in the suborder;
late Paleocene to middle Pleistocene.
Tyrannosauridae: a family of large predatory dinosaurs in the superfamily
Tyrannosauroidea; restricted to North America and Asia, Early to Late Cretaceous.
Tyrannosauroidea: a superfamily of theropod saurischian dinosaurs; arising in
the Middle Jurassic, the early tyrannosauroids, such as Guanlong and Eotyrannus,
were relatively small and had well-developed forelimbs with three fingers; the later
genera became the largest known land predators ever; global distribution,
165–66 Ma.
Tyrannosaurus rex: the type species of the family Tyrannosauridae, T. rex reached
lengths up to 45 feet and 12–15 feet high at the hips; estimates of its weight vary
from 7 to 18 tons. Like the more recently discovered Albertosaurus, a close relative,
T. rex was among the most powerful predators in the history of life. It was character-
ized by a large skull more than 3 feet long, with several dozen serrated teeth up to 6
inches long. It had broad skull attachments for its powerful jaw muscles, and very
large vertebrae that supported its massive head. Bipedal, with very strong hindlimbs,
it had relatively weak forelimbs only 3 feet long; each manus had two claws.
Although the teeth did not meet, coprolites indicate that T. rex may have chewed and
crushed even the bones of small prey by a shearing action of its teeth.
T. rex is noteworthy for an unusual pattern of holes in its skull that may indicate
a more advanced nervous system than most other dinosaurs, and it may have had
stereoscopic color vision, although nothing is known of its visual acuity. Its life-span
has been estimated at 27–30 years, and its normal walking speed was about 5 miles
an hour, but it probably reached speeds as high as 18 miles an hour. The first partial
skeletons of T. rex were found in Montana by Barnum Brown in 1900 and 1902, and
about a dozen partial skeletons have been found since in South Dakota, Wyoming,
Montana, and Alberta. One of the most complete skeletons, referred to as “Sue,” is
now in the Field Museum in Chicago. North America, Late Cretaceous, 68–66 Ma.
U
Ugrunaaluk: a monospecific genus of hadrosaurs in the subfamily Saurolophinae;

described in 2015, U. kuukpikensis was a duckbill dinosaur that lived above the Arctic
Circle; several thousand bones belonging to the species have been found on the
Colville River in Northern Alaska in strata from 69 Ma; it was 25–30 feet long and
7 feet high at the hip, and bore crests along its spine; it is so similar to Edmontosaurus
that some researchers consider them the same species; Late Cretaceous.
Uintacrinida: a small order of stemless crinoids that includes only two genera,
Uintacrinus and Marsupites; North America and Europe, Late Cretaceous,
87–84 Ma.
Uintatheriidae: a family of large, primitive herbivorous mammals in the order
Dinocerata; the uintatheres were replaced in their large-browser niche late in the
Eocene by the brontotheres; North America and China, late Paleocene to late Eocene.
Uintatherium: the type genus of the mammal family Uintatheriidae; up to 13 feet
long and weighing as much as 2 tons, Uintatherium resembled the modern rhino
ceros in shape but was not related; its skull was very thick and bore 6 frontal ossi-
cones; United States and China, middle to late Eocene.
Umoonasaurus: a monospecific genus of Early Cretaceous plesiosaurs in the fam-
ily Leptocleididae; U. demoscyllus was identified by Australian researchers in 2006
along with the larger plesiosaur Opallionectes; it was about 8 feet long and had
three crest-like ridges on the skull; it swam in a shallow sea of Central Australia
about 115 Ma.
ungulates: a general term for the many herbivorous hoofed mammals of the
Cenozoic; Late Cretaceous to Recent.
uniformitarianism: Until the eighteenth century, the accepted view was that life on
Earth has been affected throughout its history by sudden catastrophic events — such
as great floods, widespread volcanic eruptions, and rapid formation of mountain
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which no longer occur. In the period after such catastrophes, new and c ompletely
different life forms were thought to have appeared to repopulate the devastated
regions. As knowledge of the fossil record increased in the eighteenth and nineteenth
centuries, this view became untenable and the theory of uniformitarianism gained
credibility, advanced especially by James Hutton and Charles Lyell. Uniformitarianism
stressed the idea that changes in the past were gradual and were caused by essentially
the same forces of change observable in the present. From about 1850 to 1970 this
view was almost universally accepted, with gradualism considered a hallmark of
evolutionary change. In the last few decades a more complex view has emerged,
accepting the importance of occasional catastrophic events that cause relatively rapid
changes in the conditions of life. See also punctuated equilibrium.
Uniramia: “one branch,” a term sometimes used for a grouping of arthropods
including myriapods, onychophorans, and hexapods (insects).
Unspecialized Descent, Law of: Also called Law of the Unspecialized, it was
formulated by E.D. Cope. It holds that the typical taxa of any temporal unit are not
closely related to the most highly developed and specialized taxa of the previous
epoch but are rather descended from the least specialized taxa. Otto Schindewolf
made it an important part of his anti-Darwinian theory of typostrophism, summariz-
ing it thus: “Evolution builds upon that which has remained simple.” Although the
concept contains an obvious general truth, its use in typostrophism has been widely
refuted as over-simplified.
Uptonia jamesoni: a species of ammonoids in the order Ammonitida and family
Polymorphitidae, an important index fossil in the Pliensbachian; evolute, strongly
ribbed, about 4 inches wide; Europe and Greenland, Early Jurassic.
Uraraneida: an order of arachnids in the subclass Tetrapulmonata. As its name
indicates, the order contains the earliest araneids. Still poorly known, the uraraneids
seem to be proto-spiders, but they may lack spinnerets, and they retain a segmented
abdomen and a flagellum resembling that of scorpions. Uraraneida was established
for Attercopus fimbriunguis, and in 2016 Permarachne was assigned to the order.
North America and Russia, Middle Devonian to Permian.
Urochordata: see Tunicata.
Ursidae: the bears, a still-extant family of mammals in the clade Caniformia and
superfamily Arctoidea; the ursids evolved from canids such as Cephalogale in the
Miocene and are now found on all continents but Antarctica.
Ursus spelaeus: the cave bear, a species of omnivorous mammals in the family
Ursidae; about the size of the largest modern bears and differing only in a few char-
acters, it became extinct between 24,000 and 10,000 years ago; Europe, Pleistocene.
Utahraptor: a genus of theropod dinosaurs in the subfamily Dromaeosaurinae,
known from several specimens from Eastern Utah; estimated at up to 23 feet long
and half a ton, it is probably the largest known dromaeosaur, and it seems to be of a
heavier build than its cousins Deinonychus and Velociraptor; like other dromaeo-
saurs, it hunted in packs; United States, Early Cretaceous, 130–125 Ma.
V
Vaceletia: a still-extant monospecific genus of sphinctozoan sponges; the

s phinctozoans were thought to be extinct since the Cretaceous until V. crypta was
discovered off the northeastern coast of Australia in 1977; important because of a
rich Mesozoic fossil record, it may be related to the ceractinomorph demosponges;
global distribution, Cambrian to Recent.
Vagaceratops: a monospecific genus of herbivorous horned dinosaurs in the sub-
family Chasmosaurinae; found in Alberta’s Dinosaur Provincial Park, V. irvinensis
was given its “wandering” appellation because of its similarity to Kosmoceratops,
known only from southern Utah; the similarities extend to the neck frill, square in
Vagaceratops but topped the same way by several hornlets, some curving forward;
it was 15–20 feet long and had no other horns besides those on the frill; Canada,
Late Cretaceous, 75 Ma.
Valenictus chulavistensis: an extinct species of walrus in the still-extant family
Odobenidae and subfamily Odobeninae. Its tusks are only slightly shorter than
those of the modern walrus Odobenus, and it is very similar to Odobenus except that
it has no teeth besides the tusks. Valenictus is one of at least 24 species of walrus
that evolved from bear-like land animals since the late Oligocene, with only one
species still surviving. See also Dusignathus; Southern California, Pleistocene,
3 Ma.
Varanops: a genus of primitive synapsid reptiles in the family Varanopseidae;
about 4 feet long, it is a larger version of its relative Varanosaurus; global distribu-
tion, early Permian.
Varanosaurus: a genus of pelycosaurian synapsid reptiles in the family
Ophiacodontidae; from 3 to 4 feet long, with a set of sharp teeth, including two pairs
of longer teeth at the tip of its long snout, adapted to grasp prey; North America,
early Permian.

302 V
Varanus: a genus of very large monitor lizards in the family Varanidae; synonym:
Megalania. Ancestral to the modern Komodo dragon, it was about 20 feet long and
weighed about a ton. Australia, Asia, Europe, and Africa; Miocene to Recent.
Variscan orogeny: the mountain-building process, caused by tectonic shifting,
associated with the convergence of Laurasia and Gondwana in the Devonian and
Carboniferous to form Pangaea; also called Hercynian orogeny; see
paleogeography.
Vaughaniidae: a family of tabulate corals in the superfamily Favositicae; charac-
terized by shallow, rounded corallites and the absence of tabulae; type species
Vaughania cleistoporoides Garwood; Europe, Early Carboniferous.
Vegavis iaai: a genus of duck-like birds in the suborder Anatoidea, known from
only one specimen but generally considered the first anseriform bird; about 1 foot
long, V. iaai is the oldest known waterfowl; CT scans indicate that it had a voice box
and could probably vocalize as modern birds do; found on Vega Island in Antarctica
and named for the Instituto Antártico Argentino, which described the first of two
specimens in 2005; Late Cretaceous, 68–66 Ma.
Velociraptor: a Mongolian genus of turkey-sized carnivorous dinosaurs in the fam-
ily Dromaeosauridae. About 6 feet long but weighing less than 35 pounds, it was a
bipedal feathered predator, characterized by a large, forward-pointing claw on the
top of each pes. This sickle claw may have been used for attack, as was probably
true of Utahraptor and Achillobator, larger dromaeosaurs up to 20 feet long.
Disagreement continues as to whether the claw was sharp enough to rip skin open
or may have been used in tree-climbing to a perch to pounce on prey. Mongolia,
Vendian: see Ediacaran.
Ventastega curonica: a species of early stegocephalian tetrapods; from about
365 Ma, it is similar to Tiktaalik, another early tetrapod that was terrestrial as well
as aquatic; about 3 feet long, relatively large for the time, it is one of the earliest
tetrapods yet found; Latvia, Late Devonian.
Ventriculites: a genus of cup- or vase-shaped sponges in the class Hexactinellida;
characterized by siliceous spicules fused into a rigid skeleton; North America and
Europe, Jurassic to Eocene.
Verbeekinidae: a family of highly specialized foraminiferans in the superfamily
Fusulinoidea; characterized by planispirally coiled tests; global distribution, early
to late Permian.
Vernanimalcula: a monospecific genus of microscopic acritarchs that display
some bilateral symmetry; V. guizhouena, found in the Ediacaran Doushantuo
Formation of China, was dated in 2004 at between 600 and 580 Ma, 40 million
years before the earliest undisputed bilaterians; it was first described as a bilaterian,
V 303
but the claim has been widely disputed, so that even the kingdom in which
Vernanimalcula should be placed is in doubt.
Vertebrata: the subphylum of chordate animals that have a vertebral column con-
taining the spinal cord, including jawless, bony, and cartilaginous fish and jawed
vertebrates. There are about 66,000 species of vertebrates, accounting for 5 percent
of all described animals. All vertebrates have an internal skeleton, a central nervous
system running dorsally the length of the body, and gills or gill arches. (In mammals
the jaw, the thyroid, the larynx, and the ossicles of the ear all correspond to the gill
arches of earlier vertebrates.) For the earliest vertebrates, see Myllokunmingia,
Haikouichthys, and Zhongjianichthys.
vertisol: a clay soil that is subject to extreme shrinking and expanding with envi-
ronmental changes; taphonomically important in evaluating the original position of
fossils deposited in it.
vestigial structures: body parts of organisms that seem to have no function, such
as hip bones of snakes and rudimentary hindlimbs of whales. Such structures are to
be expected in the process of natural selection and are among the more obvious
evidences of evolution. See also analogous and homologous structures.
Veterupristisaurus: a genus of theropod dinosaurs in the family
Carcharodontisauridae; described in 2011 as the oldest carcharodontosaurid and
estimated to have been about 30 feet long, it is known from spinal fragments found
in an Upper Jurassic formation dated at around 150 Ma; Tanzania, Late Jurassic.
Vetulicolia: an enigmatic group of early to middle Cambrian organisms, tenta-
tively considered deuterostomes. Their size ranges from microscopic to 4 inches
long; they are generally elongate, with a larger anterior section which contains sev-
eral gill-like openings, and a posterior tail-like section. They have no eyes or
appendages but do seem to have a notochord and are considered chordates.
Cosmopolitan, middle Cambrian.
Vilevolodon diplomyos: a species of early haramyidan mammals in the family
Eleutherodontidae; about 3 inches long, it seems to have been a glider similar to the
larger Maiopatagium; its complex tooth crown dentition is similar to that of modern
squirrels; China, late Jurassic, 160 Ma.
Vintana sertichi: a species of groundhog-like mammals in the family
Sudamericidae, discovered in Upper Cretaceous strata on the western coast of
Madagascar in 2010. The almost intact skull increases what is known of the
Gondwanatheria, early mammals known previously only from teeth and fragments
of bone from the Late Cretaceous through the Miocene. Dated between 72 and
66 Ma and weighing about 20 pounds, Vintana was much larger than other known
mammals of the time and is notable for well-developed olfactory bulbs in the brain.
Madagascar, Late Cretaceous.
Viverravidae: a family of early mammals in the extinct superfamily Miacoidea;
related to early carnivores but believed to be unrelated to any modern carnivorans;
304 V
characterized by an elongated skull and by having only two molars in the upper and
lower jaws; North America, Europe, Asia, and Africa; early Paleocene to late
Eocene.
Viverridae: a still-extant family of omnivorous small mammals in the order
Carnivora and suborder Feliformia; although primitive, they have several living rep-
resentatives, such as civets, linsangs, binturongs, and in some classifications mon-
gooses; global distribution, late Eocene to Recent.
Volaticotherium: a monospecific genus of insectivorous eutriconodont mammals
in the proposed subfamily Alticonodontinae. The earliest-known gliding mammal,
V. antiquum was about the size of a squirrel but is not related to the modern flying
squirrel. Mongolia, Middle Jurassic, 164 Ma.
Volborthella: a problematic genus of early Cambrian fossils; its fragmentary sili-
ceous specimens are widespread in North America, Greenland, and Europe in lower
Cambrian strata, appearing earlier than trilobites and then concurrently with them
through the early Cambrian; once suggested as a cephalopod but now simply incer-
tae sedis; see also Salterella.
Volchovia: a genus of ophiocistioid echinoderms in the family Volchoviidae; its
turtle-like shell was high in the middle and flattened out toward the edges, like an
inverted shallow vase; Russia, Ordovician to Silurian.
Voltzia: a genus of early conifers, abundant in the Triassic Grès à Voltzia Lagerstätte,
to which it gave its name; generally low-growing and bushy; Europe, North and
South America, Europe, and Asia; Carboniferous to Triassic.
Vombatidae: a still-extant family of Australian marsupials in the order
Diprotodontia; the family diverged from other diprotodonts earlier than the late
Oligocene and possibly in the late Eocene; several genera of large Vombatidae
became extinct in the late Pleistocene; see Phascolonus.
Voulte-sur-Rhône: see La Voulte-sur-Rhône.
W
Waimanu: a genus of sphenisciform flightless diving birds of the Paleocene, prob-

ably a basal penguin; New Zealand, middle Paleocene.
Walcott, Charles D.: American paleontologist who discovered and investigated
the Burgess Shale site, q.v.
Wallace’s Line: the imaginary line between the different fauna characteristic of
Southeast Asia on one hand and of islands centered on Australia on the other. The
line, named for Alfred Russel Wallace, runs between parts of Indonesia, specifically
through the Lombok Strait between Bali and Lombok, eastward between Borneo
and Sulawesi, and then south of the Philippines. The deep-water barrier represented
by the line is thought to have existed for 50 million years, since the early Eocene.
With few exceptions, both mammals and birds have remained separate on the two
sides, even though the distance is quite small in places, especially in periods of great
ice formation in the polar regions.
walruses: see Valenictus.
Waptia: a genus of shrimp-like arthropods known primarily from W. fieldensis of
the Burgess Shale; an earlier species is known also from Chengjiang; about 3 inches
long, it had a thin exoskeleton and long antennae; China and North America, early
to middle Cambrian.
Weichselia: a genus of ferns in the still-extant order Gleicheniales and family
Matoniaceae; characterized by lobed leaves on 3-foot-long fronds, it spread rhi-
zomatously, forming dense swards over large areas; global distribution, Jurassic to
Late Cretaceous.
Wenlock Series: a Konservat-Lagerstätte of the middle Silurian, a site in Western
England that contains an important biota caught in fine-grained deposits of volcanic
ash interwoven with successive strata of carbonate mud; organisms represented
include polychaete worms, sponges, graptolites, microscopic radiolarians, and other
uncommonly fossilized organisms; Wenlock Epoch of the Silurian.

306 W
Western Interior Seaway: a shallow sea covering much of Central North America
in the Cretaceous; in its largest phase it was 600 miles wide and stretched from the
Arctic Ocean to the Gulf of Mexico, and was also connected by the Hudson Seaway
across Eastern Canada to the Atlantic Ocean; also called Niobrara Sea and
Cretaceous Seaway; Early Cretaceous to early Paleocene.
Whatcheeridae: a small family of temnospondyl amphibians of the Early
Carboniferous; named for What Cheer, Iowa, where a large deposit of early amphib-
ians was found in the 1990s; about 3 feet long, the type species Whatcheeria deltae
is dated to 340 Ma, about the same date as Greererpeton; the family also includes
Pederpes finneyi; North America, Early Carboniferous.
Wheeler Shale: a Konzentrat-Lagerstätte in the House Range of Western Utah,
dated at 507 Ma. Known principally for large numbers of trilobites (see Elrathia),
it has also produced specimens of Anomalocaris, Wiwaxia, and other early organ-
isms such as bivalved crustacean arthropods, chancelloriids, vetulicolians, priapulid
worms (see Selkirkia). The fauna is considered comparable to the coeval Burgess
Shale biota. North America, middle Cambrian.
White River Group: a large set of lower Oligocene deposits centered on south-
western South Dakota; the volcaniclastic strata have produced many articulated
skeletons of mammals from the period of transition from subtropical forests to
grassland, including entelodonts, oreodonts (Merycoidodon), all three of the rhino
cerotoids (see Amynodontidae, Hyracodontidae, and Rhinocerotidae), horses,
hyaenodonts, canids, and some of the last of the brontotheres; United States, late
Eocene to early Oligocene.
Wielandiella: a genus of shrubs in the family Williamsoniaceae; Europe and Asia,
Late Triassic to Middle Jurassic.
Williamsonia: a genus of trees in the family Williamsoniaceae; known worldwide
from cones and fern-like leaves; Late Triassic to Late Cretaceous.
Williamsoniaceae: a family of Mesozoic trees in the still-extant division
Spermatophyta and extinct order Bennettitales; their reproductive processes are
enigmatic, but they were similar in overall appearance to palms, growing as high as
33 feet; global distribution, Triassic to Cretaceous.
Winneshiek Site: a Middle Ordovician Lagerstätte in a meteorite crater in Iowa;
discovered in 2005, the site has produced significant eurypterids (see Pentecopterus),
conodonts, phyllocarids (see Ceratiocarididae), and other indeterminate fossils;
United States, Middle Ordovician.
Wiwaxia: a small, slug-like but armored animal of the middle Cambrian; first
found in the Burgess Shale group and identified by Walcott as a polychaete worm,
it is now sometimes classified as a mollusk, possibly a stem-polyplacophoran; it
resembled a miniature sea urchin except that it had only two rows of spikes on its
head and eight rows of carbonaceous sclerites covering its body, similar to those of
W 307
an isopod; isolated specimens have been found also in the United States, China, and
Russia, dated between 513 and 505 Ma.
Wukongopteridae: a small family of pterosaurs represented by the genera
Darwinopterus and Wukongopterus (the latter is poorly known); the family is an
example of modular evolution, having a long tail and other characteristics of the
earliest pterosaurs, combined with the skull features of the more advanced pterodac-
tyloids; China and England, Middle to Late Jurassic, 165–153 Ma.
Wynyardiidae: a small extinct family of marsupials in the still-extant order
Diprotodontia; similar to the phalangerids, the Wynyardiidae are intermediate
between the primitive polyprotodonts and the advanced diprotodonts; Australia,
Oligocene to Miocene.
X
Xenacanthida: an order of late Paleozoic freshwater sharks in the superorder

Euselachii; formerly called Pleuracanthodii. Eel-like and as long as 13 feet, they are
characterized by two-pronged teeth and a large serrated spine extending backward
from the rear of the braincase. They appeared in the Carboniferous and spread
worldwide in the Permian, but only a few genera survived the Permian extinctions
before disappearing in the Middle Triassic.
Xenarthra: a clade of unusual placental mammals of the Americas that includes
the living sloths, anteaters, and armadillos, as well as many extinct ancestors,
including the glyptodonts, pampatheres, and ground sloths. They were formerly
referred to as paratheres, or as edentates because of the lack of enamel on their teeth.
Many genera are marked by varying degrees of body armor in the form of dermal
ossicles. The anteaters of Africa and Asia (such as the aardvarks, echidnas, numbats,
and pangolins) are unrelated, having evolved convergently to fill the same ecologi-
cal niche. North and South America, Paleocene to Recent.
Xenodiscus: a genus of ceratitic ammonoids in the superfamily Xenodiscoidea;
evolute and thinly discoidal, 2–4 inches wide; North America, Asia, and Madagascar;
middle Permian to Early Triassic.
Xenopithecus: a proposed genus known only from a disputed jaw fragment, pos-
sibly of an early hominoid ape; Kenya, late Oligocene, 28 Ma.
Xenungulata: a still-obscure order of large South American ungulates placed in
the extinct superorder Meridiungulata; the order currently contains one family, the
Carodniidae, with three genera; its inconclusive similarities to several other very
different and widespread mammalian groups (such as the primitive Pantodonta, the
Uintatheriidae of the Eocene, and the Embrithopoda) leave it incertae sedis;
Paleocene to Eocene.

310 X
Xenusion: a genus of basal onychophorans or lobopods in the class Xenusia;

poorly known from two specimens found in glacial debris; see also Onychophora;
Europe, early Cambrian.
Xiaotingia zhengi: a species of feathered theropod dinosaurs from the Late
Jurassic, about the size of a chicken; described in 2011, its classification and signifi-
cance are still in doubt; China, 155 Ma.
Xiongguanlong baimoensis: a 5-foot-long dinosaur discovered in the southern
Gobi Desert and described in 2010. It seems to be an intermediate ancestor of the
large tyrannosaurs of the Late Cretaceous. Although earlier fossils of much smaller
ancestors of tyrannosaurs are known from England and China, Xiongguanlong is an
important mid-weight relative that exhibits some of the characteristic features of the
effective predator T. rex: broader skull attachments that made massive, more power-
ful jaw muscles possible, and thicker vertebrae to support a larger head. China,
Xiphactinus: “sword-ray,” a genus of large predatory actinopterygian fish in the
family Ichthyodectidae; between 15 and 20 feet long, it is characterized by a large
gape and four large fangs at the tip of its snout for grasping prey; a specimen at the
Royal Tyrrell Museum in Alberta, Canada, is famous for having a well-preserved
6-foot ichthyodectid fish (Gillicus) inside its stomach; global distribution, Early to
Late Cretaceous.
Xiphodontidae: an extinct family of artiodactyl ungulates in the suborder
Tylopoda. The xiphodonts were generally slender and camel-like but known only in
Europe. Since the early evolution of the camel family is restricted to North America,
they probably evolved separately, but a paucity of intermediate specimens makes
their relationship unclear. The type genus Xiphodon is well-known in the late
Eocene, and Paraxiphodon from the Eocene to the early Oligocene.
Xiphosurida: a still-extant order of marine chelicerate arthropods in the class
Xiphosura; the order includes several Paleozoic families and is represented today by
Limulus and Tachypleus, the horseshoe crabs; see also Lunataspis; global distribu-
tion, Late Ordovician to Recent.
Xylotitan: a genus of dwarf brontotheres; the most abundant mammal in the Clarno
Nut Beds of Oregon, dated about 44 Ma; North America, middle Eocene.
Y
Ya Ha Tinda: a Konservat-Lagerstätte in Alberta, first described in 2017. Located

in the Lower Jurassic Fernie Formation, the site has produced well-preserved articu-
lated marine vertebrates, mollusks, crinoids, crustaceans, and some plant fossils.
Like the Holzmaden, Strawberry Bank, and Osteno sites of Europe, it is dated at
183 Ma, near the boundary of the Pliensbachian and Toarcian Stages of the Early
Jurassic. Significantly, its fauna is remarkably similar to those of the European sites.
Canada, Early Jurassic.
Yawunik kootenayi: a species of Cambrian arthropods found in 2014 at the Marble
Canyon site of the Burgess Shale; it had an external skeleton, segmented body, and
jointed appendages; it could move its frontal appendages backward and forward
when swimming, and caught prey in the toothed claws on some appendages;
Yinotheria: a proposed subclass of mammals to include several Mesozoic families
and the still-living monotremes, the platypus and four species of echidnas; Late
Triassic to Recent.
Yixian Formation: a Lower Cretaceous (Barremian to early Aptian) formation in
Liaoning Province; Yixian fossils form the larger part of the Jehol biota, underlying
the equally important Jiufotang Formation; China, Early Cretaceous.
Younginidae: a late Permian family of small diapsid reptiles, formerly placed in
the discarded classifications Eosuchia and Younginiformes but now tentatively
placed in the clade Neodiapsida; the type species Youngina capensis lived in
Southern Africa at the end of the Permian and may have survived into the Triassic;
Africa and Madagascar, late Permian.
Yuknessia: a genus of colonial hemichordate animals in the class Pterobranchia; it
extended long branches from a holdfast and is known from the Burgess Shale and
other coeval sites; North America and Asia, middle Cambrian.
Yunnanocopia: see Lophogastrida.

312 Y
Yunnanolepis: a genus of primitive placoderms in the order Antiarchi; China,

Early to Middle Devonian.
Yunnanozoon: a genus of early Cambrian animals in the Chengjiang biota; Y. livi-
dum, similar to Pikaia of the Burgess Shale, may be the earliest-known hemichor-
date; China, 530 Ma.
Yutyrannus huali: a species of large coelurosaurian dinosaurs in the superfamily
Tyrannosauroidea, described in 2012 and dated at 125 Ma. About 30 feet long and
weighing 1.5 tons, Yutyrannus is the largest known feathered dinosaur. The three
specimens are also notable for being almost completely articulated fossils with
three-fingered hands. Feathers are clearly present in several areas of the body, the
longest being 8-inch neck feathers and 6-inch tail feathers. Yixian formation,
Liaoning Province, China; Early Cretaceous.
Z
Zalambdalestidae: a family of small, primitive eutherian mammals; now consid-

ered stem-eutherians and probably not placental; the shrew-like, mostly insectivo-
rous Zalambdalestes was about a foot long; Mongolia, Late Cretaceous, possibly to
Paleocene.
Zhongjianichthys: a poorly known genus of basal chordate animals; eel-like and
jawless, it is about an inch long and has a thick skin and a ventral fin the length of
its body; proposed as a relative of Myllokunmingia; Chengjiang biota, China, early
Cambrian.
zoarium: a collection of individual zooids that make up a compound or colonial
organism; some bryozoans form zoaria more than a foot wide.
zone fossil: see index fossil.
zooecia: singular zooecium, the chambers constructed by individual bryozoans.
Zosterophyllum: a genus of primitive vascular land plants in the subdivision
Lycophytina and extinct paraphyletic class Zosterophyllopsida; characterized by
kidney-shaped sporangia growing near the ends of the branches; global distribution,
Silurian to Late Devonian.
Zuniceratops: a monospecific genus of ornithischian dinosaurs in the suborder
Ceratopsia; Z. christopheri is known from one skull and the bones of several indi-
viduals, first found in 1996; only 10 feet long and weighing less than 300 pounds,
Zuniceratops seems to represent a transitional stage between the earliest ceratop-
sians and the larger, more familiar ones that were common in North America toward
the end of the Cretaceous; New Mexico, 91 Ma.
Zygentoma: the silverfish, a still-extant order of insects previously grouped with
the Archaeognatha in the obsolete order Thysanura; characterized by an elongate
body and at least two cerci and one medial extension on the tail; cosmopolitan,

314 Z
known definitively from the Middle Devonian but probably originating in the late
Silurian.
Zygolophodon: a genus of proboscideans in the family Mammutidae; one of the
largest terrestrial mammals, it was 14 feet high at the shoulder, weighed 15–18 tons,
and bore a pair of 13-foot tusks; North America, Europe, Asia, and Africa; Miocene
to Pleistocene.
Further Reading
Aldridge, R.J., ed. 1987. Paleobiology of Conodonts. Chichester: Ellis Horwood.

Allaby, Michael, and Ailsa. 2013. Oxford Dictionary of Geology and Earth Sciences. Oxford:
Oxford University Press.
Allman, Warren D., and David J. Bottjer. 2001. Evolutionary Paleoecology. New York: Columbia
University Press.
Arbour, Victoria M., and David C. Evans. 2017. A new ankylosaurine dinosaur from the Judith
River formation of Montana. Royal Society Open Science. https://doi.org/10.1098/rsos.161086.
Armstrong, Howard A., and Martin D. Brasier. 2005. Microfossils. Oxford: Blackwell Publishing.
Baron, Matthew G., David B. Norman, and Paul M. Barrett. 2017. A new hypothesis of dino-
saur relationships and early dinosaur evolution. Nature 543: 501–506. https://doi.org/10.1038/
nature.21700.
Beerling, David. 2007. The Emerald Planet: How Plants Changed Earth’s History. Oxford:
Oxford University Press.
Benton, Michael J., ed. 1993. The Fossil Record 2. 2nd ed. London: Chapman and Hall.
———. 1998. The quality of the fossil record of the vertebrates. In The Adequacy of the Fossil
Record, ed. Donovan and Paul, 269–303.
Benton, Michael. 2003. When Life Nearly Died. London: Thames and Hudson.
Benton, Michael J. 2009. Paleontology and the history of life. In Evolution: The First Four Billion
Years, ed. Ruse and Travis.
———. 2014. Vertebrate Palaeontology. 4th ed. Boston: Wiley-Blackwell.
Berta, Annalisa. 2012. Return to the Sea. Berkeley: U.C. Press.
Berta, Annalisa, and James Sumich. 2015. Marine Mammals: Evolutionary Biology. 3rd ed.
Amsterdam: Elsevier.
Bitter, von, H. Peter, et al. 2007. Eramosa Lagerstätte — Exceptionally preserved soft-bodied
biotas . . . (Silurian, Ontario, Canada). Geology 35: 879–882.
Blazejowski, B., et al. 2016. Ancient animal migration: A case study of eyeless, dimorphic
Devonian trilobites from Poland. Palaeontology 59: 743–759.
Boardman, Richard S., Alan Cheetham, and Albert J. Rowell. 1987. Fossil Invertebrates. Palo
Alto: Blackwell Scientific.
Botting, Joseph. 2016. Diversity and ecology of sponges in the Early Ordovician Fezouata biota,
Morocco. Palaeogeography, Palaeoclimatology, Palaeoecology 460. https://doi.org/10.1016/j.
palaeo.2016.05.018.
Briggs, Derek, and Jean-Bernard Caron. 2017. A large Cambrian chaetognath with supernu-
merary grasping spines. Current Biology 27 (16): 2536–2543.e1. https://doi.org/10.1016/j.
cub.2017.07.003.
Briggs, Derek, and Peter R. Crowther. 2001. Palaeobiology II. Oxford: Blackwell Science.

https://doi.org/10.1007/978-3-319-73055-4
316 Further Reading
Briggs, Derek, et al. 1994. The Fossils of the Burgess Shale. Washington: Smithsonian Institution
Press.
———. 2016. Bivalved arthropods from the Middle Ordovician Winneshiek Lagerstätte, Iowa,
USA. Journal of Paleontology 89 (6): 991–1006.
Brown, Caleb M., et al. 2017. An exceptionally well-preserved three-dimensional armored dino-
saur reveals insights into coloration and Cretaceous predator-prey dynamics. Current Biology
27 (16): 2514–2521.e3. https://doi.org/10.1016/j.cub.2017.06.071.
Brusatte, Stephen, and Zhe-Xi Luo. 2016. Ascent of the mammals. Scientific American 314 (6):
30–35.
Brusca, Richard, and Gary J. Brusca. 1990. Invertebrates. Oxford: Sinauer Associates.
Buffetaut, Eric. 2005. A new sauropod dinosaur with prosauropod-like teeth from the Middle
Jurassic of Madagascar. Bulletin de la Societé Géologique de France 176 (5): 467–473.
Burnham, David A., et al. 2000. Remarkable new bird-like dinosaur from the Upper Cretaceous of
Montana. U. of Kansas Paleontological Contributions, no. 13, March 2000.
Burrow, Carole J., and Michael A. Murphy. 2016. Early Devonian vertebrates from the northern
Roberts Mountains, Nevada. Journal of Paleontology 89: 734–740.
Butterfield, Nicholas J. 2000. Bangiomorpha pubescens: implications for the evolution of sex, mul-
ticellularity, and the Mesoproterozoic/Neoproterozoic radiation of eukaryotes. Paleobiology.
https://doi.org/10.1666/0094-8373 (2000) 026.
Carpenter, Kenneth. 1999. Eggs, Nests, and Baby Dinosaurs: A Look at Dinosaur Reproduction.
Bloomington: Indiana University Press.
Carroll, Robert L. 1988. Vertebrate Paleontology and Evolution. New York: W.H. Freeman and
Co.
———. 1997. Patterns and Processes of Vertebrate Evolution. Cambridge, U.K.: Cambridge
University Press.
Chang, Mee-Mann, et al. 2008. The Jehol Fossils. Amsterdam: Academic Press.
Chiappé, Luis, and Lawrence Witmer, eds. 2002. Mesozoic Birds. Berkeley: University of
California Press.
Clack, Jennifer. 2012. Gaining Ground: The Origin and Evolution of Tetrapods. 2nd ed.
Bloomington: Indiana University Press.
Clack, Jennifer, et al. 2016. Phylogenetic and environmental context of a Tournaisian tetrapod
fauna. Nature Ecology & Evolution 1 (0002). https://doi.org/10.1038/s41559-016-0002.
Clark, John, and Stella Stiegeler. 2000. The Facts on File Dictionary of Earth Sciences. New York:
Infobase Publishing.
Clarke, Tom. 2002. Oldest fossil footprints on land. Nature. https://doi.org/10.1038/news.020429-2.
Clarkson, E.N.K. 1998. Invertebrate Palaeontology and Evolution. 4th ed. Oxford: Wiley-Blackwell.
Cleal, Christopher J., and Barry A. Thomas. 1999. Plant Fossils. Woodbridge: Boydell Press.
———. 2010. Botanical nomenclature and plant fossils. Taxon 59: 261–268.
Colbert, Edwin. 2001. Evolution of the Vertebrates. New York: Wiley.
Cowen, Richard. 2013. The History of Life. 5th ed. New York: Wiley.
Davidson, Jane P. 2008. A History of Paleontology Illustration. Bloomington: Indiana University
Press.
Deméré, Thomas, et al. 2008. Morphological and molecular evidence for a stepwise evolutionary
transition from teeth to baleen in mysticete whales. Systematic Biology 57 (1): 15–37.
Donovan, S.K. 1994. The Palaeontology of Trace Fossils. New York: Wiley.
Donovan, S.K., and C.R.C. Paul. 1998. The Adequacy of the Fossil Record. New York: Wiley and
Sons.
Eldredge, Niles. 1991. Fossils: The Evolution and Extinction of Species. Princeton: Princeton
University Press.
Elliot, David K. 2012. Earth Science: Earth’s Surface and History, 2 vols. Ipswich: Salem Press.
Ellis, Richard. 2001. Aquagenesis: The Origin and Evolution of Life in the Sea. New York: Viking.
Erickson, G.M., et al. 2017. Dinosaur incubation periods directly determined from growth-line
counts in embryonic teeth show reptilian-grade development. Proceedings of the National
Academy of Sciences 114 (3). https://doi.org/10.1073/pnas.1613716114.
Further Reading 317
Erwin, Douglas H. 2006. Extinction. Princeton: Princeton U. Press.

Erwin, Douglas H., and James Valentine. 2013. The Cambrian Explosion. Greenwood Village:
Roberts and Co..
Fara, E. 2002. Sea level variations and the quality of the continental fossil record. Journal of the
Geological Society, London 159: 489–491.
Farlow, James O., and M.K. Brett-Surman, eds. 1997. The Complete Dinosaur. Bloomington:
Indiana University Press.
Fedonkin, Mikhail A., et al. 2007. The Rise of Animals: Evolution and Diversification of the
Kingdom Animalia. Baltimore: Johns Hopkins University Press.
Feist, Monique. 2014. Monoecious Sycidium and other charophytes. Review of Paleobotany and
Palynology 200: 88–195.
Feldmann, Rodney M., et al. 2017. Earliest occurrence of lophogastrid mysidacean arthropods
from the Anisian Luoping biota, Yunnan Province, China. Journal of Paleontology 91 (1):
100–115.
Foelix, Rainer F. 2011. Biology of Spiders. 3rd ed. Oxford: Oxford University Press.
Foote, Michael, and Arnold Miller. 2007. Principles of Paleontology. 3rd ed. New York:
W.H. Freeman and Company.
Forey, Peter L. 2015. History of the Coelacanth Fishes. Berlin: Springer Verlag.
Fortey, Richard. 1998. Life: A Natural History of the First Four Billion Years of Life on Earth.
New York: Knopf.
———. 2015. Fossils: The Key to the Past. 5th ed. Ithaca: Cornell University Press.
Foster, John. 2014. Cambrian Ocean World. Bloomington: Indiana University Press.
Frickhinger, Karl Albert. 1995. Fossil Atlas: Fishes. Translated by R.P.S. Jefferies. Melle: Mergus
Verlag.
Gabbott, Sarah E., et al. 2016. The Late Ordovician Soom shale Lagerstätte: an extraordinary
post-glacial fossil and sedimentary record. Journal of the Geological Society 174 (1). https://
doi.org/10.1144/jgs.2016-076.
Galton, Peter M., and Larry D. Martin. 2002. Enaliornis, an Early Cretaceous Hesperornithiform
Bird from England. In Chiappé, Mesozoic Birds.
Garrison, Nicole L., Juanita Rodriguez, et al. 2016. Spider phylogenomics: untangling the spider
tree of life. PeerJ 4: e1719. https://doi.org/10.7717/peerj.1719.
Gass, Kenneth. 2015. Solving the Mystery of the First Animals on Land. Manchester: Siri Scientific
Press.
Gaudin, Timothy J., et al. 2016. Skeletal anatomy of the North American pangolin Patriomanis
americana (Mammalia, Pholidota) from the latest Eocene of Wyoming (USA). Smithsonian
Contributions to Paleobiology 98: 1–102.
Geisler, Jonathan H., et al. 2017. The origin of filter feeding in whales. Current Biology 27 (13):
2036–2042.e2. https://doi.org/10.1016/j.cub.2017.06.003.
Gensel, Patricia G., and Dianne Edwards. 2001. Plants Invade the Land. New York: Columbia
University Press.
Glut, Donald. 1997. Dinosaurs: The Encyclopedia. North Carolina: McFarland and Co..
Gon, Sam O. 2015. A Guide to the Orders of Trilobites. Trilobites.Info. http://www.trilobites.info/
trilobite.htm.
Gould, Stephen J. 1987. Time’s Arrow, Time’s Cycle. Boston: Harvard University Press.
———. 1989. Wonderful Life: The Burgess Shale and the Nature of History. New York:
W.W. Norton.
———. 1993. Eight Little Piggies. New York: W.W. Norton.
———. 1996. Full House. New York: Harmony Books.
———. 2002. The Structure of Evolutionary Theory. Cambridge: Harvard University Press.
Graham, Linda E. 1993. Origin of Land Plants. New York: Wiley.
Grimaldi, David. 2009. Fossil record [of insects]. In Encyclopedia of Insects, ed. Resh, 455–463.
Academic Press.
318 Further Reading
Grimaldi, David, and Michael S. Engel. 2005. Evolution of the Insects. Cambridge: Cambridge
University Press.
Hanai, Tetsuro, et al., eds. 1988. Evolutionary Biology of Ostracoda. New York: Elsevier.
Hegna, Thomas A., et al. 2017. Pyritized in situ trilobite eggs from the Ordovician of New York.
Geology 45 (3). https://doi.org/10.1130/G38773.1.
Hembree, Daniel I. 2007. Phylogenetic revision of Rhineuridae ... from the Eocene to Miocene of
North America. University of Kansas Paleontological Contributions 15: 1–20.
Henke, Winfried, and Ian Tattersall, eds. 2007. Handbook of Paleoanthropology, 3 vols. Berlin:
Springer Verlag.
Hilton, Richard P. 2003. Dinosaurs and Other Mesozoic Reptiles of California. Berkeley:
University of California Press.
Hopkins, Samantha. 2005. The evolution of fossoriality and the adaptive role of horns in the
Mylagaulidae (Mammalia: Rodentia). Proceedings of the Royal Society B 272: 1705–1713.
Hopson, James A. 1994. Synapsid evolution and the radiation of non-eutherian mammals. In
Major Features of Vertebrate Evolution, ed. Prothero and Schoch, 190–219.
Jackson, Frankie D., et al. 2013. Influence of vertisol development on sauropod egg taphonomy.
Palaeogeography, Palaeoclimatology, Palaeoecology 3 (386): 300–307.
Jaeger, Edmund C. 1959. A Source-Book of Biological Names and Terms. Springfield: Thomas.
Janvier, Philippe. 1996. Early Vertebrates. Oxford: Oxford University Press.
Jenkins, D.G., and John W. Murray, eds. 1989. Stratigraphical Atlas of Fossil Foraminifera. 2nd
ed. London: Ellis Horwood.
Kah, Linda, et al. 2009. Reinterpreting a Proterozoic enigma: Conophyton–Jacutophyton stro-
matolites of the Mesoproterozoic Atar Group, Mauritania. Perspectives in Carbonate Geology:
A Tribute to the Career of Robert Nathan Ginsburg (Special Publication 41 of the IAS) 98:
277–295.
Kemp, T.S. 1999. Fossils and Evolution. Oxford: Oxford University Press.
———. 2005. The Origin and Evolution of Mammals. Oxford: Oxford University Press.
Knoll, Andrew H. 2003. Life on a Young Planet. Princeton: Princeton University Press.
Ksepka, Daniel T., Thomas A. Stidham, and Thomas E. Williamson. 2017. Early Paleocene land-
bird supports rapid phylogenetic and morphological diversification of crown birds after the
K–Pg mass extinction. Proceedings of the National Academy of Sciences 114. https://doi.
org/10.1073/pnas.1700188114.
Kuhn-Schnyder, Emil, and Hans Rieber. 1984. Handbook of Paleozoology. Baltimore: Johns
Hopkins University Press.
Lambert, David. 2002. The Encyclopedia of Prehistory. New York: Facts on File.
Laubichler, Manfred D., and Karl J. Niklas. 2009. The morphological tradition in German pale-
ontology: Otto Jaekel, Walter Zimmermann, and Otto Schindewolf. In The Paleobiological
Revolution, Essays on the Growth of Modern Paleontology, ed. Sepkoski and Ruse, 278–300.
Chicago: University of Chicago Press.
Le Herisse, Alain, et al. 2012. The end of a myth: Arpylorus antiquus a Paleozoic dinoflagellate
cyst. Palaios 27 (6): 414–423.
Levi-Setti, Riccardo. 1993. Trilobites. Chicago: University of Chicago Press.
———. 2014. The Trilobite Book. Chicago: University of Chicago Press.
Lieberman, Bruce S. 2000. Paleobiogeography: Using Fossils to Study Global Change, Plate
Tectonics, and Evolution. New York: Kluwer Academic.
Lincoln, Roger, et al. 1998. A Dictionary of Ecology, Evolution and Systematics. Cambridge
University Press.
Losos, Jonathan B. 2014. The Princeton Guide to Evolution, ed. et al. Princeton: Princeton
University Press.
Lu, Junchang, et al. 2016. A Late Cretaceous diversification of oviraptorid dinosaurs: evidence
from a new species preserved in an unusual posture. Scientific Reports. nature.com. doi: https://
doi.org/10.1038/srep.35780.
Further Reading 319
Macdougall, J. Douglas. 2004. Frozen Earth: The Once and Future Story of Ice Ages. Berkeley:
University of California Press.
MacFadden, Bruce J. 1992. Fossil Horses: Systematics, Paleobiology, and Evolution of the Family
Equidae. Cambridge, U.K.: Cambridge University Press.
Maisey, John G. 1996. Discovering Fossil Fishes. New York: Henry Holt.
Marshall, Stephen A. 2006. Insects: Their Natural History and Diversity. Buffalo: Firefly Books.
Martindale, Rowan C., T.R. Them, B.C. Gill, S.M. Marroquín, and A.H. Knoll. 2017. A new Early
Jurassic (ca. 183 Ma) fossil Lagerstätte from Ya Ha Tinda, Alberta, Canada. Geology 45: 255–
225. https://doi.org/10.1130/G38808.1. http://geology.gsapubs.
McGhee, George R., Jr. 1996. The Late Devonian Mass Extinction. New York: Columbia U. Press.
McLaughlin, Kathleen. 2017. The footprints of giants. Science 356 (6344): 1224–1226.
McLellan, Bruce, and David Reiner. 1994. A review of bear evolution. International Conference of
Bear Research and Management 9 (1): 85–96.
Miller, Randall F. 2003. The oldest articulated chondrichthyan from the Early Devonian period.
Nature 405: 501–504.
Miller, William, ed. 2007. Trace Fossils. Amsterdam: Elsevier.
Milsom, Clare, and Sue Rigby. 2010. Fossils at a Glance. Oxford: Wiley-Blackwell.
Monks, Neale, and Philip Palmer. 2002. Ammonites. London: Natural History Museum.
Morris, Christopher, ed. 1995. Academic Press Dictionary of Science and Technology. San Diego:
Academic Press.
Morris, Simon Conway. 1998. The Crucible of Creation. Oxford University Press.
Moysiuk, Joseph, et al. 2017. Hyoliths are Paleozoic lophophorates. Nature 541: 394–397. https://
doi.org/10.1038/nature 20804.
Murdock, Duncan J.E., Sarah E. Gabbott, and Mark A. Purnell. 2016. The impact of tapho-
nomic data on phylogenetic resolution: Helenodora inopinata (Carboniferous, Mazon Creek
Lagerstätte) and the onychophoran stem lineage. BMC Evolutionary Biology 16: 19. https://
doi.org/10.1186/s12862-016-0582-7.
Murray, John W., ed. 1985. Atlas of Invertebrate Macrofossils. New York: Wiley and Sons.
Neuendorf, Klaus, James Mehl, and Julia Jackson. 2011. Glossary of Geology. Washington, D.C.:
American Geosciences Institute.
Ni, X., et al. 2016. A late Paleocene probable metatherian (?deltatheroidan) survivor of the
Cretaceous mass extinction. Scientific Reports 6 (38547): 1–9. https://doi.org/10.1038/srep
38547.
Nichols, Douglas J., and Kirk R. Johnson. 2008. Plants and the K-T Boundary. Cambridge:
Cambridge University Press.
Niklas, Karl J., and Tom L. Phillips. 1976. Morphology of Protosalvinia from the Upper Devonian
of Ohio and Kentucky. American Journal of Botany 63 (1): 9–29.
Ogg, James G., Gabi Ogg, and Felix M. Gradstein. 2016. A Concise Geologic Time Scale.
Amsterdam: Elsevier.
Owen, Elizabeth, and Eve Daintith. 2004. The Facts on File Dictionary of Evolutionary Biology.
New York: Infobase Publishing.
Pagel, Mark. 2002. Encyclopedia of Evolution, 2 vols. Oxford: Oxford University Press.
Palmer, Douglas. 1999a. Atlas of the Prehistoric World. New York: Discovery Books.
———, ed. 1999b. The Marshall Illustrated Encyclopedia of Dinosaurs and Prehistoric Animals.
London: Marshall.
———. 2009. Evolution: The Story of Life. Berkeley: University of California Press.
Palmer, Douglas, et al. 2009. Prehistoric Life: The Definitive Visual History of Life on Earth.
London: Dorling Kindersley.
Parker, Sybil. 2017. McGraw-Hill Dictionary of Scientific and Technical Terms. 7th ed.
Peecook, Brandon R., and Christian A. Sidor. 2015. The first dinosaur from Washington state and
a review of Pacific Coast dinosaurs from North America. PloS One 10 (5): e0127792. https://
doi.org/10.1371/journal.pone.0127792.
320 Further Reading
Penney, David. 2016. Amber Paleobiology: Research Trends and Perspectives for the 21st Century.
Manchester: Siri Scientific Press.
Poinar, George, and Roberta. 1999. The Amber Forest. Princeton: Princeton University Press.
Prothero, Donald R. 2007. Evolution: What the Fossils Say and Why It Matters. New York:
Columbia University Press.
———. 2015. The Story of Life in 25 Fossils. New York: Columbia University Press.
Prothero, Donald R., and Scott Foss, eds. 2007. The Evolution of Artiodactyls. Baltimore: Johns
Hopkins University Press.
Prothero, Donald R., and Robert M. Schoch. 1994. Major Features of Vertebrate Evolution.
Paleontological Society Short Courses in Paleontology, No. 7.
Raup, D.M. 1983. On the early origins of major biologic groups. Paleobiology 9: 107–115.
Reader, John. 2011. Missing Links: In Search of Life’s Origins. Oxford: Oxford University Press.
Resh, Vincent H., and Ring T. Cardé. 2009. Encyclopedia of Insects. Amsterdam: Academic Press.
Retallack, Gregory J. 1998. Fossil soils and completeness of the rock and fossil records. In The
Adequacy of the Fossil Record, ed. Donovan and Paul, 133–164.
———. 2001. Soils of the Past: An Introduction to Paleopedology. Oxford: Blackwell Science.
———. 2013. Ediacaran life on land. Nature 493 (7430): 89–92.
Retallack, Gregory J., et al. 1996. Reconstructions of Eocene and Oligocene plants and animals of
central Oregon. Oregon Geology 58 (3): 51–59.
Rich, Patricia V., et al. 1996. The Fossil Book: A Record of Prehistoric Life. Dover publications.
[New edition of the Fentons’ book from 1958, 1986, and 1989.]
Romano, Carlo, et al. 2017. Marine early Triassic Actinopterygii from Elko County (Nevada,
USA): implications for the Smithian equatorial vertebrate eclipse. Journal of Paleontology 1.
https://doi.org/10.1017/jpa.2017.36.
Rudkin, David M., et al. 2008. The oldest horseshoe crab: a new xiphosurid from Late Ordovician
Konservat-Lagerstätten deposits, Manitoba, Canada. Paleontology 51 (1): 1–9.
Ruse, Michael, and Joseph Travis, eds. 2009. Evolution: The First Four Billion Years. Cambridge:
Belknap Press.
Sanchez, Sophie, Paul Tafforeau, Jennifer Clack, and Per Ahlberg. 2016. Life history of the stem
tetrapod Acanthostega revealed by synchrotron microtomography. Nature 537: 408–411.
https://doi.org/10.1038/nature.19354.
Sarjeant, W.A.S. 2013. Fossil and Living Dinoflagellates. Amsterdam: Elsevier.
Schindewolf, Otto. 1950. Die Grundfragen der Paläontologie. Stuttgart: Schweizerbart Verlag.
Trans. By Judith Schaefer (1993), Basic Questions in Paleontology. University of Chicago
Press.
Schoch, Rainer. 2014. Amphibian Evolution: The Life of Early Land Vertebrates. London: Wiley
Blackwell.
Schopf, J.W., ed. 1983. Earth’s First Biosphere: Its Origin and Evolution. Princeton: Princeton
University Press.
———. 2009. Emergence of Precambrian paleobiology: a new field of science. In The
Paleobiological Revolution, ed. Sepkoski and Ruse, 89–110. Chicago: Chicago University
Press.
Schopf, J.W., and Cornelius Klein. 1992. The Proterozoic Biosphere. Cambridge University Press.
Schultka, Stephan. 1991. Erster nachweis der gattung Euthycarcinus (Arthropoda) aus dem
obercarbon von Ibbenbüren (Nordrhein-Westfalen, Deutschland). International Series of
Monographs in Pure and Applied Biology. https://doi.org/10.1007/BF 02989847.
Selden, Paul A. 1996a. First fossil mesothele spider, from the Carboniferous of France. Revue
Suisse de Zoologie 2: 585–596.
———. 1996b. Fossil mesothele spiders. Nature 379: 498–499.
Selden, Paul A., and John Nudds. 2008. Fossil-Lagerstätten. Geology Today 24 (4): 153–158.
———. 2012. The Evolution of Fossil Ecosystems. Amsterdam: Elsevier.
Selden, Paul A., and D. Penney. 2011. Fossil spiders. Biological Reviews 85: 171–206.
Further Reading 321
Selden, Paul A., et al. 2008. Fossil evidence for the origin of spider spinnerets, and a proposed
arachnid order. Proceedings of the National Academy of Sciences 105 (52): 20781–20785.
Sepkoski, J.J. 2002. A Compendium of Fossil Marine Animal Genera. Bulletins of American
Paleontology 363: 1–560.
Sepkoski, David. 2012. Rereading the Fossil Record. Chicago: University of Chicago Press.
Sepkoski, David, and Michael Ruse, eds. 2009. The Paleobiological Revolution: Essays on the
Growth of Modern Paleontology. Chicago: University of Chicago Press.
Sidor, C.A. 2003. The naris and palate of Lycaenodon longiceps (Therapsida: Biarmosuchia), with
comments on their early evolution in the Therapsida. Journal of Paleontology 77 (5): 977–984.
Singer, Ronald. 1999. Encyclopedia of Paleontology, 2 vols. Chicago: Fitzroy Dearborn.
Smith, Andrew B. 1994. Systematics and the Fossil Record. Oxford: Blackwell.
Smith, K.T., and A. Scanferla. 2016. Fossil snake preserving three trophic levels and evi-
dence for an ontogenetic dietary shift. Palaeobio Palaeoenv 2016. https://doi.org/10.1007/
s12549-016-02441.
Song, Haijun, Paul B. Wignall, Jinnan Tong, and Hongfu Yin. 2013. Two pulses of extinction dur-
ing the Permian-Triassic crisis. Nature Geoscience 6. https://doi.org/10.1038/ngeo1649.
Sperling, E.A., K.J. Peterson, and M. Laflamme. 2011. Rangeomorphs, Thectardis (Porifera?)
and dissolved organic carbon in the Ediacaran oceans. Geobiology 9: 24–33. https://doi.
org/10.1111/j.1472-4669.2010.00259.x.
Stanley, Steven M. 1987. Extinction. New York: Scientific. American Library no. 20.
Stevens, Margaret S., and B. James. 2007. Family Merycoidodontidae. In The Evolution of
Artiodactyls, ed. Prothero and Foss, 157–168.
Stewart, Wilson N., and Gar W. Rothwell. 1993. Paleobotany and the Evolution of Plants. 2nd ed.
Cambridge: Cambridge University Press.
Stöcker, Friedrich. 1986. ABC Biologie. Translated and revised by Thomas A. Scott (1995) as
Concise Encyclopedia: Biology. Berlin: De Gruyter.
Sweet, Walter C. 1988. The Conodonts. Oxford: Oxford University Press.
Sweet, Walter C., and Philip Donoghue. 2001. Conodonts: past, present and future. Journal of
Paleontology 75 (6): 1174–1184.
Talent, John A., ed. 2012. Earth and Life: Global Biodiversity, Extinction Intervals and
Biogeographic Perturbations through Time. Dordrecht: Springer Verlag.
Taylor, Paul D. 2004. Extinctions in the History of Life. Cambridge: Cambridge University Press.
Taylor, Paul D., and David N. Lewis. 2005. Fossil Invertebrates. Harvard University Press.
Taylor, Paul D., and Aaron O’Dea. 2014. A History of Life in 100 Fossils. Washington: Smithsonian
Books.
Taylor, Thomas N., Edith L. Taylor, and Michael Krings. 2009. Paleobotany: The Biology and
Evolution of Fossil Plants. 2nd ed. Amsterdam: Academic Press.
Tidwell, William D. 1998. Common Fossil Plants of Western North America. Washington:
Smithsonian Institution Press.
Torsvik, Trond H., and Robin Cocks. 2017. Earth History and Palaeogeography. Cambridge:
Cambridge University Press.
Traverse, Alfred. 2007. Paleopalynology. 2nd ed. Dordrecht: Springer Netherlands.
Trewin, Nigel H. 2013. Scottish Fossils. Edinburgh: Dunedin Academic Press.
Tsur, Samuel A. 1999. Elsevier’s Dictionary of the Genera of Life. Amsterdam: Elsevier.
Tudge, Colin. 2000. The Variety of Life. Oxford: Oxford University Press.
Vacelet, Jean, Philippe Willenz, and Willard D. Hartmann. 2010. Living hypercalcified sponges.
Treatise Online, Part E, rev. 4: 1.
Vannier, J., et al. 2016. Exceptional preservation of eye structure in arthropod visual predators
from the Middle Jurassic. Nature Communications 7. https://doi.org/10.1038/ncomms.10320.
Van Roy, Peter, et al. 2010. Ordovician faunas of Burgess shale type [Fezouata]. Nature 465:
215–218. https://doi.org/10.1038/nature.09038.
Varricchio, David J., and Frankie D. Jackson. 2016. Reproduction in Mesozoic birds and evolution
of the modern avian reproductive mode. The Auk 133 (4): 654–684.
322 Further Reading
Varricchio, David J., Frankie D. Jackson, John Borkowski, and John Horner. 1997. Nest and egg
clutches of the dinosaur Troodon formosus and the evolution of avian reproductive traits.
Nature 385: 247–250.
Varricchio, David J., J.R. Moore, G.M. Erickson, M.A. Norell, F.D. Jackson, and J.J. Borkowski.
2008. Avian paternal care had dinosaur origin. Science 322 (5909): 1826–1828.
Vaux, Felix, Steven Trewick, and Mary Morgan-Richards. 2016. Lineages, splits and divergence
challenge whether the terms anagenesis and cladogenesis are necessary. Biological Journal of
the Linnean Society 117 (2): 165–176. https://doi.org/10.1111/bij.12665.
Viira, Vive, and R.J. Aldridge. 1998. Upper Wenlock to Lower Pridoli (Silurian) conodont biostra-
tigraphy. Journal of Micropalaeontology 17 (1): 33–50.
Wacey, David. 2009. Early Life on Earth. In Springer Topics in Geobiology Series. New York:
Springer Sciences.
Weishampel, David B., et al. 1990. The Dinosauria. Berkeley: University of California Press.
Werdelin, Lars, and W.J. Sanders. 2010. Cenozoic Mammals of Africa. Berkeley: U.C. Press.
Whatley, Robin, and Caroline Maybury. 1990. Ostracoda and World Events. London: Chapman
and Hall.
Wheeler, Quentin D. 2010. Do we need to describe, name, and classify all species? In Beyond
Cladistics: The Branching of a Paradigm, ed. Williams and Knapp. Berkeley: University of
California Press.
White, Mary E. 1990. The Flowering of Gondwana. Princeton: Princeton University Press.
White, Tim. 2009. Ladders, bushes, punctuations, and clades: hominid paleobiology in the late
twentieth century. In The Paleobiological Revolution: Essays on the Growth of Modern
Paleontology, ed. Sepkoski and Ruse, 122–148. Chicago: University of Chicago Press.
Whittington, Harry B. 1992. Trilobites. Martlesham, U.K.: Boydell Press.
Williams, David M., and Sandra Knapp, eds. 2010. Beyond Cladistics: The Branching of a
Paradigm. Berkeley: University of California Press.
Williams, Matt, Michael J. Benton, and Andrew Ross. 2014. The Strawberry Bank Lagerstätte
reveals insights into Early Jurassic life. Journal of the Geological Society 172: 683–692.
https://doi.org/10.1144/jgs.2014-144.
Wilson, Edward O. 1999. The Diversity of Life. New York: Norton.
Wilson, Gregory P., Eric G. Ekdale, John W. Hoganson, Jonathan J. Calede, and Abby Vander
Linden. 2016. A large carnivorous mammal from the Late Cretaceous and the North
American origin of marsupials. Nature Communications 7: 13734. https://doi.org/10.1038/
ncomms.13734.
Winchester, Simon. 2001. The Map That Changed the World. New York: Harper Collins.
Xing, Lida, et al. 2017. A mid-Cretaceous enantiornithine (Aves) hatchling preserved in Burmese
amber with unusual plumage. Gondwana Research 49: 264–277. https://doi.org/10.1016/
j.gr.2017.06.001.
Zheng, Daran, et al. 2016. A new damsel-dragonfly from the Lower Jurassic of Northwestern
China and its paleogeographic significance. Journal of Paleontology 90 (3): 485–490.
Zhu, Min, et al. 2009. The oldest articulated osteichthyan reveals mosaic gnathostome characters.
Nature 458: 469–474.
Zimmer, Carl. 2014. The Tangled Bank: An Introduction to Evolution. Greenwood Village,
Colorado: Roberts and Company.

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Robert L.

Technical Editor: Brittney A. Oleniacz

ISBN 978-3-319-73054-7 ISBN 978-3-319-73055-4 (eBook)

Library of Congress Control Number: 2017963120

© Springer International Publishing AG 2018

Printed on acid-free paper

This Springer imprint is published by Springer Nature

This little book is a moderately comprehensive reference handbook covering the

Aachenosaurus: name given to a piece of petrified wood found in the Aachenian

© Springer International Publishing AG 2018 1

Acanthothiris: an obscure genus of articulate brachiopods in the still-extant order

Mammalodontidae. British Columbia, Washington, Oregon, Mexico, and Japan;

in 2015 as only 5–6 inches long, the shrew-like A. scansorius is the earliest-known

Agriotherium: “hunting beast,” a genus of early bears in the subfamily

allochthonous: referring to a fossil assemblage consisting of organisms removed

amniotes: animals (reptiles, birds, and mammals) which evolved ways of

Amphistium: a genus of flatfish showing the movement, over millions of years, of

Anarcestes: a genus of small ammonoids in the order Agoniatitida and family

Ancylopoda: a suborder of perissodactyl ungulate mammals characterized by

Anoplotheriidae: a family of artiodactyl ungulate mammals in the suborder

flight. Publicized in 1999 by National Geographic, it was soon discovered to be a

Archipolypoda: a superorder of millipedes in the subclass Chilognatha, possibly

early hominids’ development of bipedalism. Other paleontologists, however,

Asioryctitheria: a proposed order of basal eutherian mammals; insectivorous and

Atelostomata: a superorder of irregular echinoids characterized by the absence of

Australopithecus: a genus of primates in the family Hominidae. With specimens

Avalofractus: an Ediacaran genus of rangeomorphs; 2 inches long, its alternating

background extinction see extinction.

© Springer International Publishing AG 2018 31

Balaenidae: a still-extant family of whales in the suborder Mysticeti, including the

and rays, resemble wings; an example of convergent evolution compared to the

Bellinurus: an extinct genus of freshwater arthropods in the order Xiphosurida;

body plan: see Cambrian and Ediacaran biota.

Brachyceratops: an obscure genus of ceratopsian dinosaurs, known only from

Brontotheriidae: “thunder animal,” a large family of herbivorous perissodactyl

Burgess Shale: name given to one of the most important Konservat-Lagerstätten,

Caelifera: the grasshoppers and locusts, a suborder of chewing herbivorous insects

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Calcinea: a still-extant, probably paraphyletic subclass of sponges in the class

The marine environment was dominated at first by reef-building Archaeocyatha and

the modern lancelets, including dozens of modern genera in the Branchiostomiformes

Ceratopsia: “horned face,” a large suborder of herbivorous ornithischian ­dinosaurs

For the evolution of marine mammals, see Aegyptocetus, Ambulocetus,

Chancelloriidae: a poorly known group of sponge-shaped Cambrian organisms,

Chilopoda: a still-extant class of myriapods that includes the centipedes;

Cidaroida: a still-extant order of primitive sea urchins in the subclass

Clarkecaris: a genus of crustaceans in the order Anaspidacea and proposed family

Clypeus ploti: “swimming shield,” a species of irregular echinoids in the family

Coelophysis a genus of early saurischian dinosaurs, theropods in the family

Collyrites: a genus of small irregular euechinoids in the family Collyritidae,

continental drift: see paleogeography.

3,000 feet; see Rugosa, Tabulata, and Scleractinia; worldwide, Cambrian to

Corythosaurus: “Corinthian-helmet lizard,” a hadrosaurid dinosaur in the

Cryptovolans: see Microraptor.

photosynthesis, they may have significantly changed the composition of the

Dactylioceras: a genus of planulate ammonoids in the order Ammonitida; only

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Daphoenodon: a genus of large carnivorous canids in the family Amphicyonidae;

c­oincided with the worldwide Quaternary extinctions of many large mammals;

Dendropithecus: a genus of early primates in the superfamily Hominoidea and

Ceratopsia: “horned face,” a large suborder of herbivorous ornithischian dinosaurs

coincided with the worldwide Quaternary extinctions of many large mammals;

s ymmetry and is composed of bundles of muscles stacked horizontally, with short