VOL. 9 NO.

I - SPRING 2000 Page 14

Ethology and Attachment: A Historical Perspective

Eckhard H. Hess and Slobodan B. Petrovich'

An outline is presented ofthe assumptions underlying Suppose you were asked, 'Why does your thumb move in a
earlier and contemporary ethology. An example ofelhologi­ different way from the other fingers?' You might give an answer in
co! analysis is presented, wilh a locus on the on/ogeny, terms of the anatomy of the hand· the differences in skeletal
structure and muscle attachments between the thumb and the other
mediating mechanisms ofcausation, junction and evolution of
fingers: that would be an answer concerned with the immediate
cricket songs. In a his/arico/frame, conceptual and method­
causation of thumb movement. You might give an answer in terms
ological origillS 0/modem ethology were sketched out and the of the hand's embryology describing how, as the finger rudiments
significance a/important trends explored The conclusion of developed, one came to have a different structure from the others. Or
the analysis of this chapter is that researchers of human
This article is intended solely for the personal use of the individual user and is not to be disseminated broadly.

you might give a functional answer-an opposable thumb makes it

development studying attachment under Ihe aegis ofethologi­ easier for us to pick things up, climb trees, and so on. Or finally you
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ca/theory have moved in a direction that diverges/rom the might say that we are descended from monkey-like creatures, and
conceptualizations and research emphases ojcontemporary monkeys have opposable thumbs, so of course we do too. This
ethology. This paper describes elements ofthe ethological would be an answer in terms of evolutionary origi n. All of these
approach to attachment in an attempt tofacilitate an interdis­ answers would be correct: no one would be complete. In the same
way, ethologists are interested in questions of all four types of
ciplinary exchange among ethologists, and developmental
behavior. Indeed they believe that, although logically distinct and
psychologists. Its purpose is to share a historical perspectwe
independent, questions concerning immediate causation, develop­
and habits ofthought, and 10 communicate theoretical and ment, function and evolution are sometimes inter-fertile (p. 21).
methodological developmenls and that have had an Impact on
the ethological study of behavior. As an example, we will briefly review the literature of cricket
songs. The research involved is representative of ethological
At the, outset, our goal is to tell what ethology is about, in methodology. OUT review includes the types of questions
a historical context. As an example, the treattnent considers asked, the experimental subjects employed, the nature of the
the development, mediating mechanisms of causation, as well behavioral response studied and its measurement, comparative
as the function and evolution of the cricket's song. Then we analyses within and across species, as well as how ecological
extrapolate sOllie conceptual and methodoiogical lessons of example also can illustrate the process characteristic of
interest and of use to a wider audience. Our treatment ontogeny, causation, function, and evolution of species-typic
proceeds with an appraisal of the contributions of ethology to isolation and of identification in simpler invertebrate systems.
the study of human attachment. Finally. we focus on some of Invertebrates, and insects in particular, tell an interesting story
the issues of relevance both to ethology and developmental (e.g., Wilson, 1975) and their message (even though unheard
psychology. and evaluate whether or not human-development of in this volume) is important for 3n understanding of
theory and research on anachment in the frame of ethology species-typic behavioral development.
have diverged from the emphases of contemporary ethology.

CriCket-Song Study as an Example of

What Ethology Is About: A Historical
Behavioral Analyses in Ethology
Perspective On Some Conceptual And
Methodological Extrapolations There are approximately 3,000 species of crickets, of
which field crickets make up a special group of about 400.
Ethology has been described as the biology of behavior The field crickets are the most familiar. Relatively large, they
(Eibl-Eibesfeld� 1975; Hinde, 1982; Tinbergen, 1963). While are yellowish-brown insects known for their loud, musical
chirping. Male crickets produce sounds by rubbing together
ethology has a relatively long and interesting history
(Burghardt, 1986; Jaynes, 1969), for its most recent recogni­ stridulating areas located on the forewings and utilize a rapid
tion it owes much to the contributions of a small group of flunering motion to produce a typical vibrato chirp. The
investigators, among whom Lorenz (1965, 1970, 1971, 1974), receiving auditory organs are tympana located within slits on
Von Frisch (1967), and Tinbergen (1951, 1972) have received the forelegs. Most cricket species chirp at night, some during
widest recognition. the day, and others both day and night. In general, under­
Currently, the ethological literature on various aspects of standing of neurophysiological mechanisms involved in
animal (including human) behavior is so voluminous and cricket bioacoustics has few parallels, if any, in the animal
varied that it leaves one wondering what it is that ethologists literature (Alexander, 1966; Bentley & Hoy, 1974; Ewing &
do not study. Faced with similar concerns, Tinbergen (1963) Hoyle, 1964; Huber, 1962; Rose, 1986).
suggested that. once behavior is adequately described and In a southeastern region of the United States during
operationalized, ethologists study its ontogeny or develop­ summer there are as many as 20 different species of tree
ment, its immediate causation or mechanisms, its adaptive crickets producing discrete sounds, mostly the male's calling
significance or function, and its evolutionary origins. In the song, the function of which is to attract the female for mating,
words of Hinde (1982): How does a female distinguish the sounds of a conspecific?
Studies have demonstrated that males of each species have a
particular pulse rate in their song, and it is this pu lse rate that
 Page 15 VOL. 9 NO. I - SPRING 2000

provides a female with discriminative cues. It is also interest­ finnulus. These two sibling species of ground crickets mature
ing to nOle that the metabolic and physiological processes in a at the same lime, overlap geographically and ecologically, but
cricket are functionally affected by outside temperatures, so Sing different songs. Fulton was able to develop FI and F2
that a pulse rate in the song changes with temperature, earning hybrids, carry out F I backcrosses with parental species, and
some species the appropriate label of "thennometer crickets." analyze the songs of various crosses, Fulton's results were
'The refinement of the evolved system is remarkable when one generally elear CUI and slnlightforward. Pulses in the song of
considers that physiological mechanisms which detennine F I hybrids were delivered at a rate intennediate between those
females' responsiveness to a signal change at the same time in in the songs of the two parental generations. The songs of
a fashion that parallels the males' pulse rate. The sound­ backcross progeny were more like the parent utilized in the
producing repertoire of the male cricket serves a number of backcross. Subsequent literature on other species has fwther
runctions. elucidated the genetic detennination of the song panern of
I, facilitating and establishing sexual contact (the calling each cricket species. The songs art phenotypic expressions of
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song); different genotypes, thereby offering evidence that links

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2. mediating sexual attraction at a relatively short together genetic infonnation, developmental processes,
distance (the courtship song); structural and functional organization of the neuroendocrine
3. signaling departure of a cOW1ed female (the cowtship­ system, and behavior (e.g., Bentley & Hoy, 1974;
interruption song); Schildberger, 1984).
4. repelling or dominating other males (the aggressive In summary, crickets are sensitive to stimuli in other
sound); sensory channels: acoustic, chemical, visual, tactile, and
5. maintaining contact between a mated pair (the thenna!. This review has used one example to demonstrate
postcopulatory song); how discrete acoustic signals function in species-typic
6. a wide range of what appear to be recognition sounds isolation and identification, while it also orrers oven and
(e.g., Alexander, 1966, 1968). covert evidence for the proximate and the ultimate cauSlltion
How does this brief commentary on cricket bioacoustics of such behaviors.
illustrate the importance of acoustic communication in cricket
speciation and evolution? What are some of the factors that On the Relationship Between Elhological Theory
maintain the species-specific integrity of a gene pool of some
and Research: Levels of Organization-Levels of
20 different species of tree crickets that are not geographically
isolated? The species-specific characteristics of the male Analysis
calling song and the recognition of that song by a conspecific
female were identified as an imponant isolating mechanism The development and the use of theory have been valued by
(Alexander, 1966; Dixon & Cade, 1996; Walker, 1957; researchers across disciplines and areas of inquiry. The
Wiedennann & Loher, 1984). characteristic thinking has been that theory generates research
Viewed in the context of our understanding of evolution­ models and questions, thereby requiring that the empirical
ary processes, aickets tell an interesting overt and covert story answers to those questions be referred back to evaluate merits
in evolutionary tenns. Among the 3000 cricket species, many of a particular model or, if need be, to modify or even discard
are isolated by their geography and habitat. When a number an existing theOf)'. Disciplined empiricism requires a theory,
of species occupy the same habitat, then temporal, ethological, however infonnal or preliminary it may be or however
or mechanical isolating mechanisms maintain species integri­ difficult an investigator may find testing assumptions stem­
ty. Thus, one species will chirp at night and another during ming from it.
the day (temporal isolation). If more than one species occupy The appreciation of what ethology is about is more
the same habitat and "sing" at the same time, then the differ­ meaningful ifone is reminded of the early intellectual anteced­
ences in the pulse rate (ethological isolation) maintain species ents of present-day ethology. The clash involving an empha­
identity. Acoustic signalS and communication serve in the sis on laboratory-discovered facts as contrasted to naturalistic
prezygotic isolation of closely-related species. observation culminated in three famous debates at the French
The literature on the ontogeny of acoustic communication Academie des Sciences around the year 1830, in which the
in crickets also deserves more attention from behavioral naturalistic evolutionary point of view suffered a profound
scientists than it has received to date. It should be kept in defeat. BaronCuvier had laboratory facts on his side, but as
mind that many insects mature without hearing the signals of we have learned subsequently, by arguing for the immutability
their own species, and that they sense many sounds that have of the species, he was wrong in principle. whereas Geoffroy
absolutely no resemblance to signals that they as mature adults Saint-Hilaire was right in principle without the appropriate
must eventually produce. As Alexander (1968) has pointed facts (Jaynes, \969). The debates contributed to po\arilation
OUI, there must have been intensive selection pressure for between the two camps, with Cuvier's side insisting on the
resistance to irrelevant acoustic innuences and toward flXed laboratory analysis and founding comparative psychology,
relationship between acoustic genotype and acoustic pheno­ while Geoffroy Saint-Hilaire's camp emphasized naturalistic
type. observations and established ethology. Comparative neuro­
Experiments investigating the genetic correlates of physiologist and protege ofCuvier. Pierre Fk>urens, the author
communication signals in several species of crickets orrer of Psychologie Comparee (1864), is credited with developing
further support to this thesis (e.g., Alexander, 1966, 1968; a comparative psychology that synthesized the mechanistic
Benlley& Hoy, 1974; Fulton, 1933). For example, as early as neurophysiological approaches of Des-cartes' human psychol­
1933, Fulton hybridized Nemobius allardi and Nemobius ogy with Cuvier's animal psychology. It is worth noting,
 VOL. 9 NO. I - SPRING 2000 Page 16

however, that during that same year and consistent with the might provide the variations on which natural forces could
intellectual bias of his school, Flourens (1876) published exert selection pressure.
another book, leading French science's attack on Darwin's The major breakthrough and the beginnings of the modem
Origin of Species (1859). The comparative psychology that synthesis surfaced in the 1930s, when R. A. Fisher (1930)
developed in North America around the tum of the century published The Genetical Theory of Natural Selection,
embraced the Darwinian view afthe world, but it remained a Dobzhansky (1937) produced Genetics and the Origin of
laboratory science, and its failure to appreciate the importance Species, followed by Oparin's (1938) The Origin a/Life,
af the ecological·naturalistic dimension of behavior contribut­ Mayr's (1942) Systematics and the Origin a/Species, and
ed to its decl ine (e.g., Lockard, 1971). Huxley's (1942) Evolution: The Modern Synthesis. These
By comparison, throughout the nineteenth century the works brought together diverse areas of human knowledge and
naturalistic bias was advanced by other prominent biologists. inquiry. Organic evolution began to be viewed as a by·
Alfred Giard (1904) emphasized ethology and E. Haeckel product of the chemical evolution of matter and biophysics,
(1898) pushed for "oecology" (presently eoology), then and biochemistry and molecular biology surfaced as the new and
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now defined as the study af the relationships among organisms exciting areas of inqui!)'. The new neo-Darwinian synthetic
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and environments. It is no accident that the more recent theory of organic evolution made sense out of taxonomy. It
pioneers of ethology sought to avoid a dichotomy between explained the fossil record as well as the fibless of adaptations
field and laboratory research, and they succeeded in doing so between organisms and their habitats. The cell theory put
under the conceptual framework of evolutionary theory (e.g" forward convincingly in 1839 by Gennan microscopists,
Eibl-Eibesfeldt, 1975; Hess, 1973; Jaynes. 1969; Lorenz, Schleiden and Schwann, was given a new vision: The cell is a
1981; Schneirla, 1966; Thorpe, 1963; Tinbergen, 1951). Mendelian unit carrying the genetic code of stored variability
that is crucial to evolution and, at the same time, is a
Levels of Organization-Levels of Analysis physiochemical entity obeying the laws of physics and
chemistry. The bridge between particle physics and human
Any behavioral problem can be conceived as varying along evolution and ecology was fonned. The door was left open
for the new generation of Nobel laureates such as Watson and
dimensions identified as levels of analysis. Each level can be
defined in tenns of its position on an infonnation continuum. Crick (1953), who, by their elucidation of the double-helical,
physiochemical structure of the DNA mole<:ule and its role in
The major unifying and consensually valid theme in the
ethological perspective is the synthetic theory of organic heredity, provided one of the major empirical validations for
evolution. the new synthesis.
When Darwin and Wallace in the 1850s proposed their Unfortunately, the behavioral sciences were largely left
out of the modem synthesis (Dawkins, 1986; Hess, 1973;
theory of evolution by natural selection of the fittest and by
Lockard, 1971; Lorenz, 1965; Wilson, 1975). The reasons
specific examples demonstrated how these processes could
were many. The pursuit of the mysteries ofHfe focused the
account for the evolution of organisms, they planted the seeds
of the powerful scientific and intellectual conceptualization concerns of the biological sciences on the molecular universe,
that is still unfolding. From Malthus, Wallace and Darwin thereby leaving the behavioral territory to psychology,
knew that organisms reproduced in far greater numbers than sociology, anthropology, and psychiatry). In tum, many
could be sustained by a particular environmental setting. professionals in these disciplines found the nativist, material·
isl, detenninist implications of the modem synthetic thea!)' of
Their observational evidence was that populations remain
relatively constant. They therefore concluded that a large organic evolution to be either irrelevant or difficult to accept
proportion of the offspring must fail to survive. Moreover, and incorporate procedurally, professionally, politically, and
they knew that animals compete for the available resources of personally. For example, until very recently the lack of
the environment and thereby participate in an active "struggle emphasis on the role of hereditary factors in behavior has been
for existence" one of the hallmarks of North American psychology and
(Darwin, 185911869). As Darwin (185911869) indicated: sociology. Thus, many behavioral scientists were swprised by
the "unconventional" decision of the Nobel Foundation in
... owing to this struggle for life. any variation, however slight and 1973 to award the prize for physiology and medicine to three
from what-ever cause proceeding. if it be in any degree profitable to ethologists, Karl Von Frisch, Konrad Lorenz, and Nikko
an individual of any species, in its infinitely complex relations t o Tinbergen, thereby acknowledging the efforts of those
other organic beings and to external natw-e, will tend to the preserva­ individuals in bringing the study of behavior under the
tion of that individual and will generally be inherited by iLS offspring. umbrella of the synthetic theo!)' of organic evolution. With
The ofsf pring, also, will thus have a bett.er chance of surviving for, of the subsequent ad-vent of sociobiology (e.g., Wilson, 1975)
the many individuals of any species which are periodically bom, but
and cultural materialism (Harris, 1966, 1979), the initial
a small number can survive. (p. 61)
surprise gave way to exchanges characteristic of a paradigm
clash (e.g., Cavalli-Sforza & Feldman, 198 I; Gould, 1980;
Even though it was most important for the evolulionary theory
Rose, Lewon-tin, & Kamin, 1984; Lumsden & Wilson, 1981;
that heritable variations be present in each generation, Darwin
Trivers, 1985).
nevertheless freely conceded his ignorance of the me<:hanisms
Current ethology is occupied with four hierarchical
of inheritance. It was not until about 1900 that Mendel was
biological questions and concerns: What are the ontogeny,
rediscovered and that Hugo de Vries proposed his mutation
causation, function, and evolution of behavior? Explanation
theory by pointing out the likely possibility that the obvious
and understanding require that attention be given to each of
morphological changes he observed in the evening primrose
these questions and concerns and to the various levels of
 Page 17 VOL. 9 NO. I - SPRING 2000

interrelalionship among them. The magnitude af the hierar­ The attachment theoty proposed by Bowlby (1969/1982)
chical concerns requires a breadth of synthesis that transcends was developed in an attempt to extend and improve traditional
levels of analysis from genotype to behavior and ecology-a psychoanalytic approaches. The three volumes of Attachment
synthesis that transcends the extremes of levels of biological and Loss (1969, 1973, 1980) provide a modem synthesis that
organization. goes well beyond the modesty of Bowlby's original claims.
In genera� the consensus among ethologists has been that: (a) Nevertheless, Ainsworth's (1969) observation that "In effect
organic evolution has been a by-product of the chemical what Bowlby has attempted is to update psychoana1ytic theory
evolution of matter, (b) animal species, including Homo in the light of recent advances in biology" (p. 998) still rings
sapiens, are the products ofnalUl1ll selectioo, and (c) genes are true. Bowlby's (196911982) synthesis of psychoanalytic
chemically code for phenotypic expressions. In tenns of thought and ethological research was very compatible with the
reproductive success, natural selection favors those animals ethology of the 19505 and 1960s period. Patterns of infant­
whose genes, through their phenotypic expressions, success­ adult attachment were approached from a comparative cross­
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fully interact with the environment afme ecosystem. The species perspective as evolved species-typical behavioral
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above-listed considerations stem from the world view shared adaptations. Bowlby was careful to distinguish between
by ethologists (Dawkins, 1986). Even so, some considemtions "teleological assumptions" under which the purposes of
are often neglected. We now attempt to relate these consider­ behavior are assumed and "teleonomy" under if which the
ations to levels of analysis in the behavioral sciences. contingencies facilitating the survival value of behavioral
The ethological model incorporates in a hierarchical adaptations may be demonstrated. Bowlby attributed the
fashion levels of organization from subatomic particles to evolutionary origins of attachment behavior to predatory
ecosystems. No level of organization or analysis is conceived selection pressures. This conceptualization of human attach­
as more "important" or "adequate" than another, since a ment was articulated within the framework of biopsychosocial
position on the infonnation continuum is not in itself a systems theory (Bowlby, 196911982; Bischof, 1975) that
criterion for importance or adequacy. The reduction of a invites comparisons with levels of organilJltion and analysis
behavioral problem to a neurophysiological one, or of a that we have identified as characteristic of the ethological
neurophysiological one to a biochemical one, does not in itself approach.
generate a more fundamental or a more important explanation If one reviews the methods and practices of present-day
of the original behavioral problem. Surely, we recognize that adherents of the ethological theory of attachment, there is
the water molecule has characteristics and properit es indepen­ found a mismatch between those conceptualizations and
dent of those of hydrogen and oxygen. At the same time, we objectives of modem ethology (as we have elaborated them)
must note that knowing the characteristics of hydrogen and (e.g., Ainsworth, Blebar, Waters, & Wal� 1978; Brethenon &
oxygen does provide us with some important infonnation Waters, 1985; Sroufe & Watm, 1977). In a striking contrast
about water. Thus, it follows that the usefulness and appropri­ to the cricket song example, these contributions are character­
ateness of her particular level of analysis is circum.scribed by ized by the paucity of research on ontogeny, including
theoretical orientation, parameters of the problem under mechanisms of causation, function, and evolution of attach­
investigation, and contextual circumstances, as well as by ment. Considerations of genetic, neurophysiological, neuroen­
general purposes of the discipline or the investigatOr. Thus, as docrine, functional analyses, the latter including such molar
our introductory example indicates, a student in modem processes as perception, preverbal, nonverbal and verbal
ethology investigating the behavioral biology of the cricket communication, and learning, and evolutionary processes of
song would fmd it necessary to acquire at least some sophisti­ attachment are missing or are dealt with superficially.
cation in language and the tools of genetics, newophysiology In contemporary ethology, Darwinian formulations such
and neuroanatomy, quantitative behavioral analysis, as "adaptations for the good of the species" have given way to
systematics, ecology, and evolution. considerations of evolutionary strategies of ultimate causation
and conditional probabilities in proximal development that are
Ethology And Attachment derived from theory and empirical evidence from both
experimental and field ethology, population genetics, evolu­
The attachment behavior of young precocial fowl toward tionary biology, behavioral ecology, and developmental
biologically-appropriate adults or surrogates including psychobiology. Among the researchers of human attachment,
hwnans, was noted two thousand Y"'"' ago (Hess & Petrovich, these developments have received but scant attention. We
1977). Konrad Lorents (1935) paper on companions as
noted in an earlier section that Bowlby (1969/1982) was
careful to distinguish between teleological and teleonomic
factors in the socialization of birds represents the flf'St post­
Darwinian experimental attempt to deal extensively with the assumptions. Admittedly, "adaptive" is a troublesome term
phenomenon of "imprinting" (see Petrovich & Gewirtz, incurring problems of teleology in its use, if ecologic-teleono
Chapter S; for a review of the literature from a historical mic contingencies of survival value are not specified. More­
perspective, see Hess, 1973; Hess & Petrovich, 1977). The over, given the biological history of Homo sapiens, various
more recent interest in human attachment has been sparked by modes of adaptation may be outcomes of specific experiences
the elegant contributions of Bow I by (1958, 196911982, 1973, rooted in learning and tradition as of genetically programmed
1980) and by the derivative approach and refinements of
processes. At some level of analysis, however, the modem
Ainsworth (1969, 1982) and her associates (Ainsworth, view holds that conditional responses are an outcome from the
Blehar, Waters, & Wall, 1978). Among students of human coaction. of these processes (probablistic epigenesis). Even
development, these contributions have come to be known as so, analyses of these processes in a given ecological setting are
the ethological approach to attachment. required. Lack of sensitivity to these issues is noteworthy
 VOL. 9 NO. I - SPRING 2000 Pig, 18

among human developmentalisls investigating attachment Bentley, D., & Hoy. R. R. (1974� The: neurobiology of cricket
under the aegis of ethology. song. Sclefllijic AmericaT/, 23/.3444.
Contempomry ethology is neutral about the relative Bischoff, N. (1975). A systems """""'" lowanIlhc functional
connections of attachmenl or fear. ChJ/d lR' elopmenl. 46. 801-
..
conbibutions of laboratory and field research. The consensual
817.
view holds that the problems of interest to researchers are to
Bowlby, J. (1958). The nalure of thc child's tie to his mother.
be found in nature, A laboratory is a tool that allows the
inlernolJonol Journal o/Psychoonolysis. 39. 350-373.
investigator an opportunity to test specific hypotheses under Bowlby. J. (196911982). A"achmenl and loss Vol I
controlled conditions and investigate experimentally puzzling Allachmenl. New York: Basic Books.
aspects of behavioral development. In tum, laboratory Bowlby. J. (1974). Auachmel/l and loss. Vol. 2. Separalian
solutions are evaluated in (enns of their putative biologicaU alUiel)l (md anger. New York: Basic Books.
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fitness of behavioral adaptations found in nature. By compari· and depression. New York: Basic Books.
son, the application of the Strange-Situation laboratory Bretherton, I. (1985). Allachmem theory: Relros�cl and
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prospeCt. In I. Brctherton & E. Waters (Eds.). Growing points of

procedure for assesis ng attachment has dominated the method­
This document is copyrighted by the American Psychological Association or one of its allied publishers.

attachment theory and research: Monographs o/ the Society/or

ological landscape in the human-development approach to
�seQl'Ch in ChJld /)noelapmem (pp. 3-35), 50. Nos. 1-2. Serial No.
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1984). Iransmlsslon and ellolulion. Princeton, NJ: Princeton University
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For example, Wale..' and Deane·s (1985, p. 42) statement that Darwin, C. (1869). Ort lhe origin ofsp
ecies by fMOIIS ofnatural
"questions about what is learned during the attachment se/«tion. or tiwpresef"o'OliOl'f of/awJl'td I'QCeS in the struggl e/or lifo.
London: John Murray; New York: Appleton.
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re«ived linle attention," is likely to be shared by any etholo­ Dixon, K.A., &. Cade, W. (1986). Some factors influencing
gists interested in molar and ecological dimensions of early ma.le�maIe aggression in the field cricke Gryllus Ittle�r (time of day,
socialization. age, weight and sexual maturity. Animal DeMiliOI'. 34.340-346.
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(1969/1982) attributed the evolutionary origins of anactunent
Journol 0/ Expuimenlal Biology, 43, 139-1 '3.
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Fisher, R. A. (1930). 17w �neticaJ theory of natural se/eclion.
by researchers of human development and by authors of texts
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most plausible ultimate explanation of the origins of attach­ Frisch, K. Von (1967). The dance language and orielflalion of
ment is that behaviors denoting attachment increase the beel. Cambridge, MA: Harvard Unillersity Press.
inclusive fitness of the individual whose mode of reproduction Fullon. B.B. (1933). Inheritance of song in hybrids of IWO
is characterized by intricate patterns of parental investment subspecies of Nemobius /asciatus (Orthoptera). Annols O/IM
(Petrovich & Gewirtz, 1985; Petrovich, Gewinz, & Hess, Entomological $ociel)l ofAmerica. 16. 368-376
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