Journal of the History of Biology  Springer Science+Business Media B.V.

2017
DOI 10.1007/s10739-017-9491-x

The Development of Sociobiology in Relation to Animal Behavior

Studies, 1946–1975

CLEMENT LEVALLOIS
Emlyon Business School
23, Avenue Guy de Collongue
69130 Ecully
France
E-mail: [email protected]

Abstract. This paper aims at bridging a gap between the history of American animal
behavior studies and the history of sociobiology. In the post-war period, ecology,
comparative psychology and ethology were all investigating animal societies, using
different approaches ranging from fieldwork to laboratory studies. We argue that this
disunity in ‘‘practices of place’’ (Kohler, Robert E. Landscapes & Labscapes: Exploring
the Lab-Field Border in Biology. Chicago: University of Chicago Press, 2002) explains
the attempts of dialogue between those three fields and early calls for unity through
‘‘sociobiology’’ by J. Paul Scott. In turn, tensions between the naturalist tradition and
the rising reductionist approach in biology provide an original background for a history
of Edward Wilson’s own version of sociobiology, much beyond the William Hamilton’s
papers (Journal of Theoretical Biology 7: 1–16, 17–52, 1964) usually considered as its key
antecedent. Naturalists were in a defensive position in the geography of the fields
studying animal behavior, and in reaction were a driving force behind the various
projects of synthesis called ‘‘sociobiology’’.

Keywords: Social behavior, Animal behavior studies, Sociobiology, Edward O. Wilson,

Ethology, Comparative psychology, Ecology, J. P. Scott, Stuart A. Altmann

In the immediate post-World War II period, animal social behavior was

investigated by several overlapping scientific communities, at the fron-
tier between biology and psychology. Ecologists, ethologists, and
comparative psychologists represented distinct research traditions,
ranging from naturalists to white-coat laboratory workers and black-
board mathematicians – leading to widely divergent representations of
how ‘‘social’’ behavior in animals was to be accounted for. Working in
very different settings, both indoor and outdoor, these students of
animal behavior interacted via key individuals and episodic debates,
alternating between conflicting attitudes and calls to reach a synthesis.
 CLEMENT LEVALLOIS

Against this background, Edward O. Wilson’s sociobiology, while

similarly devoted to ‘‘the systematic study of the biological basis of all
social behavior’’ (Wilson, 1975, p. 4), is often conceived by the histo-
riography as a relatively separate and later development. The goal of
this paper is to flesh out the relation between sociobiology and the
contemporary established fields studying animal social behavior in post-
WWII America.
The result is a narrative where the history of sociobiology is woven
with the history of ecology, ethology and comparative psychology. We
identify the naturalists (and their concerns about animal behavior
studies performed in the field) as a driving force in what became the
sociobiological synthesis. Shifting attention to the various issues the
naturalists’ debates animated provides a richer understanding not just
of sociobiology’s formation but also of animal social behavior and of
the subsequent history of post-1950s biology.
An attempt to write the history of sociobiology is bound to be sur-
rounded by ambiguity: is it to be the history of a field, a term, or even a
book? Some reviewers of Wilson’s Sociobiology: The New Synthesis
(Wilson, 1975) suggested that the title of the book accomplished a
performative act. Because it was a new label, with a definition attached
to it, and a 697-page book embodying its principles, a new field called
‘‘sociobiology’’ would have thence been created in 1975.1 Alternatively,
one could consider that sociobiology, being defined by Wilson as the
investigation of the biological origins of social behavior, is a subject
with a history at least as old as Darwin’s Descent of Man (1871) –
rendering the history of so large a topic very diluted, and the origins of
the label ‘‘sociobiology’’ relatively unimportant.2 One could still choose
to see sociobiology as mere réchauffé of late nineteenth, early twentieth
century ‘‘social Darwinism,’’ because of its potential or actual conser-
vative implications (Cherry, 1980; Lerner, 1992). However, most usually
1
‘‘Then one day here rose up a man from among [natural historians]. He had been
called Entomologist, Ecologist, and even Biochemist. But that was not enough. All grew
quiet as he raised his golden pen: ‘There shall be a new science,’ he said, ‘and it shall be
called SOCIOBIOLOGY’’’ (West Eberhard, 1976, p. 89). ‘‘A subject called sociobiology
exists by virtue of the Cartesian syllogism, ‘I have a textbook written about me,
therefore I am a discipline.’’’ (Samuelson, 1975). ‘‘A biochemist recently asked me to
define sociobiology. The only simple answer to the question was ‘The branch of biology
covered in E. O. Wilson’s book.’’’ (Krebs, 1976, p. 709).
2
‘‘Wilson wrote the first modern synthesis of the accumulated literature on behav-
ioral ecology and its related ‘‘skin-out’’ fields, especially ecology and evolutionary
theory, and he promulgated the label ‘‘sociobiology,’’ but that doesn’t make him the
field’s father/founder. Who was? Darwin.’’ (Stuart Altmann, email to the author, March
3, 2004).
 THE DEVELOPMENT OF SOCIOBIOLOGY IN RELATION

concur with Ullica Segerstrale’s sociological account of the sociobiology

debate, which considers that the ‘‘deep background’’ of sociobiology
lays in the mid-1960s with William Hamilton’s and Robert Trivers’s
articles on the evolution of altruism (Hamilton, 1964; Trivers, 1971,
1974) – a narrative at odds with Wilson’s own view.3
These narratives leave a rich trove of materials unaddressed, worth
incorporating in a history of sociobiology: the development through the
1950s and beyond of several disciplines claiming an interest in the
biological study of the social behavior of animals, paving the way to
initiatives named ‘‘sociobiology’’, well before Wilson’s use of the term.
To the point, we claim that post-WWII animal social behavioral studies
(ethology, comparative psychology and the Chicago school of ecology)
were in a state of protracted conflict over their different traditions of
indoor and outdoor studies. Wilson’s sociobiology, we will argue, can
be profitably reconsidered as one of a series of attempts to resolve those
tensions by defending the naturalist tradition in animal studies.
Recent works examining the field-laboratory border in twentieth
century biology have emphasized the role of localization in the shaping
of biologists’ practices, leading to very different styles of research and
career paths depending on whether they worked in the field, a zoo, a
museum or in the laboratory (Livingstone, 1995; Weidman, 1996; Fin-
negan, 2008; Lightman, 2016, pp. 179–328).4 Typically, since the late
nineteenth century, field biologists struggled to retain a sense of legiti-
macy at a time when standards of quantification and control elevated
the laboratory and experimental science as the imagined future of
biology (Benson, 1991). Naturalists adapted techniques from the labo-

3
On the ‘‘deep background’’ of sociobiology, see Segerstrale (2000, pp. 79–99). In
chapters both entitled ‘‘What is sociobiology?’’, Kitcher (1985) and Alcock (2001) take a
similar view. Wilson flatly denies this historical reading of his work: ‘‘I was inspired to
write Sociobiology not by the impetus of gene-centered theory of altruism as begun by
Williams and Hamilton (and Haldane deserves some credit) although I admired the idea
greatly and was pleased to include their work in a 1965 article on social insects and then
in The Insect Societies in 1971.’’ (Wilson, email to the author, April 16, 2009). This
historical claim by Wilson should be put in the perspective of his recent reconsideration
of the role of kin selection in the origins of eusociality (Nowak et al., 2010; Gibson,
2012).
4
Our framework bears an obvious debt to Kohler (2002), which traces the history of
American (mainly plant) ecology in this perspective, up to 1950. See also Montgomery
(2005) for a focus on primatologist C. R. Carpenter in the first part of the twentieth
century, paying the same attention to scientific practice in situ; Rees (2006) on the
practices of the field of post-1950 primatologists, Burkhardt (1999) for a brief history of
ethology focusing on the problem of place, Billick and Price (2010) on ecology and
place.
 CLEMENT LEVALLOIS

ratory to the use in the field, developed their own tools, and invented
protocols to reinvent the field as a place where ‘‘nature experiments’’
could be performed. These activities which necessitated new capabilities,
such as field stations, transformed the traditional, observational natural
history into ‘‘professional’’ ecology in the early decades of the twentieth
century (Billick and Price, 2010; Kingsland, 2010). We argue that these
tensions continued to play a role in post-1950 biology, and even
heightened due to the coming of age of yet another mode of production
of knowledge in contemporary biology, namely blackboard mathe-
matical biology.
We will first depict the three main traditions of investigation of
animal societies (ecology, comparative psychology and ethology),
examining how they were situated at different places in the geography of
scientific practices in mid-twentieth century, leading to divergent and
equally problematic conceptions of the nature of ‘‘social’’ behaviors,
and the methods to study them. We will then make the argument that
this lack of unifying framework led outdoor biologists, who in this
geography suffered from a recurring deficit of scientific credibility, to
seek closer relations with experimental biology in order to develop an
alliance, that behavioral geneticist J. Paul Scott called ‘‘sociobiology’’.
Once established that Scott’s sociobiology first emerged in the 1950s as a
synthesis of laboratory and field based practices, it will be recognized
that a similar pattern explains the forging of alliances by naturalists with
mathematicians and theoretically minded biologists in the 1960s,
resulting this time in Stuart Altmann’s and then Wilson’s sociobiology.
This will lead us to reconsider in a last section the meaning of Socio-
biology: The New Synthesis, and to suggest that this project led to
reaffirming the naturalist credo in biology.

Troubles on the Lab-Field Border

Until the 1950s, ecologists at the University of Chicago were among the
most visible students of social behavior in animals (Mitman, 1992).5
Drawing on a tradition of attention to the relationships between the
organisms and their environment, students of animal behavior at the
Department of Zoology emphasized the social relations in animal

5
A mathematical approach to animal social behavior was developed at the same
time by the ‘‘Committee of Mathematical Biology’’ of Nicolas Rashevsky in the same
university. Lacking an adjunct empirical research program, it failed to gain much
traction among students of animal behavior (Abraham, 2004; Shmailov, 2016).
 THE DEVELOPMENT OF SOCIOBIOLOGY IN RELATION

communities. The physiological gradients conceptualized by Charles M.

Child, Ralph Gerard’s notion of ‘‘orgs’’ encompassing both living
beings and societies, Alfred Emerson’s attachment to the concept of
superorganism, and Warder C. Allee’s stress on the adaptive role of
cooperation in evolution were as many ways to think of animals not as a
homogeneous collection of isolated individuals, but as integrated
groupings (Mitman, 1992).6 According to the definition of the ‘‘social’’
held at Chicago, the behavior of individual animals was to be explained
as part of a subsuming whole functioning like a coherent system or
organ, displaying physiological properties (Burns, 1980; Banks, 1985;
Mitman, 1992).
Interestingly, those integrated groupings were given different names
and treatments, depending on the ‘‘practices of places’’ (Kohler, 2002)
used to investigate them. Social groups in nature were ‘‘communities’’,
while they were ‘‘populations’’ when studied in the labs, or in the field
but with standards of quantitative rigor meetings those of the lab (Allee
et al., 1949, p. 265). The problem at Chicago was that they tended to
attach as much if not more importance to the field as to the laboratory,
at a time when funding was re-orientated towards a physics-inspired
biology, designating clearly the laboratory as the preferred place to
conduct science.7 Indeed, Emerson, Allee and their colleague Karl
Schmidt were especially keen on field work: ‘‘seasoned tropical biolo-
gists’’, they all ‘‘shared a certain fondness for life in the jungle.’’ (Mit-
man, 1992, p. 103) Allee lamented that he ‘‘want[ed] students with a love
for field work which even the handicap of living in the city of Chicago
cannot quench.’’ ‘‘I am tired of working with man-made, degree-hunt-
ing graduate students in zoology. I welcome the natural naturalist who
cannot be kept out of field work even though his doctor’s or other
research lies elsewhere.’’ (Allee, 1930, p. 630).
This attitude also translated into the debates on the levels of selection
in evolutionary theory. Some among the younger generation of students
reacted negatively to the ‘‘good of the species’’ brand of argument which
was commonplace among naturalists. Based on statistical analyses of

6
Even Sewall Wright, who as a geneticist could have ignored the role of higher level
interactions, stands out for suggesting a first analytical model of group selection – which
buttressed his colleagues’ efforts at Chicago to show that pro-social behavior was a force
of evolution (Wright, 1945).
7
Once appointed Head of the Division for Natural Science at the Rockefeller
Foundation, Warren Weaver promoted a ‘‘New Biology’’, also dubbed ‘‘experimental
biology’’, which eroded the funding available to field biology (Kohler, 1976, pp. 286–
287). ‘‘Molecular biology’’ was coined by officers of the Rockefeller Foundation in their
Annual Report for 1938 (Abir-Am, 1987, pp. 32–33).
 CLEMENT LEVALLOIS

homogeneous populations in labs, rather than in the field, the renun-

ciation to its fitness by the individual for the benefit of a community was
judged a logical impossibility. ‘‘If it was biology Emerson was dis-
cussing, I would be better off selling insurance,’’ recalls evolutionary
biologist George Williams about his attendance to a conference at
Chicago in 1957 (Brockman, 1995, p. 40). In sum, Chicago ecologists
were situated right on the border between the field and the laboratory,
which put them in an unstable position. While they studied populations
in the laboratory as a complement to the investigation of natural
communities in the field, the new biology used the laboratory to cancel
the noise of social interactions and isolate the evolutionary destiny of
homogeneous, asocial populations.
In contrast, the immense majority of comparative psychologists
conducted their science indoor, which spared them the legitimacy issues
faced by field ecologists. But their definition of ‘‘social behavior’’ turned
out to be equally problematic, though in a different way. Whereas
ecologists at Chicago worked in the field and in the laboratory, the
privileged techniques and location of comparative psychology were
quasi-exclusively those of the laboratory, affording a tight control of
environmental parameters and precise measurements. This rigor was
contrasted with the perceived unscientific, subjectivist methods of field
biologists, whose outdoor activities could allow neither the full
description nor the reproducibility of observations.
Most contributions in comparative psychology reported experiences
of individual animals engaged in exercises of maze-learning or sensory
discrimination under various stressful conditions. Individual behavior
was understood as a manifestation of the subject’s inner physiological
drives, susceptible of modification through physiological manipulations
and conditioning (via reinforcement or punishment). In the relatively
rare instances when social behavior was under scrutiny, comparative
psychologists proceeded by replicating their methodological approach
to the study of single animals.8 This approach to social behavior, by
comparing the ‘‘normal’’ social trait with highly perturbed states, was
conducive to a peculiar understanding of sociality and behavior. Indeed,

8
In the early days of comparative psychology, the investigation of social behavior
was included in the agenda the discipline traced for itself (Holmes, 1922). The clarifi-
cation that ‘‘minding’’ (we would say ‘‘cognition’’ today) had always been the actual
matter of interest, rather than ‘‘behavior’’ in full, is stated in a rare defense of the classic
program of comparative psychology by William Mason (Mason, 1980). On average,
between 1921 and 1974 less than 10% of the articles published in the Journal of Com-
parative Psychology were concerned with social behavior. For similar bibliometric
surveys, see Schneirla (1946); Beach (1950); Lown (1975).
 THE DEVELOPMENT OF SOCIOBIOLOGY IN RELATION

comparative psychologists regularly expressed their wish for sounder

theoretical motivations to their scientific enterprise, looking at ethology
alternatively as a potential inspiration for reform, and as a threatening
competitor.9
In comparison with the experimental approach to the study of social
behavior privileged by comparative psychologists, the defining feature
of ethologists has sometimes been said to be that they loved the animals
they studied (Dewsbury, 1992, p. 209; Burkhardt, 2005, p. 312).
Ethology achieved international scientific standing in the post-war
period with the works of Dutch Niko Tinbergen and Austrians Karl von
Frisch and Konrad Lorenz. Contrary to comparative psychologists,
ethologists did not try to identify the physiological origins of social
behavior with laboratory experiments, followed by dissections and
physiological analyses. Instead, they claimed the merits of considering a
rich variety of species and the importance of spending long hours at
animal watching, so as to surely identify the original, ‘‘untouched’’
behaving of animals. Focused on the full range of behavior rather than
on the functioning of high level cognitive abilities, ethologists down-
played the importance of learning and insisted on the instinctual, innate
patterns of behavior.10
Natural or semi-natural settings obviously allowed for an increased
attention to social behavior. In open fields and with free movements,
phenomena like imprinting, gregariousness, mating, conflict or coop-
eration were more readily observed than by confining single animals in
mazes or Skinner boxes.11 Yet, the import of ethology in the United
States did not go without concerns for its scientific credentials. Com-
parative psychologists were quick to point to their relative strength in
quantitative methods, and to dismiss the ethologists’ too strong insis-
tence on the innate nature of animal behavior, to the detriment of an
attention to the development and learning stages, necessary for the
9
The often-cited example of absurd confusion between natural and artificial settings
is the mention by a comparative psychologist of the ‘‘natural cage conditions’’ (emphasis
added, Burkhardt, 1999, p. 500). A list of calls to reform comparative psychology by
insiders includes (Schneirla, 1946; Beach, 1950; Hodos and Campbell, 1969; Chiszar,
1972).
10
On the ethology and comparative psychology traditions and the parallel divide
between naturalist and laboratory traditions, see Jaynes (1969). To Jaynes, the gist of
the difference is that naturalists study animals ‘‘not as corpses reeking with formalde-
hyde in the Cuverian tradition, but as living things in their habitat.’’ (on p. 602).
11
Comparative psychology did develop ‘‘open-field’’ experiments in the 1960s, but it
featured individual animals raised in captivity, for which the ‘‘open field’’ (a measured,
artificial perimeter) appeared as an unusual and frightening experience (e.g., Candland
and Campbell, 1962).
 CLEMENT LEVALLOIS

individuals to acquire their full pattern of behavior.12

This categorization of animal behavior studies circa 1950 in three
currents (ecology, ethology, comparative psychology) has sometimes
being criticized for its arbitrariness, since a number of individuals tried
to render them interfecund, leading to exchanges of concepts, tech-
niques, and to common publications (Dewsbury, 1992). Some com-
parative psychologists did conduct field studies, and some ethologists
had a preference for observations in semi-natural conditions, rather
than in the complete wilderness.13 We contend here that distinguishing
the features of these fields, while acknowledging the influences and
overlap between them, does not impede our comprehension of animal
social behavior studies at the time. To the contrary, identifying the
tensions between these fields, in particular the long standing competi-
tion for scientific credibility between naturalist and laboratory-based
traditions, is key to the understanding of the recurrent calls to synthesis
in animal behavior studies in the 1950s and beyond. Often presented as
idiosyncratic projects (as in ‘‘Wilson’s sociobiology’’), we want to re-
present them now as responses to a common need: building up legiti-
macy by making alliances between the laboratory and field research
traditions.

The Committee for the Study of Animal Societies Under Natural

Conditions

The first use of the term ‘‘sociobiology’’ in the post-war period nicely
illustrates the attempts for the three traditional fields investigating the
social behavior of animals to communicate on a common platform. In
12
This different appraisal of the instinctive nature of social behavior reflected the
established consensus in the post-war American society that ‘‘culture’’ was irreducible to
biological explanations – to argue the contrary was risking being accused of social
Darwinism (Bellomy, 1984). In this respect, social behavior in animals occupied an
ambiguous position: was it as strongly determined by Darwinian evolution as physical
characters were, or did it belong to a different order of phenomena entirely, as in an
embryo of culture? Behaviorists and comparative psychologists considered that envi-
ronmental factors played a larger role, and were suspicious of the determinism implied
by the evolutionary view. These divergent views were made fully apparent in the criti-
cism of Lorenz’ ethology by comparative psychologist Daniel Lehrman (Lehrman, 1953;
Burkhardt, 2005, pp. 384–390).
13
As Burkhardt notes, Lorenz studied semi-domesticated animals in the enclosed
fields adjacent to his father’s large house in Altenberg, Austria. Zoos were other priv-
ileged artificial settings frequented by naturalists (Mitman, 1996; Burkhardt, 1999,
2005).
 THE DEVELOPMENT OF SOCIOBIOLOGY IN RELATION

1946, a conference sponsored by the Rockefeller Foundation was con-

vened at the Jackson Memorial Laboratory, Bar Harbor, Maine, under
the supervision of behavioral geneticist J. Paul Scott (1998). Scott had
the profile to be a go-between for the three fields. A former student of
Allee and Wright at Chicago, he had demonstrated an interest both for
field and laboratory studies. Working on organisms ranging from fruit
flies to mammals, and director of behavioral studies in the national
center for the production of laboratory rats, he was well-prepared to act
as a synthesizer of various approaches to animal behavior studies. In-
deed, if the conference on ‘‘Genetics and Social Behavior’’ was supposed
to launch a Rockefeller project to demonstrate the genetic basis of
intelligence, Scott and the participants ‘‘hijacked’’ the conference and
made it deliver an unexpected outcome (Scott, 1985, 1989).14
The audience gathered a sample of scientists from ecology and
comparative psychology, sympathetic to a dialogue of their fields with
ethology. Hence, the comparative psychologists in attendance were
notable for having expressed an unusual interest in the study of animal
behavior under natural conditions, rather than in their familiar labo-
ratories. Robert Yerkes, who chaired the conference, was the towering
figure of comparative psychology, and came from a time when the
discipline used many more animal models, in a variety of settings.15 C.
R. Carpenter, a former fellow of Yerkes’s ‘‘monkey farm’’, was also
among the audience. He was praised by Scott who had an enormous
respect for his work on the howling monkeys, stressing that it was ‘‘the
first major study of mammalian society under natural conditions.’’
(Carpenter, 1934; Scott, 1985, p. 414). The audience also comprised two
other figures in comparative psychology, both in the forefront of the
criticism of the overwhelming position of laboratory studies in their
field. Frank Beach, Curator of the Department of Animal Behavior at
the American Museum of Natural History until this year, was receptive
to Tinbergen’s ethological approach to animal behavior studies, and
would address a scathing criticism of comparative psychology as per-

14
Founded in 1929 by C. C. Little to study the heredity of cancer, the Jackson
Laboratory had become one of the main suppliers of inbred mice for cancer research in
the United States. It remains an important center in mammalian genetics to this day
(Mitman, 1992; Paul, 1998; Rader, 2004).
15
Yerkes had founded in 1911 the Journal of Animal Behavior, first American journal
devoted to the study of the topic. The journal was discontinued in 1917, to reappear in
1921 as the Journal of Comparative Psychology.
 CLEMENT LEVALLOIS

Figure 1. The fate of comparative psychology, envisioned by Beach. Note the T-maze
and the running wheel. From Beach (1950, p. 118). Published by APA, reprinted with
permission

formed by its flagship journal, that he suggested should be renamed

‘‘The Journal of Rat Learning’’ (see Figure 1).16
T. C. Schneirla was the other prominent comparative psychologist
present at the conference with also a strong interest in field studies.
Specializing in ants, he had co-authored a classic textbook in compar-
ative psychology replete with studies of rat maze learning, but had also
published an acclaimed study of army-ants, observed in the field in
Barro Colorado Island in 1932 and 1936 (Schneirla, 1933, 1938; Maier
and Schneirla, 1935; Tobach, 2000).17
Reflecting the participants’ pluralistic approach to the study of social
behavior, the Bar Harbor conference called for a conjoint development
of field and laboratory studies. An outcome of the conference was the
creation of a ‘‘Committee for the Study of Animal Societies under
16
In 1947, Beach met with Tinbergen who was visiting the United States. The six-
lecture series given by Tinbergen at the American Museum and Columbia University
would constitute the material for The Study of Instinct, published in 1951 (Burkhardt,
2005, pp. 292, 362–366).
17
Barro Colorado Island, together with Cayo Santiago Island (see below in the text),
have been major centers for the study of animal social behavior and for the socialization
of the scientists involved in those studies. Allee spent 3 months on Barro Colorado in
1924, Emerson 6 months in 1935, and Schmidt some time circa 1930 (Schmidt to
Schneirla, March 1, 1948, Schneirla Papers, Box M579, Folder ‘‘Business Support for
Research’’, Theodore C. Schneirla Papers, Archives of the History of American Psy-
chology, The University of Akron, hereafter cited as ‘‘Schneirla Papers’’).
 THE DEVELOPMENT OF SOCIOBIOLOGY IN RELATION

Natural Conditions’’ (CSASNC) to translate this goal into practice. By

wedding the best practices of comparative psychologists to those of the
naturalists, the result would be a synthesis ‘‘between the fields of biology
(particularly ecology and physiology) and psychology and sociology.’’
‘‘Many names have been given to it, but perhaps the best and most
descriptive is ‘sociobiology’,’’ Scott indicated at the first conference
organized by the CSASNC (Scott, 1950, pp. 1003–1004).
This blend of practices would need a suitable place to flourish, at
mid-range between the laboratory and the complete wilderness. Island
biological stations fitted these constraints. Straightforwardly, insularity
meant that one of the great features of the laboratory, isolation, was
reproduced in preserved natural settings – without the artificiality of
man-made walls and cages. Precise census work could then be per-
formed, opening the way to demographic measurements and controlled
manipulations of the structure of social groups. Hence, the CSASNC
urged that Barro Colorado Island should ‘‘be put on a permanent basis
of support as a center of tropical research’’ (Scott, 1947, p. 208). Islands
would also play a crucial role in the genesis of Altmann’s and Wilson’s
versions of sociobiology.18
The CSASNC acted as a provider of resources for the informal network
of naturalist minded students of animal social behavior. Through a typed
newsletter sent regularly by Scott, CSASNC members and ‘‘correspon-
dents’’ were told about available grants, places suitable for naturalistic
study, and job openings.19 The newsletters also performed an organiza-
tional function, several including a listing of members and correspondents,
or serving as call for papers and programs for the yearly meetings of the
CSASNC. Sub-committees were created to collect a bibliography on field
instruments, or to encourage field trips to Barro Colorado Island. In 1948,
the first conference of the CSASNC in New York led in 1950 to the pub-

18
Barro Colorado Island was formed by the flooding of a valley, at the creation of the
Panama Canal. It was transformed into a natural reserve in 1923. It is then a perfect
example of the man-made character of the ‘‘natural’’ places singled out by sociobiol-
ogists. On the convenience of islands for census work on ants, see Schneirla to Dia-
mond, October 20, 1953, Box M585.2, Folder ‘‘Public relations’’, Schneirla Papers.
19
Six newsletters were issued between June 10, 1949 and November 1956: Box M579,
Folder: ‘‘Committee for Study of Animal Behavior’’, and in box M585, Folder
‘‘CSASUNC’’, both in the Schneirla Papers. The newsletter (n.d.) including the minutes
from the CSASNC meeting in New York December 29, 1949 lists 28 members (elected
to the committee) and 56 correspondents (simply receiving the committee reports and
news).
 CLEMENT LEVALLOIS

lication of papers on the ‘‘Methodology and Techniques for the Study of

Animal Societies,’’ edited by Scott. This collection provided the first ele-
ments of a synthesis between laboratory and field studies. Field workers
showcased an updated toolbox for outdoor studies, demonstrating a con-
ciliatory attitude towards laboratory studies.20 In Scott’s perspective,
sociobiology was to be the organizational vehicle through which naturalists
could gain a renewed visibility for field work, at a time when they felt
academic journals were becoming less open to this type of research:
All of this [the activities of the CSASNC], I think, will tend to work
in the direction of both building up interest in animal behavior and
sociobiology and thereby making it possible to do what we talked
about at the [New York] conference, namely to organize a new
journal for this type of work. In spite of the cordial attitude of the
editors of various existing journals, my own experience and that of
others who have worked in the field leads me to believe that it is
often very difficult to fit papers to the individual requirements of
these journals.21
Hence sociobiology, as it was first coined and developed by the
CSASNC, arose as a form of appeasement to contemporary tensions
between field and laboratory studies in animal social behavior studies. It
was adamant that the naturalist tradition was an asset rather than a
hindrance for the scientific study of animal behavior. The task for
sociobiology was precisely to develop a rigorous approach to field
studies in order to match the methodological rigor expected by biolo-
gists working in the laboratory. This striving for an alliance between
naturalists and their colleagues working indoor was at the core of
Scott’s sociobiology, as it would be in Altmann’s and Wilson’s.
Through the late 40s and 50s, the CSASNC adopted a more and
more formal structure and it became a regular feature of the joint an-
nual meetings organized by the Ecology American Society (EAS) and
the American Society for Zoology (ASZ) to sponsor a session on
‘‘Animal Behavior and Sociobiology.’’ In 1956, a ‘‘Section On Animal

20
One attempt was particularly diplomatic, when ecologist John Calhoun suggested
borrowing from comparative psychologist their expertise at controlling conditions, but
also their favorite subject, the rat (Calhoun, 1950). Calhoun would continue to situate
his studies of rat crowding and sociopathy in increasingly controlled and symbolically
charged environments (Ramsden and Adams, 2009).
21
Scott to Schneirla, July 9, 1948, Box M579, Folder ‘‘Conferences: Behavior
Committee’’, Schneirla papers.
 THE DEVELOPMENT OF SOCIOBIOLOGY IN RELATION

Behavior and Sociobiology’’ was eventually created within the EAS,

and it gained more than 300 members in less than a year.22 It is at this
point that the young Wilson established contact with this new current of
animal social behavior studies, via his student Altmann.

Primatology on Cayo Santiago: Sociobiology, Redux

In Fall 1953, 2 years into his graduate studies at Harvard, Wilson had
attended a conference by Tinbergen and Lorenz on an American tour,
who were lecturing on the ‘‘new science of ethology’’ (Wilson, 1994, p.
285).23 Lorenz, the passionate naturalist and animal-lover, made a
decisive impression on the similarly inclined Wilson:
‘‘Then Lorenz came … He was a prophet of the dais, passionate,
angry, and importunate. He hammered us with phrases soon to be-
come famous in the behavioral sciences: imprinting, ritualization,
aggressive drive, overflow; and the name of the animals: graylag goose,
jackdaw, stickleback. He had come to proclaim a new approach to the
study of behavior. Instinct had been reinstated, he said; the role of
learning was grossly overestimated by B.F. Skinner and other
behaviorists; we must now press in a new direction. He had my com-
plete attention. Still young and very impressionable, I was quick to
answer his call to arms. Lorenz was challenging the comparative
psychology establishment. … My thoughts now raced. Lorenz has re-
turned animal behavior to natural history. My domain. Naturalists, not
psychologists with their oversimple white rats and mazes, are the best
persons to study animal behavior.’’ (Wilson, 1994, pp. 285–287, italics in
the original).
22
See ‘‘Newsletter’’ by Scott, November 1956, Box M585, Folder ‘‘CSASNC’’,
Schneirla Papers. In 1958, the ASZ would also burgeon a ‘‘Division of Animal
Behavior’’. By 1964, members from this Division would join those of the Section On
Animal Behavior and Sociobiology to create their own society devoted to the study of
animal behavior: the Animal Behavior Society. See ‘‘The Animal Behavior Society: Its
Early History and Activities’’, Schein, May 1976, Box M3796, Folder ‘‘ABS: History
and Origins’’, J. P. Scott Papers, Archives of the History of American Psychology, The
University of Akron. The society kept true to the naturalist ethos that had inspired the
CSASNC: ‘‘It seemed most important that the society should never lose sight of its basic
unifying theme that all really relevant research on animal behaviour must eventually be
related to the natural conditions under which the behaviour has evolved. We therefore
wrote into the constitution the statement that ‘The Editor of the Journal shall encourage
balances publication of both field studies and laboratory studies having fundamental
relationship to the natural life of animals.’ ’’ (Collias, 1991, p. 627).
23
This series of conference has had a considerable impact not just on Wilson, but on
the American public in general (Emlen, 1996, p. 178; Burkhardt, 2005, pp. 370–407).
 CLEMENT LEVALLOIS

Wilson was not aware that his feelings against ‘‘rats and mazes’’ com-
parative psychology and behaviorism, or his enthusiasm for naturalist
studies, were particularly close to those of the CSASNC members.24 But
he did not remain isolated for long. In 1955, Wilson had just finished his
Ph.D. dissertation on the ant genus Lasius when the Harvard admin-
istration asked him to supervise a post-graduate student, 1 year younger
than him. This student, Stuart Altmann, had specialized in the study of
social behavior of rhesus macaques, and had thereby developed strong
connections with the primatologists involved in the CSASNC.25
Carpenter had given Altmann the census data of another colony of
rhesus monkeys he had implanted himself in the late 1930s on Cayo
Santiago Island, off the coasts of Puerto Rico. In the 1940s, the bio-
logical station of Cayo Santiago had suffered a lack of funds, inter-
rupting census work while the population of monkeys was decimated by
shortage of food and removals for civil and war-related research
(Rawlins and Kessler, 1986, p. 29). In 1956, behavioral research on
social primates resumed with Altmann traveling to Cayo Santiago, his
doctoral supervisor Wilson accompanying him:
‘‘My interest in sociobiology was not the product of a revolution-
ary’s dream. It began innocently as a specialized zoology project one
January morning in 1956 when I visited Cayo Santiago, a small
island off the east coast of Puerto Rico, to look at monkeys… The
two days Stuart [Altmann] and I lived among the rhesus monkeys of
Cayo Santiago were a stunning revelation and an intellectual
turning point… In the evenings Altmann talked primates and I
talked ants, and we came to muse over the possibility of a synthesis

24
‘‘Unfortunately, [Wilson] and I had very little contact prior to the publication of his
1975 volume. People kept telling me about a bright young man working with social
insects. But entomologists are a breed apart from other biologists; the insect world is
their universe, and Wilson never came to the Animal Behavior Society Meetings.’’
(Scott, 1985, p. 405) This is not an entirely convincing explanation, as Schneirla and
Emerson were entomologists and founding members of the CSASNC, and as Wilson
was named a Fellow of the Animal Behavior Society in 1968.
25
In 1955, Altmann had done howler monkey census work in the biological station on
Barro Colorado Island, where Schneirla had done his renowned field work in the 1930s and
1940s. This also put Altmann in touch with Carpenter, who had done the original census in
Barro Colorado in 1932 and 1933. Collias, another founding member of the CSASNC, had
also done census work on this population of howler monkeys in 1951. The early career of
Altmann and his contacts with Carpenter are accounted in Haraway (1981, 1983, 1989).
 THE DEVELOPMENT OF SOCIOBIOLOGY IN RELATION

of all the available information on social animals. A general theory,

we agreed, might take form under the name of sociobiology.’’
(Wilson, 1994, pp. 308–310)26
The naturalist approach was still at the heart of this envisioned socio-
biology, as Altmann’s starting point was the collection over 2 years of
ethograms and measurement of frequencies of behavioral displays in the
wildlife of Cayo Santiago. However, as in Scott’s case, what would be
constitutive of sociobiology was the wedding of typical naturalist
practices with those from ‘‘indoor’’ biology, as a means to overcome the
deficit of credibility of traditional naturalist studies. Hence, Altmann
did not rely on the notion of ‘‘releaser’’ as he did for the concept of
ethogram, considering that it would introduce ‘‘gratuitous’’ assump-
tions on the sequential orders of behavioral patterns (Altmann, 1965).
He would have them better ascertained from scratch by a sociometric
analysis, demanding skills borrowed from statistics.27

26
Altmann: ‘‘The term [sociobiology] was, for me, a natural extension of the term
‘‘psychobiology,’’ which Robert M. Yerkes, in the Yale Psych[ology] Dep[artemen]t,
had coined, to indicate a (hoped for) rapprochement between the two fields. Scott used
the term four times in his 1950 Foreword to the publication [Annals NY Acad Sci.,
51(6)] of the results of a conference on ‘‘Methods and techniques for the study of animal
societies.’’ Perhaps it was also used [in] chapters in that conference proceedings. The
volume is logged as number 135 in my collection, and so was one of the first reprints
that I owned, and I am fairly sure that I had a copy with me during my two years on
Cayo Santiago. Whether I got the term from there or from the obvious extension of
Yerkes’ term, I cannot say.’’ ‘‘I was, in one sense, the first sociobiologist, in that, at my
request, I was hired as a sociobiologist when I went to work at the Yerkes Regional
Primate Research Center, in 1965. So far as I know, no one else had such a job title at
that time.’’ (Altmann, email to the author, March 3, 2004).
27
The choice of Cayo Santiago for the place where to conduct the sociobiology of
primates was also revealing of the delicate balance to be struck between field and other
(lab) practices, as readily acknowledged by the scientific workers on the island. ‘‘Cayo
Santiago frees investigators from restrictions imposed by problems in the field which
limit long-term observations on groups of wild animals, and which often result in a
superficial view of the complex behavioral interactions that comprise and sustain social
network. … Cayo Santiago offers an effective compromise between laboratory and field
conditions because its free-ranging animals live relatively undisturbed in a seminatural
habitat and are easily seen.’’ (Rawlins and Kessler, 1986, p. 13). Similarly, Barro Col-
orado Island was often presented as a hybrid of ‘‘jungle’’ and ‘‘laboratory’’, especially
when addressing to a lay audience (Allee and Allee, 1925; Coursen, 1956). This hybrid
nature was periodically renegotiated, as when tracts of forest were cut down to make
place for a new laboratory building (see Catharine Reed to Schneirla, June 8, 1956;
Schneirla to Reed, June 22, 1956, Box M585.1, Folder ‘‘Barro Colorado I’’, Schneirla
Papers).
 CLEMENT LEVALLOIS

On top of the blending of different methodological traditions to collect

and establish the empirical validity of field observations, Altmann and
Wilson considered that an improved theoretical framework was necessary
to extend the lessons from this case study of a single animal society to a
comparative study of all social behaviors in animals. Altmann’s own
preference was with a cybernetic approach to communication. Looking
for an elementary language able to univocally describe the wealth of
social networks existing across species, he came to the view that the
theory of information and communication, as developed by Norbert
Wiener, was such a tool (Altmann, 1962, 1965, 1967; Haraway, 1981).28
This signaled an important shift in animal behavior studies, with the field
and the laboratory being now supplemented by an additional legitimate
place of production of knowledge: the blackboard. Conclusions derived
from the development of mathematical models, traced on a blackboard by a
researcher with possibly no prior training in zoology – and more probably
in physics or engineering, would gain greater and greater credit, in a
remarkable and rapid shift from the situation of the last years (Keller, 2002,
pp. 79–89). At first, Wilson followed a route similar to Altmann’s, relying
on the concept of feedback loops and the formalism of information theory
when analyzing mass-foraging in ant societies (Wilson, 1962; Haraway,
1981). But the competitive environment at the Department of Biology at
Harvard seems to have pushed Wilson to find a more definitely authori-
tative tool to achieve a unified theory of animal social behavior.

The Sociobiologist as a ‘‘Mathematician Naturalist’’

Since the 1930s, institutional and financial resources for biological sci-
ence had been increasingly redirected away from natural studies, to fund
what would be called ‘‘molecular biology’’. Warren Weaver, head of the
Natural Science Division within the Rockefeller Division, was instru-
mental in imposing the vision that biology should get its inspiration
from physical sciences. In his view, ‘‘ecology was glorified natural his-
tory.’’ (Mitman, 1992, p. 105).29 This stance, and the incontestable re-
sults achieved by molecular biologists, seriously threatened the future of
28
CSASNC’ sociobiologists could accommodate with Altmann’s framework, as when
John Emlen (ornithologist, participant in the 1946 Barr Harbor conference) adopted the
language of control and communication to describe a ‘‘program of study into the general
sociobiological mechanisms of cultural diffusion and maintenance in primate social groups.’’
(Gordon Stephenson and John Emlen, circa 1971, cited in (Haraway, 1983, p. 188).
29
On Weaver, the Rockefeller Foundation and the development of molecular biol-
ogy, see Kohler (1976, 1991); Abir-Am (1982); Kay (1996).
 THE DEVELOPMENT OF SOCIOBIOLOGY IN RELATION

animal studies in biology departments.30 The atmosphere of these days

at Harvard is nicely crystallized in an oft-quoted anecdote told by
Wilson about the Department of Biology, which also hosted James
Watson, co-discoverer of the structure of the DNA in 1953:
‘‘One day at a department meeting I naively chose to argue that the
department needed more young evolutionary biologists, for bal-
ance. At least we should double the number from one (me) to
two… I proposed, following standard departmental procedure,
that [a specialist in environmental biology] be offered joined
membership in the Department of Biology.

Watson said softly, ‘Are they out of their minds?’

‘What do you mean?’ I was genuinely puzzled.

‘Anyone who would hire an ecologist is out of his mind,’ responded

the avatar of molecular biology.’’(Wilson, 1994, pp. 219–220).
Molecular biology had renewed the faith in reductionism, and made
animal behaviorists studying whole organisms in the field look simply
wrong headed. Complex social forms would be deciphered by looking at
their putative genetic basis, which implied skills in physical chemistry
(and the mathematical foundations it supposed) that most naturalists
lacked. Wilson himself did not feel at ease with mathematics but was
eager to catch up, sensing that in these ‘‘molecular wars’’, naturalist
studies could be saved by (and had no choice but) strengthening their
scientific status by drawing on the prestige of some mathematical
biology (see Figure 2). As he puts it, the alternative was to be relegated
in museums and looked down as mere ‘‘stamp collectors.’’ After his
encounter with Altmann and his reaction against behaviorism, this was
another strong motive to work at a synthesis, aiming at preserving the
naturalists’ credo and upgrading it to the required scientific standards of
the day. Sociobiology would be the comparative study of the biological
roots of social behavior where the wide knowledge of species and
observation skills of the naturalists would be an asset, and a unifying
30
On the conflict between the naturalist tradition and the rising molecular biology,
see Hagen (1999) in systematics, Johnson (2008) in zoology, and Dietrich (1998) in
evolutionary biology. Of course molecular biology is a laboratory and experimental
science, but it draws on physics, chemistry and mathematics in a way that traditional
experimental biology (e.g., morphology) never did. See Haraway (1989, pp. 394–395) for
a similar account of Wilson’s rivalry with Watson and the role it played in the socio-
biology project.
 CLEMENT LEVALLOIS

Figure 2. Wilson pictured working on his mathematics in 1961, symbolically and lit-
erally hanging on the border between indoor and outdoor biology. From Naturalist
by E.O. Wilson (on page 243). Copyright  1994 Island Press. Reproduced by per-
mission of Island Press, Washington, D.C.

theory of social behavior formulated in advanced mathematics expres-

sions – providing an authoritative analytical framework on par with the
scientific status of molecular biology.
The time was ripe for such an alliance between naturalists and
mathematicians. Since the early twentieth century, population ecology
had developed into a mathematical science. Inspired by the successes of
differential calculus in physics and chemistry, mathematical formalisms
had slowly been imported in biology to study problems in demography,
epidemiology, and ecology (Kingsland, 1995). Alfred Lotka (1880–
1947) was an early promoter of this approach, and if he failed to reach
true recognition outside demography in his lifetime, his work inspired
many natural and social scientists in the post-war period.
One of them was ecologist G. E. Hutchinson, whose student Robert
MacArthur became a collaborator of Wilson.31

31
Wilson and MacArthur belonged to a larger informal group of biologists and
mathematicians interested in the mathematization of natural science, comprising Ri-
chard Lewontin, Richard Levins, and Egbert Leigh.
 THE DEVELOPMENT OF SOCIOBIOLOGY IN RELATION

Although [MacArthur] had majored in mathematics at Marlboro

College, and had a conspicuous talent for it, his heart was in the
study of birds. He was a naturalist by calling, and seemed happiest
when searching for patterns discovered directly in Nature with the
aid of binoculars and field guides… As a mathematician-naturalist
he was unique […].
(Wilson, 1994, p. 244, our italics).Hence in Wilson’s memory, what
MacArthur chiefly brought to their collaboration was the ability to think
about ecological problems as a physicist or mathematician, while
retaining at heart an identity of a naturalist. This fitted perfectly Wilson’s
agenda: to bring the physicist’ mechanist metaphor and mathematical
sophistication to the rescue of naturalist studies.
However, models in population ecology focused on interspecies rela-
tionships, and remained silent on the more essential forms of social
behavior observable at a lower, intra-species level. To account for the latter,
Wilson followed a similar strategy: he relied on recent developments in
mathematical modeling by the British naturalist William Hamilton (Se-
gerstrale, 2013), whose mathematical analysis of altruism in social insects
had come to his attention. Hamilton provided a simple rule which rein-
terpreted diverse forms of social behavior among kin as the outcome of a
cost–benefit Darwinian calculation at the gene level, with the benefits
brought to inclusive fitness outweighing the costs to individual fitness borne
by the performer of the altruistic act. This rule was presented using a
maximization principle and sophisticated mathematical formalizations not
typical of a naturalist’ training. It epitomized the values of abstract rea-
soning and generalization by the enunciation of laws in the tradition of
mathematico-physics becoming ever more present in contemporary biology.
Thence, Hamilton’s rule helped reintroduce social behavior in the
series of topics studied by modern biology. This gene-centered view of
sociality also replaced the ‘‘survival of the species’’ brand of argument
that had made Williams gasp. Naturalists had found a new legitimacy in
concentrating their efforts on social groupings; they had reintegrated the
orthodox Darwinian framework.
It is not often stressed that in addition to the presentation of his
mathematical model, Hamilton’s papers contained a lengthy and tech-
nical discussion of its relevance for many instances of social species,
supported by a wealth of observational studies (Hamilton, 1964). The
social aspects of behavior being most easily studied in the field, this
comforted the naturalists in the position of main providers of the
empirical material supporting mathematical models of sociality, and
(especially if they agreed to train in math) as the most able to extend
 CLEMENT LEVALLOIS

these models to a variety of situations. Indeed, beyond the presentation

of elementary principles in theoretical population biology in its first
chapters, Sociobiology provided an empirical validation of these prin-
ciples by illustrating them with an encyclopedic survey of observational
studies of social species.32
Wilson followed a similar approach when he included the notion of
evolutionary stable strategies (ESS), recently developed by George
Price and John Maynard Smith (in close relation to Hamilton), among
the mechanisms of social evolution presented in the first chapters of
Sociobiology (Maynard Smith and Price, 1973; Harman, 2010; Wilson,
1975, p. 129). An ‘‘evolutionary strategy’’ designates a highly stylized
representation of a type of animal behavior in relation to conspecifics
or to the broader environment, limited to a small repertoire of options
(such as fight or flight) with associated payoffs, measured in terms of
fitness. It is denoted as ‘‘stable’’ if the individuals adopting it and
largely present in a population cannot be replaced by individuals
adopting a different strategy. Drawn from game theory, an interdis-
ciplinary theoretical framework developed since the 1940s by mathe-
maticians, economists and biologists to model social interactions
(Erickson, 2015), the notion of ESS facilitates the study of how
populations composed of animals adopting competing types of
behaviors evolve, through the resolution of mathematical models or
through simulations. These advantages come at the price of a de-
contextualization of the behaviors under study, and a difficult trans-
position of the assumption made on the rationality of the human
players to a game played by animals.
The un-natural assumption of rationality led even mathematically
inclined biologists like Richard Lewontin and Richard Levins not to
pursue their initial interest in game theory for animal behavior studies
(Erickson, 2015, pp. 221–222). In contrast, Wilson provided a stage in
Sociobiology for a dialogue between game theoretic and naturalist ap-
proaches to animal social studies. Game-theoretic models need the
scaffold of field observations, both for the calibration of their param-
eters (definition of the types of behaviors and of the payoff matrix) and
to gauge their external validity, as witnessed by the references to a
number of naturalist studies in the original paper on ESS by Maynard
32
The visuals of the book bring testimony of the importance of the naturalist tra-
dition in Wilson’s sociobiology. The wildlife illustrator Sarah Landry ‘‘went beyond the
effort required for an ordinary book on animal behavior, traveling to zoos and aquaria
to sketch captive animals and visiting herbaria to render in detail the plant species found
in natural habitats of the animal societies.’’ (Wilson, 1994, p. 324). In essence, the
illustrator tuned her work to the naturalist’s spirit of the book (Myers, 1990).
 THE DEVELOPMENT OF SOCIOBIOLOGY IN RELATION

Smith and Price. The fact that game theory was integrated in a synthesis
of animal social behavior studies under the authority of naturalists
bolstered the relevance of the latter in the competitive landscape of
contemporary biology.

Conclusion

Sociobiology: The New Synthesis has been extensively studied for the
debate it initiated on the relationships between natural and social sci-
ences. This concentrates the historical significance of Wilson’s book on
one type of synthesis which the book claimed to achieve: the sociobiol-
ogy of all social animals, from ants to human beings, developed in its
twenty-seventh chapter. In this study, we put the focus on a different
kind of synthesis pursued by Sociobiology, recasting it in the continuity
of two scientific projects bearing such a name, which developed in post-
war United States. We argued that animal social behavior was investi-
gated by ecologists, ethologists and comparative psychologists, each
characterized by different practices of places. In these fields, researchers
with an inclination for the collection of observations in natural condi-
tions were feeling an increasing pressure to assert the legitimacy of
outdoor studies, perceived not to meet the standards of rigor of modern
science. Scott’s CSASNC, which morphed into the Section On Animal
Behavior and Sociobiology (forerunner of the Animal Behavior Society),
was an organizational effort explicitly directed at promoting studies of
animal behavior wedding techniques and protocols from the laboratory
and the field. Altmann developed his own sociobiology project while
observing primates for two years on Cayo Santiago Island, a type of field
station which the CSASNC had called to develop for allowing obser-
vations blending natural and controlled conditions. Altmann’s own
version of sociobiology was still rooted in naturalist studies of animal
behavior, but this time the deficit in legitimacy was not so much felt in
relation to superior laboratory methods, than in regard to the reduc-
tionist program of molecular biology, which was so dominant in the
immediate post-war period, especially at the Department of Biology at
Harvard where Altmann was conducting his PhD under the sponsorship
of Wilson. In this context, Altmann’s sociobiology developed by com-
bining extensive field studies with a heavy emphasis on the mathematical
analysis of information and communication in animal groups.
In the light of these two preceding episodes, Wilson’s sociobiology
appears to follow a recurring pattern: the development of a defensive
alliance between the naturalist approach to animal behavior studies and
 CLEMENT LEVALLOIS

new forms of scientific endeavor threatening this naturalist tradition.

While Scott was most concerned with laboratory settings replacing
studies under natural conditions, Wilson risked to be sidelined by
molecular biology, which favored sophisticated theoretical develop-
ments and a reductionist analysis of phenomena. Embracing these new
developments by discussing them on par with classic studies of animal
societies, as illustrated with the centrality of mathematical models of kin
selection in Sociobiology, enhanced the relevance of the type of
knowledge produced by naturalists: centrality of field work, attention to
levels of social interaction at the group and interspecies level, and the
study of evolutionary dynamics in broad, natural ecosystems.33
In this frame, the historical meaning of sociobiology can be recon-
sidered: in addition to the ‘‘bombshell’’ effect of the book Sociobiology,
sociobiology as a series of post-war projects also represented a more
discrete but effective attempt to reaffirm the relevance of the naturalist
tradition in animal behavior studies and biology at large.

Acknowledgements

I wish to thank two anonymous reviewers for useful suggestions. I

gratefully acknowledge the kind assistance of Lizette Royer on the ar-
chives of Frank Beach, J. Paul Scott and T. C. Schneirla at the
University of Akron. The intellectual stimulation of Philippe Fontaine
is deeply appreciated. All remaining errors are mine.

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