"It Felt More like a Revolution." How Behavioral Ecology Succeeded Ethology, 1970-1990.

As soon as ethology's status diminished in the early 1970s, it was confronted with two successor disciplines, sociobiology and behavioral ecology. They were able to challenge ethology because it no longer provided markers of strong disciplinarity such as theoretical coherence, leading figures and a clear identity. While behavioral ecology developed organically out of the UK ethological research community into its own disciplinary standing, sociobiology presented itself as a US competitor to the ethological tradition. I will show how behavioral ecology took the role of legitimate heir to ethology by rebuilding a theoretical core and an intellectual sense of community, while sociobiology failed to use its public appeal to reach disciplinary status. Meanwhile, ethology changed its disciplinary identity to encompass all biological studies of animal behavior.

Disciplinary Identities: People, Places, Publications

“Many of us have been surprised at the unconventional decision of the Nobel Foundation to award this year's prize […] to three men who had until recently been regarded as ‘mere animal watchers,’” began Nikolaas Tinbergen his Nobel Prize Lecture, delivered in Stockholm on 23 December 1973. Fellow ethologists saw the award of the Nobel Prize in Physiology or Medicine to Konrad Lorenz, Nikolaas Tinbergen, and Karl von Frisch as honoring them for creating the discipline of ethology, celebrating it as a fresh and distinctly biological perspective on animal behavior that far surpassed the subjective impressions of “mere animal watching” in the past, transforming natural history into proper science. However, as the Nobel Prize outwardly celebrated a strong discipline with charismatic leaders and clear conceptual content, ethology had already lost a lot of its disciplinary strength.

The ascension of ethology to the status of a discipline in the early 20th century and its prime in the 1940s and 1950s were soon memorialized and analyzed by ethologists, ethologists turned historians, and historians of science. There is vast scholarship on how ethology was purposefully established as a discipline, the influence and idiosyncrasies of its founding figures, and the distinct approach it took to animals as its subjects. Richard Burkhardt examined it from a disciplinary perspective, asking: “How did the science of ethology come into being?” However, the disciplinary trajectory of ethology in the last quarter of the 20th century has received less interest. This paper examines a period when the field of animal behavior studies became more crowded and ethology experienced the threat of disciplinary competition and asks: How did the discipline of ethology move out of being?

It is not possible to answer this question by exclusively looking at ethology. There is agreement on two points regarding ethology's decline: It occurred by the mid‐1970s and involved the rise of two other disciplines, sociobiology and behavioral ecology. The conventional wisdom goes that ethology was replaced by either sociobiology or behavioral ecology, or both. This is a compelling narrative, as both sociobiology and behavioral ecology have their roots in ethology and animal behavior. However, there is a more complex relationship between these disciplines than a simple matter of succession. This complicated process of disciplinary reconfiguration has received little attention and the changing tides of these disciplines, their core theoretical and methodological content, and their personnel, as well as how these materialized in their publication structure have yet to be analyzed. This paper aims to close this gap in historiography while following a disciplinary perspective approach where disciplines as a whole rather than individual contributions are examined by focusing my analysis on selected disciplinary publications.

Tracing the development of one or even several disciplines requires taking a bird's eye view of developments. This is done by looking at signature publications as well as their reception within the community from the macro level rather than examining the specific practices or results of any given researcher or group. Burkhardt explores the disciplinary beginnings of ethology along the central categories people, places, practices, politics; my approach here replaces the latter two categories with publications. While people make disciplines, publications make disciplines visible to the larger scientific community. Establishing journals, writing textbooks and monographs, and organizing conferences and publishing their proceedings, therefore, can be understood as strategic moves in achieving disciplinarity. Not all publications are created equal when it comes to disciplinary reflections. Certain types of publications stand out: My focus is on textbooks, as they form the basis of a discipline, as well as long‐running series, proving a thorough line of disciplinary reflection on different themes and trends. Book reviews play an extraordinary part in disciplinary transformation as they allow for more explicit disciplinary reflections and boundary work since a reviewer is invited to place the scholarship into its context. The disciplinary politics of the time are reflected in these publications.

The intertwined histories of sociobiology, behavioral ecology, and ethology are a ripe field for investigating the dynamics of disciplinary rearrangement and competition. There are different ways to define disciplinarity, and even though the scholarship has not settled on one definition, there is a consensus that it involves uniting a set of scholars with a subject matter and a research method. The following section (2.) assesses the disciplinary status of ethology at what seems to be its disciplinary peak. Then, each of the following sections explores how strengthening an element of disciplinarity – social coherence (3.), unifying theoretical core (4.), and advancing through competition (5.) – leads to discipline formation. The final section (6.) answers the question of what happened to ethology and if, indeed, it ceased to be.

An Insecure Discipline: Ethology Beyond the Nobel Prize, 1973–1976

The perception of disciplinary coherence at the time of the Nobel Prize was in stark contrast to the true state of the discipline. Looking deeper into the ethological community, feelings about the state of the discipline were not unequivocally celebratory and were, in fact, more insecure. However, this was not always the case. Ethology had its roots in the 1920s and 1930s and its most impressive successes in the following decades and in its prime, ethologists knew precisely what they stood for: During this time, especially Lorenz and Tinbergen had crafted the image of ethology as a merger of the naturalist tradition with more scientific approaches, theoretical foundations, and exact terminology, providing complex causal mechanisms to explain animal behavior. Central concepts revolved around ideas of innateness and instinct, behavior that animals did not learn. This focus on the innate was used to present ethology as an alternative to the branch of psychology interested in animal behavior and learning: comparative psychology. These two disciplines were thought of as competitors, often based on a caricature of their actual complexities and nuances: Here was American‐based comparative psychology, preoccupied with just one species in the confines of the laboratory, running white rats through mazes, exclusively focused on behavior modification through punishment and reward; there was European ethology, observing instinctive behavior in natural settings, without interventions and dealing with the whole of the animal world. Comparative psychology provided an optimal adversary to early ethology's emerging identity. However, by the time of the Nobel Prize in 1973, the boundaries between ethology and comparative psychology had become porous as core concepts of ethology, such as “instinct”; they were first questioned and then dismantled, especially in the English‐speaking part of the ethological landscape.

Ethologists’ disciplinary insecurity was reflected in two edited volumes that appeared in the 1970s: Perspectives in Ethology (1973), edited by ethologists Patrick Bateson and Peter Klopfer, and, three years later, Growing Points in Ethology (1976), edited by Bateson again, this time with Robert Hinde. All three of them were leading figures in establishing ethology at Madingley Field Station of Cambridge University beginning in the 1950s. Despite the recent Nobel Prize, these volumes did not set a jubilant and self‐confident tone, distancing themselves from ethology's past, stating dissatisfaction with the current status quo and offering little future perspective beyond “encouraging scientists […] to let their hair down a little and write in a less inhibited way.” They seemed unsure of ethology's future potential, or, as Hinde and Bateson phrased it with the metaphor of growth: “As things stand, the roots ethology has put down seem healthy. But are they growing towards soil that will continue to nourish them? Are the shoots heading toward the light?”

In exploring ethology's future, Bateson and Klopfer wanted to leave its past behind as back then, “much of the evidence was anecdotal, much of the thinking intuitive,” and Bateson and Hinde were equally keen to “not to review past achievements or reenact past battles.” This dismissal was mainly directed at Konrad Lorenz. As early as the 1950s, his theory of instinct, a central concept for ethology, underwent scrutiny and criticism that eventually turned into outright rejection. This process started as an external critique from the other side of the Atlantic and disciplinary divide, spearheaded by Daniel Lehrman, an American comparative psychologist, which initially could be attributed to the larger disciplinary rivalry. However, while Lorenz stood firm against Lehrman's arguments, Tinbergen and his colleagues at Oxford and Cambridge embraced both Lehrman and his critique and turned away from early ethological theories. Lorenz perceived this intellectual development as a personal attack and publicly declared his unhappiness with the modern field of ethology in the UK, declaring it had unduly given up on innateness in the introduction to his long‐awaited textbook.

This marked a rift in the development of UK and continental ethology. In the UK, Tinbergen's group had grown more interventionist, so the differences to comparative psychology were lessening. His circle, including Robert Hinde, became open advocates of integrating comparative psychology into ethology; the latter even wrote a book in 1966 that was an explicit synthesis of these two disciplines, which is credited with having “swept away much of the old theorizing by Lorenz and Tinbergen.” Lorenz's status in UK ethology had degraded. While Klopfer and Bateson did not call out Lorenz by name, Arthur Riopelle was direct and brutal in his conclusion that ethology had no use for Lorenzian thought anymore: “This superb collection of essays doesn′t afford much solace to tired old notions. The old Lorenzian notions of causation of behavior and evolutionary functions are passed over with scarcely a glance.”

With Lorenz firmly in the past, ethologists in the 1970s were looking to forge a future path for ethology. However, in their assessment, the current state of the discipline did not provide a satisfactory basis: They contrasted the intuitive thinking of early ethology with the current focus on more rigorous and sophisticated quantitative methodology, which they saw as an overcorrection to the point of stifling the innovative and creative energy of the discipline: “Radical or novel theories must be fully formed if they are not to be laughed to scorn, and as new ideas peep out of the ground in the conceptual flowerbed, it is easy enough to shout ‘Weeds!’ and dig them up by the roots,” wrote Klopfer and Bateson in Perspectives. Hinde and Bateson make a similar point: “Vast mounds of impeccably collected data could easily overwhelm creative thinking and a preoccupation with method could displace a concern for problems.” Cambridge psychologist Michael Morgan agreed with their concern in his reviews: “There is thus a great deal of heart searching and a striving to put the subject on what a mathematician would call ‘a sound axiomatic footing,’” but he is not impressed with the results: “One feels not altogether sure about the value of this search for rigor.”

Older ethologists articulated a tension between, on the one hand, wanting to re‐create the sense of excitement of earlier days, but this time with more scientific rigor and less loose thinking, and on the other hand being overwhelmed by the new increasingly data‐driven and mathematical methodology. Ethology seemed to be stuck, having lost its conceptual identity and doubting its ability to forge a new one. In his review, George Barlow picks up on this: “One does well to ask: Who is an Ethologist? It is hard to say.” Several reviewers also note with astonishment how hollow former adversarial distinctions have become. “The title of the book could equally be Perspectives in Psychology,” writes Riopelle; Beer provides a more vivid description: “Perspective drawing is a way of making parallel lines appear to meet. This book illustrates admirably how the distinction that used to be easy to make between European ethology and American comparative psychology has diminished almost to a vanishing point.” In order to re‐establish a disciplinary identity around a shared but not too complicated theory in the future, the editors suggested that ethology was merely in need of an intellectual energy burst, a forum where scholars can be more speculative to re‐invigorate the discipline as “embryonic ideas which have tremendous potential may not be easily distinguishable from embryonic nonsense.” However, the results were not convincing. While reviewers lauded the volumes for their high quality, it remained unclear “where ethology is headed today.”

Where does this leave ethology in 1973? Both Growing Points and Perspectives made it clear that the honors of the present represented the intellectual past of the discipline. The foundational figures and theories were in the past, and the future direction of the discipline was unclear, with especially the older generation questioning the push for more mathematical approaches. With methodology and theory changing and one of the defining characteristics, the line between ethology and comparative psychology, eroded, the question was urgent: What about exploring animal behavior shaped ethology into a coherent discipline?

A New Golden Age: Young Ethologists at Cambridge and Oxford, 1975–1978

While these publications reflect the weaknesses of ethology in the mid‐1970s – a conceptual identity lost between the overhaul of classical ethology and the spread of more mathematical and quantitative methods – surprisingly, they reveal an ongoing strength of ethology: its strong social identity.

Cambridge social psychologist Martin Richards ruminates on this phenomenon extensively in his review of Growing Points. Like many reviewers of this volume, he notes the lack of conceptual coherence. “The diversity begs the question of the definition and scope of ethology – what makes the ‘growing points’ that are discussed ethology rather than comparative psychology, genetics or developmental psychology among many possible contenders?” and then answers his own question: social belonging. “What is perhaps remarkable about ethology is the way in which workers of such varied interests continue to identify with one another and form a cohesive social group.” Richards himself is familiar with the close‐knit ethological community as he did his Ph.D. under Hinde's supervision at Madingley Station in Cambridge, one center of ethological sociality. Several of these centers were a key factor in creating a sense of community that was strong enough to secure the disciplinary identity through a phase of theoretical insecurity. As Peter Klopfer puts it: “The history of 20th‐century ethology is essentially contained in the reports and records of three institutions,” Lorenz’ Max‐Planck‐Institute in Starnberg, Tinbergen's Animal Behavior Group in Oxford, and the Madingley Field Station at the University of Cambridge.

The Growing Points volume was, in fact, dedicated to one of these central institutions: It consisted of papers presented in 1975 in celebration of the research station in Madingley of the Cambridge department. Madingley became a focal point for UK ethology, a pet project of William Thorpe, founded in 1950 with Robert Hinde as its first director. Thorpe was heavily influenced by Lorenz and Tinbergen and while Hinde was not officially supervised by Tinbergen, he can still be counted as a Tinbergen student. Thorpe and Hinde were instrumental in establishing ethology with an ornithological perspective at Cambridge; Hinde and his student Bateson were responsible for creating a focus on neuroscience, physiology, and development at Cambridge that stressed complicated interrelations between different ecological and behavioral factors. And beyond Cambridge, Hinde's synthetical account of animal behavior became the “great blue bible” of ethology.

The anniversary of this foundational institution provided ethologists in the 1970s with a chance to strengthen their social ties as well as reflect on the direction of the discipline. Since the theoretical approaches of the participants of the conference at this time were diverse, it was the personal connection that decided who was an ethologist and could contribute to the conversation, as Richards noted in his review:

Despite the fact that the contributions are in no sense a cross‐section of ethologists (all are people who have worked at Cambridge or have been closely associated with the Sub‐Department and, for instance, no continental researchers are represented) the diversity of work discussed is enormous and a summary is impossible in the space available.

Therefore, Growing Points as a conference proceeding, in particular, reflects the social dimension of the discipline. And in this social dimension, ethology at Cambridge was going strong to the point where Klopfer identifies it as the place where “signs of ethological ferment are most in evidence.” Despite Klopfer's assessment of Cambridge as the center for a movement of transforming ethology, Oxford soon overtook Cambridge in relevance. Over the course of the 1970s, it leveraged its social cohesion and strong group identity into leading the charge in both an intellectual and generational change within ethology.

When Tinbergen came to the University of Oxford in 1948, first as a university lecturer and then, as of 1966, as Professor for Animal Behaviour, his focus was twofold: Firstly, supervising students and building a vibrant research community; secondly, shifting ethology towards questions of the evolutionary significance of behavior. Since the late 1960s, Tinbergen became increasingly concerned that there was too much physiological work and too little on evolution and function work, which required intense investment of time and energy out in the field. Tinbergen, himself a keen field naturalist, made it his priority to steer modern ethology toward the study of behavior and environmental pressures, named behavioral ecology. There was allegedly even an informal understanding that Thorpe's and Hinde's Field Station at Madingley was to focus on behavioral mechanisms while Tinbergens's group at Oxford was to focus on behavioral function. Oxford's overzealous adoption of the adaptationist outlook was soon lovingly mocked by Cambridge ethologists, ascribing Oxford the motto: “Never use a causal explanation if a teleological one will do instead.” Even though his own research interests turned increasingly toward human behavior in the late 1960s and did not display the overriding adaptationist focus Oxford became famous for, Tinbergen had firmly established his Oxford research group as a place for connecting evolution, ecology, and ethology. This research group was instrumental in transforming ethology from within over the course of the 1970s.

Who was this group that was ringing in a new era of ethology? The generational and intellectual transformation is best represented by Tinbergen's right‐hand man, Mike Cullen. Cullen had completed his Ph.D. under Tinbergen, married fellow postdoctoral group member Esther Sager , and became “an extraordinary influence on the development of the scientific discipline of ethology.” This assessment is taken from Cullen's obituary in The Guardian, penned by Richard Dawkins and John Krebs, where they also name him “the unsung hero of that golden age.” That this little‐known ethologist has such a prominently penned and placed obituary is a testament to the depth of gratitude the authors felt they owed him.

Cullen increasingly took over the mentoring role from Tinbergen since the mid‐1960s, when Tinbergen became less involved with his students and increasingly focused on his work on autism in children, often interrupted by bouts of depression. Luckily, Cullen was a born mentor for this group of talented youngsters and almost all members of the group mention his enormous influence. He was widely admired for his mathematical abilities and analytical rigor: Tim Clutton‐Brock recalls him as a “continuous source of stimulation challenging us to focus on testing hypotheses rather than on description.” Strict but humorous and extremely dedicated to his students, “he would spend whole days on a student's problems, designing new methods to solve them and coming up with elegant mathematical solutions.”

Cullen's influence led to a shift in approach that is exemplified by an anecdote from Krebs: When choosing a Ph.D. topic, Krebs was interested in focusing on a theoretical question from population ecology. At the time, an important debate was raging between David Lack from Edward Grey Institute of Field Ornithology and Vero Wynne‐Edwards: Was population size limited by external ecological or internal evolutionary factors? Wynne‐Edwards was proposing a group‐selectionist evolutionary argument that population was self‐regulating its size for the good of the species, while Lack was categorically opposed to this view, pointing instead to ecological factors such as food shortages as imposing external limitations on population size. Krebs had seeds of an idea for an alternative evolutionary argument that did not rely on group selection and proposed the project to Lack himself. As Krebs did not know what animal he wanted to work on, Lack introduced him to his group as “This is David Krebs. He wants to do a D.Phil., but he doesn't know what he wants to work on.” While Krebs had a clearly defined problem, this did not count in Lack's eyes, as his approach was based on picking a species first. Krebs concludes: “My approach of defining a problem and not a species did not fit comfortably with the Edward Grey Institute in the mid‐1960s.” Working from a theoretical problem was, however, indicative of Cullen's influence, who ended up supervising Krebs’ Ph.D. project.

Despite Cullen's enormous social impact on the direction of ethology, he did not succeed Tinbergen when he officially retired in 1974 and did not rise to particular ethological fame. Whereas other stages of disciplinary development are visible in signature publications, in Cullen's case, his signature was his lack of publications. This is often cited as a reason he did not succeed Tinbergen , and yet as Krebs and Dawkins put it: “If he had followed what is now a common practice of putting his name on all the papers of his students and co‐workers that he had helped, he would have stood out as one of the most prolific ethologists of his time.” Cullen's influence is typical for a mentor figure, essential to discipline building in giving younger generations a voice without taking credit.

Mike's entire life revolved around student projects. But he never asked for credit, and rarely even put his name on a paper. He was much more involved in the thesis of almost every single student than Niko [Nikolaas Tinbergen] ever was, and without him, the scientific activity of Niko's group would not have been anywhere near the level it reached with Mike.

The Oxford group became a central location for ethological intellectual exchange and discussion. Tinbergen was known for his exacting and demanding questions for speakers and exerted his intellectual influence in colloquia and seminars on their scientific thinking. “In the actual discussions, the younger members tended to be much more hard‐hitting than Niko himself; they were a merciless, highly intelligent, and very articulate audience.” A new generation of ethologists was being raised in the group and their work was soon to be commended for its quality. The later generation of Tinbergen's group included future leaders of the field, such as Richard Dawkins, John Krebs, Nicholas Davies, Hans Kruuk, Colin Beer, Nicholas Blurton‐Jones and Tim Clutton‐Brock. This generation of Tinbergen profited from an extremely stimulating set of guest speakers that presented what was the cutting edge of ethological, evolutionary, and ecological theory. Robert Trivers presented his work on parent‐offspring conflict in the early 1970s, William Hamilton spoke on kin selection, and John Maynard Smith and Geoffrey Parker were there to elucidate their newest work on using game theory in evolutionary biology. As John Davies remembers: “It wasn't necessary to travel to meet stars of behaviour – we simply had to wait in Oxford for them to visit.” Rigorous questioning and examination of guest speakers was par for the course, as was the intensive exchange of ideas between graduate students, taking inspiration from the new and exciting theoretical work.

Davies recalls his time at Oxford in the late 1970s as knowing no art, only artists, meaning that their sense of group identity was based on their shared interests and activities, the new research direction being best described by what people were doing, and only then finding the thematic common ground. With hindsight, one can see that slowly the group identity was forming into a discipline centering around behavior, ecology, and evolution. A decisive factor was the young and dynamic research community that Tinbergen had established in Oxford. By the mid‐1970s, George Barlow noted in his review of Growing Points what had become “ethology's most rapidly growing and potentially most important area: behavior as a mechanism in ecology.” And despite the social continuation of ethology, the theoretical innovations of the younger generation “felt more like a revolution than a gentle gradual metamorphosis.”

A Discipline Emerges: Assembling Behavioral Ecology, 1978

Throughout the 1970s, this new approach to animal behavior, increasingly relying on mathematical models adopted from population ecology and population genetics, more informed by evolutionary theory and population thinking, formed a distinct disciplinary identity that would eventually lead it to overshadow ethology itself. This transformation realized some of the fears of earlier ethologists that complex theory and inaccessible mathematical formulations were taking over the discipline. However, the researchers at Oxford saw these developments as necessary steps to revitalize ethological research. Richard Dawkins’ contribution to Growing Points makes this clear. He identifies the main problem of ethology as not having produced any “great explanatory principles of late.” With this diagnosis and the will to fix the problem, Dawkins was right at home in Oxford. There they shared an air of excitement at the rapid pace of change and felt that they were at the “cusp of a new wave of behavior studies.” Much like earlier ethologists, members of this group were later to credit themselves that the late 1960s and 1970s were truly the time “that led natural history to blossom into a formal science.” Looking back in 1981, Nicholas Davies summarizes behavioral ecology and its progress in the 1970s in the following manner: “During the last decade behavioral ecology has emerged as an exciting field of study, so named because the way in which behavior contributes to survival and reproduction depends on an animal's ecology, the food it eats, its predators, the architecture of its habitat, and so on.”

What catalyzed a lot of this progress was the above‐mentioned debate about the viability of group selection as a means of population size regulation, as proposed by Vero Copner Wynne‐Edwards. Wynne‐Edwards’ arguments for group selection provoked objections in equal measure from ecologists like David Lack, and evolutionary theorists like George Williams. Both argued forcefully against the group selectionist logic that birds controlled their reproduction due to altruistic behavioral tendencies, explaining that such altruism would be mercilessly rooted out by the logic of natural selection. Williams’, Lack's, and William Hamilton's work on this logic became central to the theory of kin selection or inclusive fitness that circulated in the discussions of the Oxford group and inspired Dawkins to work on what was to become the international bestseller The Selfish Gene (1976). With its publication, Oxford was soon the center for what came to be known as the gene's eye view of evolution. This logic of how genes for certain behavioral tendencies increased or decreased within a population strongly informed research questions in behavioral ecology. Behavioral ecology focused on understanding selective forces that have shaped behavior over evolutionary time in its current ecological context. The logic of the gene's eye view also lent itself to being connected with a second trend of understanding animal behavior as if it were a strategic choice in order to increase reproductive success within a population. In this view, strategies for reproductive success depended not only on the ecological factors surrounding a given population but also the behaviors of others, turning the quest for optimal outcomes into a social game. These behavioral choices and strategies could best be understood by adopting models from economics, game theory and cost‐benefit analysis optimality theory. John Maynard Smith, George Price, Geoffrey Parker, John Krebs, and Nicholas Davies all made pioneering contributions in applying these theories and methods to animal behavior in the late 1960s and early 1970s.

Both of these trends created an understanding of natural selection as an optimizing process that yielded results in behaviors that were optimally adapted to their ecological and social environments, applying this thinking to behaviors such as foraging or mating. Parker describes these theoretical advances as the “two stages of ethology's metamorphosis into behavioral ecology.” Both trends shift the focus away from ethology's interest in species‐typical behavior to research questions that switch between a gene‐centered and population‐based perspective: The individual animal became the vessel for scheming and strategic genes and its behavior was seen as a means to resolve conflicts and competition to further its inclusive fitness.

These theoretical innovations based on a population‐focused, gene‐centered and strategic view of animal behavior allowed the development of clear and specific hypotheses that could be tested by collecting long‐term data sets and analyzing them with statistical methods. Davies recounts that the theories were merged with the Tinbergian tradition of extensive field studies, careful experimentation, and a comparative perspective to create the foundation of behavioral ecology. Davies and Krebs laid the foundations of this new discipline in their defining 1978 textbook: Behavioral Ecology: An Evolutionary Approach .

There is a general consensus that behavioral ecology as a discipline was shaped by the Krebs and Davies textbook (Figure 1). While there were other textbooks at the time that covered similar ground, Behavioural Ecology was by far the most reprinted and widely used. As one reviewer put it:

Figure 1

Cover of the first edition of Behavioural Ecology: An Evolutionary Approach. The cover shows a hawk chasing doves and a game‐theoretic matrix with hawks and mice, thereby emphasizing the analytical approach of behavioral ecology. Krebs, John and Nicholas Davies, Behavioural Ecology: An Evolutionary Approach (Oxford: Blackwell Scientific Publications, 1978). Copyright © 1978 by Blackwell Scientific Publications. Used by permission. All rights reserved.

The names Krebs and Davies are virtually synonymous with the term behavioral ecology in the eyes of most undergraduate students. References to one author or both in various associations litter the examination answers of uncertain candidates on the (half reasonable) assumption that they must have said something about the topic at hand somewhere.

Unlike the edited volumes earlier in the decade that showed a diversification of the discipline of ethology that lacked a coherent core, in 1978 Krebs and Davies could present a unity of conceptual viewpoint in behavioral ecology that was remarkable. While this continuation of ethology was too theoretical for some, its origins and identity were clearly represented. Reviewers knew where it came from: “Between 1965 and 1975, theoretical ecology, population genetics, ethology, and theoretical evolutionary biology fused,” and what it stood for: “emphasis is laid on the theoretical development of quantitative predictions which can be tested, and are tested, by observation and experiment both in field and laboratory.”

One ambition Krebs and Davies had for their textbook becomes clear in a review by Davies of Douglass H. Morse's textbook Behavioral Mechanisms in Ecology (1980). While he is generally complimentary, he complains that it “does not convey excitement, because it does not present empirical data in light of theory.” This makes clear that behavioral ecology as it was conceived in Oxford was not only about new theory or methodology but also about a communal feeling of excitement about these discoveries. The textbook was not only meant to serve as an introduction but also as an induction into the Oxford vibe. The textbook and its shorter companion piece, Introduction to Behavioral Ecology (1981), written jointly by Krebs and Davies and aimed at an undergraduate audience, were received as intended: “A textbook that kindles enthusiasm,” representative of “a hot field,” full of recent discoveries, and, overwhelmingly British.

In addition to its content, Introduction to Behavioral Ecology also represented the disciplinary identity of behavioral ecology as a vivid scholarly community with its center in the UK. The textbook was a collaborative effort by multiple authors, most of them British. Each author provided a chapter that was most closely related to their individual research, providing their own perspective rather than being summarized. Krebs stressed this sense of community when he recalls that they “built around exciting ideas and people that we knew,” and as the field was small enough, it was easy to be “on top of the literature and know all key people personally.” Readers quickly gained a sense that a tight‐knit group of scientists was at work here, maybe even overly so to the point of being somewhat exclusionary: Reviewers complained about the frequent use of nicknames and cliquish jargon, that was hard to understand for “non‐English workers – or even non‐Oxford behavioral ecologists.” Donald Dewsbury, an ethologist and historian of ethology, wrote in a review: “Heavily British in tone, this work provides a valuable and exciting set of sociobiological investigations that should be on the bookshelf of every specialist in the area.” While he correctly identifies the British roots of the textbook, this sentence is indicative of a problem that behavioral ecology had to confront: Their intellectual agenda was subsumed under the banner of sociobiology, a new discipline proposed by Edward O. Wilson in 1975.

Dealing with the Competition: The Challenges of Sociobiology, 1975–1985

When Edward O. Wilson published Sociobiology: The New Synthesis in 1975, he did not take aim against behavioral ecology as a discipline. Quite the opposite: Wilson saw sociobiology and behavioral ecology as two distinct disciplines that would together succeed ethology. His famous dumbbell diagram showed ethology stagnant, whereas behavioral ecology and sociobiology were to grow in originality and, by extension, relevance (Figure 2). He predicted ethology was “destined to be cannibalized by sociobiology and behavioral ecology.” At first glance, this should put behavioral ecologists and the burgeoning group of sociobiologists on the same page, ready to take over from ethology. However, most behavioral ecologists strongly identified with the ethological tradition and Wilson's book included an open attack on ethology as using outdated methodology, lacking mathematical rigor and theoretical grounding. His view of ethology is scathing: “The future, it seems clear, cannot be with the ad hoc terminology, crude models, and curve fitting that characterize most of contemporary ethology.”

Figure 2

The dumbbell diagram from Sociobiology: The New Synthesis. Although read by many as a prediction about disciplinary development, the diagram's heading clarifies that this represents Wilson's subjective take on the amount of important ideas generated in these fields. This leaves the future disciplinary status of ethology open, while making the claim that ethology has run out of important ideas to contribute to the study of behavioral biology. Wilson, Edward O., Sociobiology: The New Synthesis (Cambridge, MA: The Belknap Press of Harvard University Press, 1975). Copyright © 1975 by the President and Fellows of Harvard College. Used by permission. All rights reserved.

This attack did not sit well with ethologists in general and alienated the group of Oxford scholars who were turning to behavioral ecology: As Wilson was ready to plunge the dagger into the heart of ethology and take over, behavioral ecology presented itself to be the “legitimate heir” to ethology, drawing heavily from the Tinbergian tradition. Sociobiology, therefore, was seen as competition by behavioral ecologists: Their disciplines were similar enough to be confused with each other from the outside but different enough from the inside to insist on distinction. The challenge that behavioral ecology faced was to represent its distinct disciplinary identity when sociobiology's huge popular and media appeal gave the impression that it was, in fact, a new revolutionary discipline. Writing about sociobiology, behavioral ecologists were quick to distance themselves from the Wilsonian project.

When asked to review Sociobiology: The New Synthesis for the journal Animal Behavior, most of them commented on Wilsons's disciplinary ambitions. Robert Hinde dedicated almost his entire review to a defense of ethology's achievements and ended by criticizing Wilson for the “sales talk with which Wilson dishes up the discipline of sociobiology.” John Crook similarly accused Wilson of disciplinary politics: “This naming and renaming of fields is of course a political game […]. Nominal matters such as these are matters of fashion and his treatment of them reveals the nature of Wilson's advocacy.” They made the point that Wilson was pursuing other, non‐scientific goals, such as gaining public acclaim due to being considered the father of a discipline. Mary‐Jane West‐Eberhard even devotes the introduction to her review to what she calls “The sociobiological fairytale.” It is worth reading in full how she humorously accuses Wilson of unnecessarily trying to create a discipline by public proclamation:

Once upon a time there was a small community of modest scholars called natural historians, who devoted their lives to philosophy and the contemplation of humble plants and animals. With the passage of time and the invention of Science they began to take on new names. Some called themselves Systematists; others Ecologists; and still others Population Biologists and Ethologists. Only their enemies called them natural historians. All of the new sciences grew and became rich. However, there was one small group without a name. They went about dressed in the castoff clothing of the titled sciences, and often failed to recognize each other, even when they hurried along the same paths. So they suffered greatly. Sometimes they had to learn to collect birds or identify ants in order to get jobs. Then one day there rose up a man from among them. 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.”

She goes on:

Edward Osborne Wilson, the kindly bespectacled father of sociobiology, has assumed god‐like powers with this book […]. It is enough to make any working animal behaviorist‐ecologist tremble a bit with anticipation (what treasures of information will it hold?), with fear (will I be codified in? or out?), and with the athletic strain of holding its flopping five pounds at reading level.

Behavioral ecologists shared this impression of unnecessary spectacle. To them, Wilson's work was hardly new or original; he was just re‐packaging elements of earlier disciplines under a new label. While they appreciated the effort Wilson put into organizing the scattered empirical literature on animal behavior, they were often confused about the way Wilson arranged the theoretical framework. His synthesis was exceedingly idiosyncratic from the point of view of the Oxford orthodoxy. For instance, while Wilson included kin selection, it did not function as a central organizing principle of the book. He also did not exclude the possibility of group selection and, worst of all, even classified kin selection as a case of group selection, which Hamilton and Krebs pointed out in their review. Krebs also noted a lack of modern methodological apparatus: for instance, the incorporation of optimality models and game theory. While the former is completely missing, the latter warranted only a brief mention in the section on ritualized combat. Krebs concluded: “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.’”

As they felt that Wilson was not representative of their work, they did not want to use the moniker “sociobiology.” In general, they felt that their ambitions were broader, focusing on all behavior in relation to the environment, whereas Wilson limited himself to social behavior. And even when UK‐based scholars actually worked on the topic of social behavior, they were apprehensive of calling it sociobiology due to the political controversy attached to this term. For instance, as Patrick Bateson writes, the King's College Sociobiology Study Group at the University of Cambridge were initially uncomfortable with using the label sociobiology, since “in many places ‘Sociobiology’ is either a battlecry or a term of abuse.” In this climate, the decision by Krebs and Davis to call their textbook Behavioural Ecology was seen as a competitive ploy against sociobiology. The authors deny that the book was written to spite Wilson and that the name was chosen to evade the controversy and to reflect the broader scope, but other ecologists had different opinions and suggested it was part of an Anglo‐American rivalry: “John Krebs spearhead[ed] a hostile takeover of E.O. Wilson's ‘new synthesis’ almost before the paint could dry, successfully usurping and greatly improving the emerging field we now perceive.”

And yet, despite their conscious effort to present the discipline of behavioral ecology in its own right, in the popular perception, especially in the US, the disciplinary lines between sociobiology and behavioral ecology were blurred. Even though Krebs and Davies did not describe Sociobiology in their introduction as an inspiration, but only mention it to differentiate their own project, this did not preclude reviewers from repeatedly naming Wilson as an influence on its development. The confusion is especially present in the reviews of the undergraduate textbook Introduction to Behavioural Ecology. US reviewers described it as a great introduction to sociobiology, a defense of sociobiology, were openly confused about the differences between the fields, or saw them as irrelevant . Looking back in 2005, Parker concedes that the distinction between the fields was not clear, many saw them as synonyms, and that in the late 1970s, sociobiology was used more.

But while sociobiology created immense attention in the mid to late 1970s, it subsequently failed to catch on. Sociobiology provoked countless pages of articles in newspapers and journals, books and reviews to be written. It was the topic of panels and entire conferences, of films and documentaries, but all the press and public attention overshadowed the fact that substantially hindered sociobiology's establishment as a discipline: There were very few actual practitioners in animal behavior calling themselves “sociobiologists.” Wilson himself did not establish sociobiological research projects or sites, recruited few researchers to the discipline and almost immediately moved on to work on modeling gene‐culture co‐evolution with Charles Lumsden. Wilson was not really in the business of social discipline building. Meanwhile, behavioral ecologists, maybe due to their overwhelming British heritage, kept calm and carried on, strengthening their disciplinary identity over time. The recipe for success was the commitment of the entire disciplinary community. While Wilson hid for four years to produce Sociobiology as a single author monograph, Krebs and Davies mention directly in the introduction that the multiple author approach was a conscious choice: “We felt that the sort of book we had in mind would have to be written by a number of authors currently working in the field rather than as a single author text.” While Sociobiology was one big monograph published in 1975, never to be updated, both textbooks by Krebs and Davies became living documents that were updated and substantially revised in each edition: “They have done it again! […] They do not just sit back and update the earlier edition. Instead […] the new edition is an almost completely rewritten book.” This meant that new scholarship was being generated, included in the disciplinary canon and then disseminated to the next generation. While sociobiology remained static, behavioral ecology became a dynamic discipline. As the second edition of Introduction to Behavioural Ecology was published in 1981, there was no mention of sociobiology in the reviews anymore.

By the mid‐1980s, behavioral ecology had successfully sharpened its disciplinary identity against its competitor, best summarized by an article that came out in celebration of Sociobiology’s ten‐year anniversary in the New Scientist written by John Krebs. He made it clear that sociobiology was merely a means to bring “the study of animal behavior from an ecological and evolutionary perspective into the public eye,” and not an original research effort: “E.O. Wilson's book Sociobiology […] was by no means the start of sociobiology and behavioural ecology.” Again, he dismissed Wilson's idiosyncratic synthesis as “an eclectic compendium of facts with little theorising.” After this assessment, he quickly moved away from the topic of sociobiology – he did not even mention the term again – and used the rest of the text to celebrate the achievements of behavioral ecology and provide more information on “What behavioral ecology is all about.” The article conveyed to the reader that the real scientific field to be celebrated is behavioral ecology, and sociobiology is merely its public relations campaign. While sociobiology was a public spectacle, behavioral ecology was a serious discipline. But did behavioral ecology's succession mean that ethology had died?

What Is Dead May never Die: Ethology in the 1980 s

In the early 1980s, the role of ethology in animal behavior studies was precarious: “Some years ago, a prominent biologist dared to ask ‘Is ethology dead?’” The rise of functional studies, in the guise of both sociobiology and behavioral ecology, left behind an “embattled ethology” that increasingly ceded territory and started a strategic retreat to a minimal disciplinary definition. Even though Konrad Lorenz claimed in 1980: “Ethology, the comparative study of behavior, is easy to define,” he himself gives a minimal and vague definition of a discipline that “applies, to the behavior of animals and humans, all those questions asked and those methodologies used as a matter of course in all the other branches of biology since Charles Darwin.” Similarly, in 1981 Samuel Anthony Barnett published a textbook that defines ethology as a major branch of biology, not limited to a certain method or school, but simply “the science of animal behavior.” This Modern Ethology had rid itself of the conceptual baggage of the past. When the Zeitschrift für Tierpsychologie was renamed Ethology in 1986, the editorial stressed the importance of the comparative approach to animal behavior, but also limited its definition of ethology to a minimal consensus: “The concept ‘Ethology’ can certainly be defined as a program, separated from the concept ‘Behaviorism.’” Finally, in an article for the New Scientist, John Durant only requires that ethology is the “study of animal behavior under more‐or‐less naturalistic conditions.” During the 1980s, the term ethology lost its original meaning and became a conceptually empty vessel for different forms of animal behavior studies that were connected to natural behaviors, a comparative and evolutionary approach, and that were not behaviorism. But it ceased to refer to the specific type of underlying theory, method, or set of concepts of classical ethology.

By the end of the decade, ethologists had become accustomed to their new disciplinary understanding and took a mostly pragmatic view on defining ethology. When the 1989 volume of Perspectives in Ethology, titled Wither Ethology?, came out, its editors (once again Patrick Bateson and Peter Klopfer) recognized the futility of providing a coherent definition of ethology: “It is possible to agonize too much about the true nature of ethology,” but recognized the need for a basic consensus to provide institutional and social identification: “Even so, people who call themselves ethologists have to live in a harsh competitive world in which funding agencies need recognizable fields to which they can allocate money and universities want subjects around which courses can be organized.”

They and others in the same volume, such as George Barlow, suggested basing ethology around a definition Tinbergen provided in his famous 1963 paper, “On aims and methods of Ethology.” In it, he lists aspects of behavior that ethologists are interested in: causation, survival value, evolution, and development. Most commonly, there are four questions derived from these areas: What is the physiological cause of the behavior? What is its survival value (or: how is it adaptive/functional)? How did it evolve? And finally: How does it develop in the individual? This paper resulted from Tinbergen's disciplinary anxieties in the 1950s, and while it provided an analytical view of the discipline, it also stressed the importance of jointly examining these aspects of behavior. It was soon used as anchoring points for ethology when it had to face conceptual uncertainty. Already in 1976, Hinde and Bateson structured the contributions along these “questions traditional to ethological research.” Rather than being an entirely empty vessel, invoking the term ethology meant invoking a specifically structured approach to animal behavior research and ethology's continued importance was due to its all‐encompassing matrix to understand animal behavior.

Other contributions in the volume used the same definition: When American ethologist George Barlow asked the question “Has sociobiology killed ethology or revitalized it?” and answered: “Ethology is dead, or at least senescent. That is, if you think of ethology in the narrow sense.” The ethology he refers to means only classical ethology; ethology as the broader project of pursuing the four questions set out by Tinbergen is alive and well. With this framework at hand, the overwhelming focus of functional studies became a mere temporary aberration of the original ethological project, not a permanent disciplinary displacement. Especially Marian Stamp Dawkins in a short contribution bemoans the one‐sided focus in recent years: “There was a time when ethology could fairly claim to be a four‐question science,” but now “Ethology may still be a four‐1egged animal, but it is an animal hopping around on one big leg, with the other three dangling somewhat ineffectively.” Therefore, she advocates for more studies in the other areas of ethology as “it is difficult to be dynamic and to run fast in the one‐legged manner that ethology seems often to adopt as its mode of progression.” However, when invoking the four‐question paradigm of ethology, all adaptationist work done in behavioral ecology was still ethology in essence. Therefore, it makes sense to retain ethology as the umbrella term for biological studies of animal behavior, as Barlow argues: “But what shall we call this field? Sociobiology is too narrow, even when it includes behavioral ecology, and so is socioecology. […] As the reader might have predicted, I would prefer retaining ethology.” Earlier ethologists had already subsumed the adaptive discipline of behavioral ecology and sociobiology under the term ethology, and it became common practice to refer to all of animal behavior studies simply as “ethology.”

How did ethology move out of being? It didn't, really. It advanced to a changed state of being: By defining its later disciplinarity along a meta‐theoretical structure and redefining its scope to be very broad, it lives on as a term for animal behavior research from a biological perspective, with few limitations applied.