Commentary: New Directions in the History of Ethology.

This welcome set of original and instructive papers illuminates and enriches the history of twentieth-century ethology in multiple ways. It adds a wealth of actors, animals, methods, and places to those featured in previous treatments of ethology's development. Some of the papers extend the chronology beyond the heyday of ethology's disciplinary construction to consider exciting developments in the 1970s and beyond. Others consider animal behavior research programs pursued contemporaneously with but independently of mainline ethology's development from the 1930s through the 1960s. Another paper takes us inside an ethologist's archive of visual images to examine the importance of such images (and such a setting) for ethological practice. Collectively, the papers provide new opportunities to contemplate how research programs and disciplines evolve; the relations between concepts, practices, and places; ethology and politics, and much more. At the same time, the individuality of the papers is conspicuous. They have not been constructed on the same model. The authors have followed their own approaches, corresponding to their own, respective interests. A short commentary is not sufficient to do justice to each of them. Rather than attempt to review them one by one, I will consider a pair of themes that may help relate the papers to each other and to the history of ethology: (1) the ongoing challenge of defining ethology and identifying who the ethologists were (or are); (2) the practices and places of animal behavior study.

Red Foxes in the Filing Cabinet: Günter Tembrock's Image Collection and Media Use in Mid-Century Ethology.

This paper considers the epistemic career of visual media in ethology in the mid-20th century. Above all, ethologists claimed close contact with research animals and drew scientific evidence from these human-animal communities, particularly in public relations. However, if we look into the toolboxes of comparative behavioral biologists, it becomes evident that scientifically valid research results were primarily obtained by experimenting with model images. These visual specimens tell a technical story of the methodological requirements in behavioral science necessary to bridge everyday observations between the laboratory and the field. By neutralizing individual traces of animal bodies as well as their observers, they prompted the abstraction of ethological hypotheses. The case study of East-German biologist Günter Tembrock (1918-2011), who maintained his own collection of newspaper clippings, drawings, photographs, and films, offers a new perspective on the methodological development of this field. Furthermore, this article contributes to a scholarly discussion geared toward expanding the spaces of ethological research. My analysis of the image collections of the Forschungsstätte für Tierpsychologie presents the archive as a relevant site of study in the history of ethology.

From Karl von Frisch to Neuroethology: A Methodological Perspective on the Frischean Tradition's Expansion into Neuroethology.

This paper examines a tradition of eusocial insect research stemming from the Austrian zoologist Karl von Frisch. As I show in this paper, one of the most enduring features of the Frischean tradition has been an experimental methodology developed by Frisch in the early 1910s. By tracing this methodology's use through Frisch's student, Martin Lindauer, and two of Lindauer's students, Rüdiger Wehner and Randolf Menzel, this paper illuminates a surprising aspect of ethology's development during the last half of the 20th century. Namely, it sheds light on how the Frischean tradition, a tradition that had a complicated relationship with ethology since the discipline's formation in the 1930s, produced scientists who became leading figures in neuroethology, the most prominent contemporary field of behavioral research to retain the label of "ethology." Some of the features that distinguished Frisch's training method from the program of classical ethology and the work of his contemporaries later helped his academic descendants adapt the method to the neuroethological program.

Karl von Frisch and the Discipline of Ethology.

In 1973, the discipline of ethology came into its own when three of its most prominent practitioners-Konrad Lorenz, Nikolaas Tinbergen, and Karl von Frisch-jointly received the Nobel Prize for Physiology or Medicine. Historians have shown how Lorenz and Tinbergen were central to the practical and theoretical innovations that came to define ethology as a distinct form of animal behavior research in the twentieth century. Frisch is rarely mentioned in such histories. In this paper, I ask, What is Frisch's relationship to the discipline of ethology? To answer that question, I examine Tinbergen's relationship to Frisch's grey card experiments between Tinbergen's time as a student at the University of Leiden in the mid 1920s and his 1951 publication of The Study of Instinct. In doing so, I highlight previously neglected affinities between Frisch's early career research and the program of classical ethology, and I show how Frisch's research meant different things at different times to Tinbergen and others working in the ethological tradition.

Social and sexual behaviors in C. elegans: the first fifty years.

For the first 25 years after the landmark 1974 paper that launched the field, most C. elegans biologists were content to think of their subjects as solitary creatures. C. elegans presented no shortage of fascinating biological problems, but some of the features that led Brenner to settle on this species-in particular, its free-living, self-fertilizing lifestyle-also seemed to reduce its potential for interesting social behavior. That perspective soon changed, with the last two decades bringing remarkable progress in identifying and understanding the complex interactions between worms. The growing appreciation that C. elegans behavior can only be meaningfully understood in the context of its ecology and evolution ensures that the coming years will see similarly exciting progress.

Innate releasing mechanisms and fixed action patterns: basic ethological concepts as drivers for neuroethological studies on acoustic communication in Orthoptera.

This review addresses the history of neuroethological studies on acoustic communication in insects. One objective is to reveal how basic ethological concepts developed in the 1930s, such as innate releasing mechanisms and fixed action patterns, have influenced the experimental and theoretical approaches to studying acoustic communication systems in Orthopteran insects. The idea of innateness of behaviors has directly fostered the search for central pattern generators that govern the stridulation patterns of crickets, katydids or grasshoppers. A central question pervading 50 years of research is how the essential match between signal features and receiver characteristics has evolved and is maintained during evolution. As in other disciplines, the tight interplay between technological developments and experimental and theoretical advances becomes evident throughout this review. While early neuroethological studies focused primarily on proximate questions such as the implementation of feature detectors or central pattern generators, later the interest shifted more towards ultimate questions. Orthoptera offer the advantage that both proximate and ultimate questions can be tackled in the same system. An important advance was the transition from laboratory studies under well-defined acoustic conditions to field studies that allowed to measure costs and benefits of acoustic signaling as well as constraints on song evolution.

Animal Behavior, Population Biology and the Modern Synthesis (1955-1985).

This paper examines the history of animal behavior studies after the synthesis period. Three episodes are considered: the adoption of the theory of natural selection, the mathematization of ideas, and the spread of molecular methods in behavior studies. In these three episodes, students of behavior adopted practices and standards developed in population ecology and population genetics. While they borrowed tools and methods from these fields, they made distinct uses (inclusive fitness method, evolutionary theory of games, emphasis on individual selection) that set them relatively apart and led them to contribute, in their own way, to evolutionary theory. These episodes also highlight some limitations of "conjunction narratives" centered on the relation between a discipline and the modern synthesis. A trend in conjunction narratives is to interpret any development related to evolution in a discipline as an "extension," an "integration," or as a "delayed" synthesis. I here suggest that this can lead to underestimate discontinuities in the history of evolutionary biology.

The Development of Sociobiology in Relation to Animal Behavior Studies, 1946-1975.

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".

The 'Disadapted' Animal: Niko Tinbergen on Human Nature and the Human Predicament.

This paper explores ethologist Niko Tinbergen's path from animal to human studies in the 1960s and 1970s and his views about human nature. It argues, first, that the confluence of several factors explains why Tinbergen decided to cross the animal/human divide in the mid 1960s: his concern about what he called "the human predicament," his relations with British child psychiatrist John Bowlby, the success of ethological explanations of human behavior, and his professional and personal situation. It also argues that Tinbergen transferred his general adaptationist view of animal behavior to the realm of human biology; here, his concern about disadaptation led him to a view of human behavior that was strongly determined by the species' evolutionary past, a position that I call evolutionary determinism. These ideas can be seen in the work he carried out with his wife, Elisabeth Tinbergen, on autism. The paper concludes that Tinbergen's vision of human nature constitutes another version of what anthropologist Clifford Geertz called in 1966 the "stratigraphic" conception of the human: a view of human nature as a composite of levels in which a universal ancestral biological core is superimposed by psychological and cultural layers that represent accidental variation at best and pathological deviation at worst.

Africanizing Science in Post-colonial Kenya: Long-Term Field Research in the Amboseli Ecosystem, 1963-1989.

Following Kenya's independence in 1963, scientists converged on an ecologically sensitive area in southern Kenya on the northern slope of Mt. Kilimanjaro called Amboseli. This region is the homeland of the Ilkisongo Maasai who grazed this ecosystem along with the wildlife of interest to the scientists. Biologists saw opportunities to study this complex community, an environment rich in biological diversity. The Amboseli landscape proved to be fertile ground for testing new methods and lines of inquiry in the biological sciences that were generalizable and important for shaping natural resource management policies in Kenya. However, the local community was in the midst of its own transformation from a primarily transhumant lifestyle to a largely sedentary one, a complex political situation between local and national authorities, and the introduction of a newly educated generation. This article examines the intersection of African history and field science through the post-colonial Africanization of Kenyan politics, the broadening of scientific practices in Amboseli in previously Western-occupied spaces to include Kenyan participants, and an increasing awareness of the role of local African contexts in the results, methods, and implications of biological research. "Africanization" as an idea in the history of science is multifaceted encompassing not just Africans in the scientific process, but it needs an examination of the larger political and social context on both a local and national level.

Professor Robert McNeill Alexander CBE FRS (1934-2016).

Robert McNeill Alexander, known to friends and colleagues as 'Neill', was a zoologist with an engineer's eye for how animals work. He used mathematical models to show how evolution has produced optimal designs. His skill was to choose appropriate models: realistic enough to contain the essence of a problem and yet simple enough to be tractable. He wrote fluently and easily: 23 books, 280 papers and a CD-ROM entitled How Animals Move.

Primate Kinship: Contributions from Cayo Santiago.

Research on Cayo Santiago and Japan deserves credit for launching the study of primate kinship and for continuing to help shape it in important ways. This review describes the origins of kinship research on Cayo Santiago, beginning with Donald Sade's pioneering work establishing the concepts of kin preferences, matrilineal dominance systems and incest avoidance. It then reviews subsequent research by later Cayo Santiago researchers and alumni, focusing primarily on maternal kinship. Together these researchers have greatly expanded our knowledge of kin preferences in rhesus in terms of (i) what age-sex classes, behaviors and types of kin show them, (ii) the ways in which kinship interfaces with rank, sex, age, and dispersal patterns, and (iii) the graded and variably limited nature of kin preferences in terms of degree of relatedness. Second, the argument for kin selection at least for some types of behavior has survived challenges posed by several alternative explanations, and has been both strengthened by recent findings of paternal kin preferences and narrowed by studies showing that unilateral altruism may extend only to very close kin. Third, work on Cayo Santiago has contributed to an appreciation that both current conditions and inherent social characteristics may influence the strength of kin preferences, and fourth, it has contributed to an understanding of the possible origins of our own species' family systems. Cayo Santiago became a leader in kinship research in large part because of management practices that produce known extended lineages. These lineages have promoted and accelerated research on kinship, prompting other researchers to investigate its importance in other groups and species, where its effects only then became clear. The extended lineages remain valuable tools for research on a species that lives in a broad range of environments in the wild, including those with key parallels to Cayo Santiago.

A 75-year pictorial history of the Cayo Santiago rhesus monkey colony.

This article presents a pictorial history of the free-ranging colony of rhesus monkeys (Macaca mulatta) on Cayo Santiago, Puerto Rico, in commemoration of the 75th anniversary of its establishment by Clarence R. Carpenter in December 1938. It is based on a presentation made by the authors at the symposium, Cayo Santiago: 75 Years of Leadership in Translational Research, held at the 36th Annual Meeting of the American Society of Primatologists in San Juan, Puerto Rico, on 20 June 2013.

Cayo Santiago--a scientific treasure.

This article briefly reviews the contributions of Clarence R. Carpenter in establishing the free-ranging colony of rhesus monkeys on Cayo Santiago, the Caribbean Primate Research Center and his legacy in primatology.

Niko Tinbergen: A Message in the Archives.

Just as biologists have their favored places for doing research, so do historians. As someone who likes working in archives, the most surprising thing the present author ever found was a particular letter that had been written to him by the ethologist Niko Tinbergen-but that Tinbergen had never sent. The letter included a detailed critique of the intellectual style and conceptual shortcomings of Tinbergen's career-long friend and colleague Konrad Lorenz. The present author first saw the letter 3 years after Tinbergen's death and 10 years after the letter was composed. Here we discuss the contents and historical context of that letter.

Understanding Societies from Inside the Organisms. Leo Pardi's Work on Social Dominance in Polistes Wasps (1937-1952).

Leo Pardi (1915-1990) was the initiator of ethological research in Italy. During more than 50 years of active scientific career, he gave groundbreaking contributions to the understanding of social life in insects, especially in Polistes wasps, an important model organism in sociobiology. In the 1940s, Pardi showed that Polistes societies are organized in a linear social hierarchy that relies on reproductive dominance and on the physiological and developmental mechanisms that regulate it, i.e. on the status of ovarian development of single wasps. Pardi's work set the stage for further research on the regulatory mechanisms governing social life in primitively eusocial organisms both in wasps and in other insect species. This article reconstructs Pardi's investigative pathway between 1937 and 1952 in the context of European ethology and American animal sociology. This reconstruction focuses on the development of Pardi's physiological approach and presents a new perspective on the interacting development of these two fields at the origins of our current understanding of animal social behavior.

"To ear is human, to frogive is divine": Bob Capranica's legacy to auditory neuroethology.

Bob Capranica was a towering figure in the field of auditory neuroethology. Among his many contributions are the exploitation of the anuran auditory system as a general vertebrate model for studying communication, the introduction of a signal processing approach for quantifying sender-receiver dynamics, and the concept of the matched filter for efficient neural processing of complex vocal signals. In this paper, meant to honor Bob on his election to Fellow of the International Society for Neuroethology, I provide a description and analysis of some of his most important research, and I highlight how the concepts and data he contributed still inspire neuroethology today.

Toward a cross-species neuroscientific understanding of the affective mind: do animals have emotional feelings?

Do we need to consider mental processes in our analysis of brain functions in other animals? Obviously we do, if such BrainMind functions exist in the animals we wish to understand. If so, how do we proceed, while still retaining materialistic-mechanistic perspectives? This essay outlines the historical forces that led to emotional feelings in animals being marginalized in behavioristic scientific discussions of why animals behave the way they do, and why mental constructs are generally disregarded in modern neuroscientific analyses. The roots of this problem go back to Cartesian dualism and the attempt of 19th century physician-scientists to ground a new type of medical curriculum on a completely materialistic approach to body functions. Thereby all vitalistic principles were discarded from the lexicon of science, and subjective experience in animals was put in that category and discarded as an invalid approach to animal behavior. This led to forms of rigid operationalism during the era of behaviorism and subsequently ruthless reductionism in brain research, leaving little room for mentalistic concepts such as emotional feelings in animal research. However, modern studies of the brain clearly indicate that artificially induced arousals of emotional networks, as with localized electrical and chemical brain stimulation, can serve as "rewards" and "punishments" in various learning tasks. This strongly indicates that animal brains elaborate various experienced states, with those having affective contents being easiest to study rigorously. However, in approaching emotional feelings empirically we must pay special attention to the difficulties and vagaries of human language and evolutionary levels of control in the brain. We need distinct nomenclatures from primary (unconditioned phenomenal experiences) to tertiary (reflective) levels of mind. The scientific pursuit of affective brain processes in other mammals can now reveal general BrainMind principles that also apply to human feelings, as with neurochemical predictions from preclinical animal models to self-reports of corresponding human experiences. In short, brain research has now repeatedly verified the existence of affective experience-various reward and punishment functions-during artificial arousal of emotional networks in our fellow animals. The implications for new conceptual schema for understanding human/primate affective feelings and how such knowledge can impact scientific advances in biological psychiatry are also addressed.