History of Ecological Sciences, Part 56: Ethology until 1973

A distinction can be drawn between ecological and psychological studies of animal behavior. The former became ethology and the latter, behaviorism. Robert Boakes, University of Sussex, wrote the history of the latter: From Darwinism to Behaviourism: Psychology and the Minds of Animals (1984). However, behavioral psychology is relevant to the study of ethology (Kruuk 2003:3, Burkhardt 2005:8), despite Konrad Lorenz reading works by "Spencer, Lloyd Morgan, MacDougall, Yerkes and Watson" and concluding that "they simply did not know what they were talking about" (quoted in Nisbett 1976:37). While Lorenz's wholesale dismissal of other approaches and findings than ethological was unfair, his comments illustrate an attitudinal gulf between two different approaches to the study of animal behavior. Yet, zoologists Peter Klopfer and Jack Hailman blended the two approaches in their text, An Introduction to Animal Behavior: Ethology's First Century (1967), as did ethologist Robert Hinde in his Animal Behaviour: a Synthesis of Ethology and Comparative Psychology (1970), Michael Fox in his Readings in Ethology and Comparative Psychology (1973), and the first two chapters in P. P. G. Bateson and Peter Klopofer's Perspectives in Ethology (volume 1, 1973) are by authors from departments of psychology. American collaborators V(incent) G. Dethier, biologist, and Eliot Stellar, physiological psychologist, coauthored Animal Behavior (1961, edition 3, 1970), which has six chapters on nervous systems in different kinds of animals and three chapters on behavior, learning, and intelligence. Michael Bright's Intelligence in Animals (1994) is a synthesis of these two approaches for a popular audience. On the other hand, Lee Drickamer has discussed comparative psychology (example: Maier and Maier 1970), ethology, and ecology as three-fold roots of the study of animal behavior, and he mentioned other investigators' ways to subdivide these subjects (Vessey and Drickamer 2010, Drickamer 2016). Those various approaches are not discussed here. There are excellent sources for history of ethology. However, Alfred Espinas' lengthy "Introduction Historique" to Des Sociétés Animales (1878:7–155) has limited interest here, because, rather oddly, it is limited to a survey of ideas on human society (to which he also devoted his final chapter, 458–513, before his conclusions). Psychologist Philip Gray, Montana State University, wrote three historical articles of narrow scope (1962, 1963, 1967), followed by a brief broad historical survey (1968). Thomas McGill, Williams College, compiled a collection of 55 extracts in Readings in Animal Behavior McGill (1965) that was topically organized and included two essays on a new science of ethology (Thorpe 1956, Hess 1962). Klopfer and Hailman's Introduction to Animal Behavior (cited above) is not exactly a history of ethology's first century, but is actually a textbook on ethology written from a historical perspective. G. Richard, Université de Rennes, wrote authoritatively "The Historical Development of Nineteenth and Twentieth Century Studies on the Behavior of Insects" (1973). An attractive, 725-page Grzimek's Encyclopedia of Ethology (1977, German, 1973), contains (English edition) two brief articles on the history of ethology; neither has references (Burghardt 1977, Heinroth 1977). Nevertheless, Katharina Heinroth's survey should be taken seriously, because it is based upon her own and her late husband, Oskar Heinroth's ethological knowledge. Konrad Lorenz wrote a historical introduction to his Foundations of Ethology (1981:1–12, German, 1978), which included history of his own studies. Cambridge University ethologist William Thorpe wrote a concise The Origins and Rise of Ethology (1979), which included references and brief account of his own work. John Durant wrote "Innate Character in Animals and Man: a Perspective on the Origins of Ethology" (1981), with references. Bernard Singer's "History of the Study of Animal Behaviour" (1982) is the only history I know that seriously surveys from antiquity to recent times. His slight confusion concerning Réaumur's publications and an incorrect date for Gray 1968 are minor errors, but documentation for his article is limited. Gordon Burghardt compiled Foundations of Comparative Ethology (1985) that contains extracts from 1835 to 1937. F. M. Scudo discussed aspects of history in "'Ethology,' 'Ecology,' and 'Philosophy''' Scudo (1989). Unique resources are two volumes of autobiographical essays: Studying Animal Behavior: Autobiographies of the Founders (Dewsbury 1985) and Leaders in Animal Behavior: The Second Generation (Drickamer and Dewsbury 2009). Irenäus Eibl-Eibesfeldt, author of an ethology textbook (1970), earlier published, with Sol Kramer, an essay explaining this new science (1958). Feminist historian of biology Donna Haraway published a detailed Primate Visions: Gender, Race, and Nature in the World of Modern Science, beginning in 1908 (1989), and Georgina Montgomery published a less ideological study than Haraway's: Primates in the Real World: Escaping Primate Folklore and Creating Primate Science (2015). In 1989, the American Society of Zoologists celebrated its centennial with six papers in a Symposium on "Animal Behavior: Past, Present, and Future," which appeared in American Zoologist (1991). Lynne Houck and Lee Drickamer edited a more chronologically broad collection than McGill's: Foundations of Animal Behavior: Classic Papers with Commentaries (1996), which included works from 1872 to 1973. Their Foundations is divided into six parts, and each part has a historical introduction by a different investigator. John Clark's "Ethology and animal behavior" (2000) is a bibliographical essay that cited and critically discussed 17 publications. There are excellent detailed biographies of the two most important founders of ethology, both well illustrated: Klaus Taschwer and Benedick Föger, Konrad Lorenz: Biographie (2003), and Hans Kruuk, Niko's Nature: The Life of Niko Tinbergen and His Science of Animal Behaviour (2003). Two years later, Richard Burkhardt, Jr. published, also in great detail, Patterns of Behavior: Konrad Lorenz, Niko Tinbergen, and the Founding of Ethology (2005), which is broader in timespan than the title indicates. Lee Alan Dugatkin, like Eibl-Eibesfeldt, has published both a textbook (2004, edition 2, 2009b) and a historical study: The Altruism Equation: Seven Scientists Search for the Origins of Goodness (2006). His textbook contains interviews with 17 ethologists. Dale Peterson's lengthy biography of Jane Goodall (2006) includes details on the history of ethology, particularly in relation to primates. Tim Birkhead, Jo Wimpenny, and Bob Montgomerie's Ten Thousand Birds: Ornithology since Darwin (2014) has three relevant chapters: "Ecological Adaptations for Breeding," "The Study of Instinct" and "Behavior as Adaptation." A Journal of the History of the Behavioral Sciences began in 1965. The series, Perspectives in Ethology, edited by P. P. Bateson and P. H. Klopfer (vol. 1, 1973, vol. 2, 1976, etc.), is not historically oriented, but as time goes by, early volumes become of historical interest. Other historical articles are cited below. This survey is selective; the 19 autobiographical essays collected by Dewsbury in Leaders in the Study of Animal Behavior (1985), are from investigators who had made substantial contributions by 1973. English ethologist William H. Thorpe (1902–86) was likely invited to submit an autobiographical essay (to make a total of 20), but perhaps he was unable to write one, due to declining health (Gillespie 1990, Costall 2004). He had briefly summarized his own work in his Origins and Rise of Ethology (1979:118–123). Some autobiographical essays collected by Drickamer and Dewsbury, Leaders in Animal Behavior: The Second Generation (2010) also include noteworthy contributors active before 1973. There are at least three dictionaries of ethology. One, edited by Harré and Lamb (1986) has entries by 46 authors that include historical information and bibliographies. The dictionary by Immelmann and Beer (1989) is mostly a translation of Immelmann's Worterbuch der Verhaltensforschung (edition 3, 1982). Edward Barrows' Animal Behavior Desk Reference: A Dictionary of Animal Behavior, Ecology, and Evolution (1995, edition 3, 2011) is the most detailed, with historical references and bibliography. With abundant historical resources, I have been selective rather than comprehensive. My end-date of 1973 is based upon the award of a Nobel Prize in 1973 to Frisch, Lorenz, and Tinbergen, which brought ethology from the periphery into mainstream biology and ecology. Conveniently, Hinde's Animal Behaviour (1970) and Irenäeus Eibl-Eibesfeld's textbook, Ethology: the Biology of Behavior (1970) summarized much of what ethologists had achieved by then; the National Geographic Society published a well-illustrated, popular survey of The Marvels of Animal Behavior (Allen 1972) by 22 authors (all academics), and the 41 chapters in Grzimek's Encyclopedia of Ethnology (Immelmann 1977:725, German, 1973) provided broad surveys of ethology around the same time (Fig. 1). The most important early source for ethology was natural history literature, and in W. H. Thorpe's conception of ethology, its contributions followed "three scientific ways of probing into the behavior of animals" (1956, reprinted 1965:35), the ways of naturalists, psychologists, and physiologists. On each flight the bee does not go on to flowers different in form; it goes for example from violet to violet, and does not touch any other before it has flown back to the hive. And having arrived at the hive they shake themselves, and three or four bees attend to each. What they pick up is not easy to see, nor has there method of working it been seen… That is, the dance was discovered in antiquity, but not its purpose. Jean-Louis Labarrière wrote a detailed discussion of "Aristote et l'éthologie" (1993), primarily focused upon Historia Animalium, books 8 and 9. Liliane Bodson (1996) concentrated on Aristotelian accounts of bird behavior. Aristotelian zoological writings considered behavior in all kinds of animals known to ancient Greeks, but with more information on birds than on any other group—birds being most easily observed. Bodson focused upon reproductive behavior, from courtship to raising young and found information reported was impressively accurate. An example she discussed in detail was European cuckoo brood parasitism (HA 618a:26-31). Jason Tipton's studies (2006, 2008) focused on habits of rocky near-shore giant goby Gobius cobitis and intertidal blenny Parablennius sanguinolentus, and upon foraging interactions between mullet (Mullus spp) and sea bream (Diplodus spp) in Aristotelian writings (Fig. 2). Animal studies received a boost from the less scientifically sophisticated Roman author, Gaius Plinius Secundus (Pliny the Elder, about 23–79 AD), whose encyclopedic Naturalis Historia was a compendium of Greek and Roman nature writings (Singer 1982:261, Egerton 2001b:245, Egerton 2012:12–13). Although not very successful in his attempt to separate fact from fiction, Pliny exerted a broad influence during the Middle Ages and later in the Latin West, though not in the Byzantine East (Egerton 2002a) or in Muslim civilization; Arabic language nature writings were influenced by Aristotelian works and Byzantine tradition (Singer 1982:262, Egerton 2002b, c, 2012:17–24). A west European emperor, Friedrich II (1194–1250), lived mainly in Sicily, where he was influenced by Latin, Byzantine, and Muslim civilizations, and he introduced falconry into Western Europe from the Muslim East (Allen 1951:398–400, Stresemann 1951:8–11+2 plates, 1975:9–12, Egerton 2003a, 2012:25–27, Birkhead 2008:137–142). His treatise on falconry included his own observations not only on hawks used for hunting, but also on birds which his hawks pursued. His natural history achievements made him an amateur avian ecologist and behaviorist. His De arte venandi cum avibus was published in 1596, but only attracted ornithologists' attention in 1788. The Latin Middle Ages also produced several encyclopedists, who blended ancient and medieval learning (Singer 1982:263). Some of them were more sophisticated than Pliny, though also accepting some stories now considered incredible. The best of these encyclopedists was German scholastic Albert of Lauingen (Albert the Great, about 1200–80), who studied in Paris about 1240, and then taught there until 1248, before settling in Cologne to teach (Wallace 1970). His De animalibus libri XXVI was his longest and most influential encyclopedia, begun between 1256 and 1260, and drawing upon Aristotelian and medieval sources and his own observations are in books XX–XXI (Allen 1951:400–401, Stresemann 1951:11–12, 1975:12–13, Egerton 2003b, 2012:27–30, Birkhead 2008:see index). However, his books XXII–XXVI are based, without acknowledgment, upon De natura rerum of his former pupil, Thomas of Cantimpré (1186/1210-76/94), who studied under him in 1245–48 before completing his studies in Paris (Aiken 1947, Kibre 1976). One of the medieval tales which Albert repeated was about the osprey (Albertus 1999:599): "It hunts only fish and has one webbed foot, for swimming, like that of a goose, while the other foot has hooked talons for seizing, like that of an eagle." A more credible report concerned geese, which bred in Sclavia at "moist, sandy, marshlike places," but at the beginning of winter they "come back to our land," seeking "food and the more temperate air" (Albertus 1999:613) (Fig. 3). It has short wings, and flies as if by bounds upwards and downwards. It lives on beetles, butterflies, and biggish insects, and not only these, but also birds after the manner of a Hawk. For it kills Reguli and Finches and (as I once saw) Thrushes; and bird-catchers even report that it from time to time slays certain woodland Pies, and can put Crows to flight. It does not seize the birds it kills with its claws, after a swift flight, as Hawks do, but attacks them stealthily and soon (as I have often had experience) aims at the throat and with its beak squeezes and breaks the skull….when prey happens to be more plentiful, it lays by some for future scarcity. For it impales and hangs the bigger flies and insects on the thorns and spines of shrubs… An illustration of this shrike is below, Fig. 5b. A fellow English physician was Edward Wooton (1492–1555), older than Turner, but whose De Differentiis Animalium (1552) appeared after Turner's Avium (Bodenheimer 1928-29:I, 230–233, Raven 1947:40–42, Wheeler 1976). It was much broader in scope than Avium: chapters 5 on "quadrupeds that bear live young," 6 on "quadrupeds that lay eggs," 7 on birds, 8 on fish, 9 on insects, and 10 on squids, crustaceans, mollusks. It was an efficient compilation from previous works, but contained few, if any, first-hand observations. Some pioneer Renaissance naturalists added illustrations to their books. German physician Otto Brunfels (about 1489–1534) compiled Herbarum vivae eicones (three volumes, 1530–36, German, two volumes, 1532–37) with traditional text, but with fresh woodcuts by artist Hans Weiditz (Stannard 1970, Reeds 1976, 1991, Greene 1983:1, chapter 5). Once Weiditz got started, he kept going until he had illustrated 47 plants not represented in classical pharmacopoeia. Brunfels apologized for including them, but added names and descriptions of them. Conrad Gessner (or Gesner, 1516–65) of Zurich, compiled a well-illustrated Historia animalium (5 volumes, 1551–87, German edition, 4 volumes, 1557–89) and a posthumously published Historia plantarum (1751–71), both of which included his own observations along with traditional learning (Allen 1951:402–403, Stresemann 1951:18–22, 1975:18–21, Greene 1983:II, 747–797, Wellisch 1984, Egerton 2003:206–208, 2012:34–40). Gessner supported his family by being both a professor at the University of Zurich and as a physician, and when an epidemic of plague reached Zurich, he died along with the patients he treated. Englishman Edward Topsell (1572–1625) published a translation of Gessner's first volume on mammals and extracts from Gessner's volumes 2, 4, and 5 as his own second volume in 1607 (Raven 1947:217–226, Bircham 2007:71–73, Birkhead 2008:36–37). An Italian naturalist-encyclopedist, Ulisse Aldrovandi (1522–1605), continued in Gessner's footsteps (Bodenheimer 1928-29:I, 247–276, Allen 1951:403–405, Stresemann 1951:22–24, 1975:see index, Castellani 1970, Beier 1973:85–86, Singer even to the of Gessner's work with He did not to all of his works before he though he did three volumes on birds and one on Studies on insects in antiquity and continued on the Middle Ages and Renaissance (Bodenheimer 1928-29:I, Beier 1973, and as a separate science with of De libri edition 3, In other volumes he upon Gessner's volumes, Gessner died before his and Aldrovandi more on his own observations for this Raven was with the Gessner was not the only from the who died before completing his volume on English Thomas (about most of his plants Raven However, he had in Europe after his had been with Gessner while he was working on insects, and brought back to Gessner's and illustrations on he to the study of insects in his later he had Gessner's to his own and when he were by a Thomas English who focused upon insects (Bodenheimer 1928-29:I, Beier the for but to a the was by who got it published in Latin in under The Aldrovandi and volumes were written at about the same though the was three before and Beier (cited above) them. volumes had and and they several other during the The volume has the of now being in In there is an illustration and of a but the natural history contains this a is this that if a the to a she one of her and him the and or (Fig. English John followed the encyclopedic tradition in his plantarum Historia plantarum (three volumes, and Historia but he that tradition in other He published under the of his former and English which was actually more than by (Raven Bircham Birkhead 2008:see and the same was of their De Historia (Raven had an early interest in insects and discussed their habits in some early but only in his did he write two on and Historia both posthumously (Raven he also published works The of in the of the and concerning the and of the World It is in this at his first to or seize upon one as its into which it not any other but its However, part of added by in its exerted a influence upon and American natural history studies (Thorpe Egerton His William wrote a natural history or a of the and of from his of edition 4, also discovered the by his own and and other Egerton His were considered important for him to be a of the Society of and his were published in English translation in that He and than and he even discovered from his own He also studied other as the insects found in on He found that lay as they and that the same in their He did not that his animals" but is for the of her to her own for if they not and by Stresemann his in only a of our and be with Oskar Heinroth and Konrad (Fig. 5). de was a who devoted most of his attention to insects Egerton His volumes, was a (Bodenheimer 1928-29:I, He was not attracted by the of and all the he he studied behavior of He wrote two for a volume which he or published, but they appeared about two later 1977, de His was most important of the and was also of like of the by a without the of any or they are by one or several that are by several for the of of The was not one or the other Wheeler both of different Réaumur's influence upon other naturalists of the and early was de Réaumur's was in the the of a a de was 1970, Egerton the of the in a very that him to with an et volumes, It included volumes on mammals and volumes on birds both volumes broader and more detailed of mammals and birds than his to animals and him that he broad based upon what now was An example of his is his discussion of He did not scientific that along with illustrations was His account of began with a discussion of and then discussed its natural history the He to three The in which the from the as in and the of the (vol. in a discussion Animals to the he that of in the known of the of them in the and less than in the if the to both World In Nature is less and more are much than of the Thomas was a in a with the of He was also in the of his on the State of and he at that American mammals were to of the and he by an with of in his and he from the and of a than any in to (Fig. A de et his in the from his What the of a was his with encyclopedists and for he wrote including one on Gray and Thorpe to have been the most important early to the study of animal behavior. His was in his Gray and Thorpe its date as only a of his had been published by then his and into the there to and of these to the and of this of and to by repeated by they from to intelligence. that and had provided and that he to have the biography of He was a science as ethology and he provided the in his books that an of they in about in the from and from In a after these had made it known to the they in great to the This of had influence on later behavioral studies. English studied in detail the natural history of his and its Burkhardt Egerton history was not an in One it mainly from natural history literature, and for from the writings of and History and of due to his observations and has in to the though as the added illustrations of birds and mammals to her edition from Thomas which is since of the of in History were to and some of observations in his own treatise illustrated his edition of with woodcuts by Thomas a later of began his to in and he began in and continued until he His were with what with several entries for each He then his on subjects with information from his natural history primarily of his own observations on the and raising young of the he with about of and also on the of plants observed. He other natural history writings and or what wrote on he studied (Fig. the a between the birds that they can bear to be in the same or of the and of of that time to to be the of and and it is to this of that I at

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