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• W2190647748 abstract "Modern population ecology began during the period 1860 to the 1920s, along with early formal development of animal ecology (Kingsland 1985:9–112, Price 2003:9–13, Egerton 2014a). Population ecology advanced on three fronts, sometimes linked: field studies, laboratory studies, and mathematical–theoretical studies (Cole 1954:106, 1958:6). It seems desirable to briefly treat here two special cases: invasive species and rare and extinct species, with bibliographic guides at these topics. There is helpful literature on the history of population ecology; yet its authors were often unaware of each other's contributions. William Thompson wrote a 26-page historical introduction to his “Biological Control and the Theories of the Interactions of Populations” (1939:301–327). Thomas Park's “Some Observations on the History and Scope of Population Ecology” (1946) has such general comments it has limited current interest. LaMont Cole (1954:105–117, 1958:2–11) set an excellent example by surveying the history of human and animal demography, observational and mathematical. David Lack's Natural Regulation of Animal Numbers (1954) encompassed animals in general; although his Population Studies of Birds (1966) was narrower in scope, its appendix, “The Theoretical Controversies concerning Animal Populations,” covers animals in general. Frank Egerton has written articles on animal demography from antiquity to Darwin (listed in McIntosh 1985:335, Egerton 2012:226). Kenneth Watt's “Use of Mathematics in Population Ecology” (1962) is a very insightful historical survey. Andy Andrewartha and Louis Birch's “The History of Insect Ecology” (1973) contains an important survey on the history of insect demography. (Although Lack, and Andrewartha and Birch, were participants in population controversies, they all attempted to present fair accounts.) J. P. Dempster (1975:121–128) provided a concise and lucid history of population theories. Evelyn Hutchinson's Introduction to Population Ecology (1978) began with a historical chapter and had historical comments and footnotes throughout the book, and five pages from it are reprinted in an anthology of his writings (Hutchinson 2010:10–14). In the same year as Hutchinson's population textbook, Robert Tamarin compiled a useful sourcebook, Population Regulation (1978) and Francesco Scudo and James Ziegler compiled The Golden Age of Theoretical Ecology, 1923–1940 (1978). The last contained two papers by Lotka from 1923 and one by Volterra from 1927; all other papers were from the 1930s. Sharon Kingsland's Modeling Nature: Episodes in Population Ecology (1985) is a valuable guide, which cited Thompson (1939), Cole (1954), Lack (1966), Hutchinson (1978), Scudo and Ziegler (1978), but missed Watt (1962), Andrewartha and Birch (1973), and Tamarin (1978). The 50-page “Afterword” which she added to the second edition (1995) is more of a bibliographic guide to recent ecology history than a continuation of her original history. Robert (“Mac”) McIntosh's Background of Ecology: Concept and Theory (1985) has a 47-page chapter, “Population Ecology.” McIntosh provided a good bibliographic guide and arguments on both sides of issues, mostly without taking sides. Naomi Cappuccino (1995) and Peter Turchin (1995) wrote brief retrospective articles in the same volume, as background for taking population studies in new directions. Mark Hixon, Stephen Pacala, and Stuart Sandin (2002) utilized Egerton (1973), Kingsland (1985), Cappuccino (1995), and Turchin (1995), but neither Andrewartha and Birch (1973) nor Tamarin (1978) in their perceptive “Population Regulation: Historical Context and Contemporary Challenges” (2002). Giorgio Israel and Ana Millán Gasca compiled an important supplement to Scudo and Ziegler's Golden Age in The Biology of Numbers: The Correspondence of Vito Volterra on Mathematical Biology (2002), which includes a dozen of his correspondents who have significance for the history of ecology. Israel and Millán Casca also provided biographical sketches of each correspondent. Peter Price's chapter, “Historical Views on Distribution, Abundance, and Population Dynamics” (2003:9–47), is an important concise survey emphasizing insects, written using Thompson (1939), Tamarin (1978), Cappuccino (1995), Turchin (1995), but neither the article by Andrewartha and Birch (1973) nor by Hixon et al. (2002) nor Kingsland (1985). Daniel Botkin is a skeptic whose book, The Moon in the Nautilus Shell: Discordant Harmonies Reconsidered (2012), contains historical discussions and cited Hutchinson (1978), but not other above-named sources. Tim Birkhead, Jo Wimpenny, and Bob Montgomerie provide a valuable chapter (10) in their history of modern ornithology (2014:355–387) on population studies, emphasizing the controversy between David Lack and Vero Wynne-Edwards. That controversy is also discussed below, as is the one in Australia between Nicholson and Andrewartha and Birch. Controversies highlight alternative explanations for phenomena, and their resolution advances science. The history of population ecology from antiquity to Darwin is surveyed briefly by Cole (1954:105–117, 1958:2–11), in much greater detail by Egerton (1967, 2012), partly by Hutchinson (1978:1–46), and is beyond the scope of this essay. Various ecologists have seen the scope of population ecology in slightly different ways. As a practicality, I focus upon certain topics and neglect others, but without intentionally defining the scope of this subject, which is the prerogative of ecologists, not historians. The existence of invasives was noted long before it became a scientific question. When Spanish conquistadores invaded Mexico and Peru in the early 1500s, they inadvertently transmitted European diseases to natives, which diseases quickly decimated the contacted populations (Crosby 1972:35–63, McNeill 1976, Bray 1996). When African slaves were imported into the Americas, they brought yellow fever and malaria. These catastrophes were also repeated in other parts of the world that Europeans penetrated, either as traders or conquerors. Additionally, Europeans noticed that local species on islands to which they brought livestock and (accidentally) rats tended to disappear. However, such early observations did not become the basis of scientific inquiry. The Caribbean Islands, once taken from natives, became sites of slave plantations that raised highly profitable crops, especially sugar cane. Unfortunately, European ships also accidentally imported brown and black species of rats, which literally ate away at the profits. In 1816, “a Jamaican horticultural publication speculated that the mongoose might ‘…extirpate the whole race of the vermin’” (quoted in Laycock 1966:111). Only in 1872 did sugar-cane grower W. B. Espeut obtain from India and release four wild males and five wild females into his fields. Espeut was pleased with the effectiveness of his mongooses, and in 1877, Jamaican stock was introduced into Puerto Rico, with others soon sent to many other islands. A problem: mongooses are diurnal, rats are nocturnal; therefore, these predators varied their diet to include diurnal prey, especially as rats became scarce (Laycock 1966:114). Poultry was, for mongooses, a satisfactory alternative to rats, but not for poultry owners. Whenever poultry was protected, native island animals, which had evolved without predators, were an easy alternative. A scientific context emerged in 1887 at the Fruit Growers Convention meeting at Riverside, California, where USDA entomologist Charles Riley (1843–95) was asked to address the crisis of a severe infestation of the cottony-cushion scale Icerya purchase in California citrus orchards (Egerton 2012:190–192, 2013:69–70). Riley told the convention that the pest was accidentally imported from Australia, and that if growers lobbied Congress for funds, he would send an investigator to discover and import enemies of this citrus pest. Funds became available, Riley sent Albert Koebele to Australia, and he succeeded in finding both a vedalia beetle that preys on this scale insect and a fly that parasitizes the scales. These enemies were imported and they controlled the scales. This was beginners luck; not only were later enemies of foreign pests often ineffective when imported, but sometimes the effort backfired (as mongooses eating poultry). Agricultural pests were a problem that prompted some biologists to study insect population dynamics. In the year 1897, three significant publications by entomologists concerned insect populations. Most mathematically ambitious was a study on “Les plantations de pins dans la Marne et les parasites qui les attaquent,” by Ad. Bellevoye, member of Société Entomologique de France, and J. Laurent, Professor at the Lycée and at l'Ecole de Médecine de Reims. Their monograph offered hypothetical calculations on the population dynamics of a bee and its parasite for a period of three years (1897:101–103, 1977). Appearing in a regional science journal, it would have attracted little attention of biologists elsewhere, unless publicized. It is impossible for me adequately to express my admiration for Marchal. Following a correspondence beginning in 1894, I have known him personally since 1902, have often visited him in his laboratory and in his home, and spent the better part of three months with him traveling in the United States. Marchal published a brief note, “L'équilibre numérique des espèces et ses relations avec les parasites chez les insectes” (1897), arguing that oscillations in populations of crop pests—Hessian fly, oat midge, army worm—was due to increase in their parasite population that followed increase in host population. When host populations crashed, parasite numbers also crashed, which then allowed hosts to increase again. Marchal called attention to Bellevoye and Laurent's paper to a broad audience by summarizing their mathematical argument in his longer “Utilization des insects auxiliaires entomophages dans la lutte contre les insects nuisibles à l'agriculture” (1907, English 1908:353–354, 1977). (a) Paul Marchal. Howard 1930: plate 19. (b). Leland Ossian Howard. Web. Third, USDA entomologist Leland Howard wrote “Study in Insect Parasitism: A Consideration of the Parasites of the White-marked Tussock Moth” (1897, 1977). Illinoisan Howard (1857–1950) had B.S. and M.S. degrees from Cornell University (1877, 1883) and had become Chief, USDA's Bureau of Entomology in 1894 (Howard 1933, Graf and Graf 1959, Mallis 1971:79–86, Hatch 1972, Sawyer 1999, Egerton 2013:73–74). He undertook this study because in 1895, “Washington suffered from an extraordinary outbreak” of Orgyia leucostigma (Howard 1897:6). Previously, the tussock moth had mainly attacked fruit trees, but after the English Sparrow spread throughout American cities, it had rid city shade trees of cankerworms, and tussock moths seemingly filled the void. The Washington, D.C., outbreak motivated Howard to study the known and potential parasites of tussock moths. In 1896, however, Washington did not experience a repeat of the 1895 outbreak, and instead of crediting the English Sparrow, he credited all those parasites of tussock moths he had just studied. Alfred Lotka (1925:90–91) quoted Howard 1897:40 and provided a graph that illustrated Howard's account. discuss[ed] the natural causes of mortality in insect populations. They divide this mortality into two large categories: “catastrophic” and “facultative.” Catastrophic refers to factors that destroy a constant percentage irrespective of the abundance of the form. Facultative refers to factors that destroy a percentage increasing as the density increases. Their study attracted broad attention, and the earliest extract in Tamarin's sourcebook, Population Regulation (1978:31–35), is from Howard and Fisk (1911:105–109). Tamarin commented (1975:29): “They believed in the balance of nature, a view prevalent among those who believe in density-dependent population regulation.” USDA entomologist Dwight Pierce (1881–1967), stationed at Dallas, Texas, led a team of three who investigated enemies of the boll weevil; their report of 1912 was similar to Howard and Fiske's in describing many boll weevil parasites, but it went further in publishing a notable diagram, “The Boll Weevil Complex,” which focused upon the cotton plant, indicating both parasites and hyperparasites (reproduced in Egerton 2007:54, 2014a:68; on Pierce and Hood, see Egerton 2014a:71). Simultaneous with studies on insect pests were studies on human predation. About 1880, the population of a subspecies of willow grouse, the red grouse Lagopus lagopus scoticus, in Scotland, began to decline, and concern over this popular game species motivated Lord Lovat to establish a Committee of Inquiry on Grouse Disease, whose report (Lovat 1911) became “one of the first population studies of birds,” (Birkhead et al. 2014:372). Another practical human predation problem was annual fluctuations in commercial catch of fish. Norwegian fisheries zoologist Johan Hjort (1869–1948), whom we met in part 51 (Egerton 2014b:397–401), was head of the Norwegian government fisheries department, 1900–1916, and since annual fluctuations in fish populations were significant enough to affect the livelihood of commercial fishermen, he studied the problem (Schlee 1973:222–229, Smith 1994). Government catch statistics on cod began in Norway in 1860, when nearly 24 million were caught. A few years later the catch dropped to about 11 million, and in mid-1890s the catch shot up to about 40 million. Hjort investigated several environmental factors and found a decisive correlation with the abundance of phytoplankton, which surely pleased German zoologist Victor Hensen, whose rather different research led him to a similar conclusion (Egerton 2014b:236). Hjort (1914) suspected that year classes that began with abundant phytoplankton could be identified in the catch for several years, and his study of annual growth lines on fish scales verified that. The enthusiastic American zoologist Raymond Pearl (1879–1940), from New Hampshire, and a graduate of Dartmouth College (B.A., 1899) and the University of Michigan (Ph.D., 1902), gradually moved from the study of invertebrate physiology to genetics to human demography (Jennings 1943, Parker 1974, Kingsland 1985:56–76, Acker 1999). Pearl notably championed the logistic (sigmoid) curve as a mathematical tool to describe human population increase (Kingsland 1982). He developed this methodology unaware that the Belgian mathematician Pierre-François Verhulst (1804–1849) had achieved the same insight in three papers: 1838, 1845, 1847 (Kormondy 1965:64–69, Hutchinson 1978:16–20). The United States had taken a national census every decade since 1790, which provided data that Pearl and his mathematician colleague Lowell Reed used for their study, “On the Rate of Growth of the Population of the United States since 1790 and Its Mathematical Representation” (1920). However, their graph of population growth included a projection into the future to about the year 2000, which illustrated Pearl's tendency to think much beyond his data. Edwin Bidwell Wilson (1879–1964), a Harvard physicist, became a critic of Pearl's work (Kingsland 1985:87–95). The outcome of their controversy, and a lesser one between Pearl and Royal Chapman, was a more sophisticated appreciation of quantifying population questions (Kingsland 1985:96–97). Annual catch of cod from Georges Bank, 1883–1967. Graham 1970:250. In the 1920s, animal ecologists were not highly trained in mathematics, and two science-oriented mathematicians, American Alfred Lotka (1880–1949), and Italian Vito Volterra (1860–1940), in mid-1920s and later, published explanations of how mathematics could be used to calculate animal population dynamics (Whittaker 1941, Scudo 1971, 1984, Gridgeman 1973, Volterra 1976, Scudo and Ziegler 1978, Kingsland 1982, 1985:25–49, 102–116, 1991:7–10, Fuchsman 1999). Pearl recruited Lotka to his Johns Hopkins laboratory for two years while Lotka wrote Elements of Physical Biology (1925). Volterra became interested in population dynamics by the influence of his soon-to-be son-in-law Umberto D'Ancona, who was studying Adriatic fishery statistics (Scudo 1971:2, 1984:23–26, Israel 1993:487–492). In 1926, the year after Lotka's book appeared, but without knowledge of it, Volterra published the first of his population studies in Italian, with summary in English (Volterra 1926a, b). The longer version is now also available in two English translations (Volterra 1931a, 1978). Kenneth Watt (1962:245–247) provided a critical discussion of strengths and weaknesses of Volterra's longer 1926 article, and found: “all the conclusions which Volterra draws out in a very thorough monograph are perfectly valid deductions from the assumptions he makes. However, the assumptions are not drawn from biological reality.” Reality would involve more variables than could be conveniently added to the calculations. Lotka saw Volterra's brief summary article (1926b) in Nature and immediately wrote a letter to Nature dated 29 October 1926 with a claim of priority, which the editor obviously sent (or a copy) to Volterra, since Lotka's letter and Volterra's reply both appeared in the first issue of 1927 (Lotka 1927, Volterra 1927). Both letters were reprinted by Israel (1993:496–497). But Lotka also wrote directly to Volterra. Between 2 November 1926 and 19 January 31, Lotka sent five letters to Volterra and Volterra sent three replies that are now also published (Israel and Millán Gasca 2002:280–288). Lotka wrote his first letter in French and later ones in English. Volterra wrote his replies in Italian. Their correspondence was entirely cordial, and the last letter was from Lotka thanking Volterra for sending him a copy of his Lecons sur la théorie mathématique de la lutte pour vie (1931). Egbert Leigh (1968) discussed at length the virtues of this treatise. In 1926, Volterra was already a well-established leading mathematician, whose work attracted much more attention than did Lotka's. Scudo (1984:20) suggested that Lotka's book was too broad in scope: “Possibly by having thus overwhelmed most readers it was on the whole a flop, as Lotka bitterly lamented shortly after in letters to Volterra.” Kingsland (1985:211), however, suggested that Lotka had published too early, and lacked a biological audience, though entomologists William Thompson, William C. Cook, and Royal Chapman would soon appreciate the utility of Lotka's work (see below). A major concern for Volterra was reconciliation of mathematical calculations and theory with real biological data. He began with a real fisheries question, and he persisted in comparing mathematical calculations with biological data (Millán Gasca 1996:5–6, page numbers of reprint, 2002). Only three of the population ecologists with whom Volterra communicated were mathematically adept—Thompson, Gause, and Kostitzin--a Canadian and two Russians. The only theoretical biology Volterra knew was Darwin's theory of evolution by natural selection. The ecologist who wrote to Volterra wrote about real problems rather than theoretical problems. Volterra's major collaborator on biological mathematics was Russian Vladimir Kostitzin (1882/1883–about 1963), who had been a revolutionary in early 1900s (Scudo and Ziegler 1976:396, Scudo 1984:31–32). Kostitzin's wife Julie, a parasitologist, moved to Paris in mid-1920s, and he followed her by 1927. In 1931, Volterra refused to sign a loyalty oath imposed by Italy's Fascist government on professors, and he moved to Paris (Volterra 1976:86), where he lectured on animals struggling for existence, which attracted Kostitzin's interest. Kostitzin had already coauthored with his wife a mathematical analysis of a parasitic relationship between hermit crabs, Eupagurus cuanensis, and parasitic barnacles, Chlorogasdter sulcatus (Kostitzin and Kostitzin 1931). In 1934 Kositzin published Symbiose, Parasitisme et Évolution, followed by Biologie mathématique (1937, English, 1939). He wrote more papers than anyone else in Scudo and Ziegler's Golden Age of Theoretical Ecology (1978). The published Volterra–Kostitzin correspondence covers 42 pages (Israel and Millán Gasca 2002:225–266); however, Volterra died in 1940, and thereafter Kostitzin corresponded with Volterra's widow, Victoria (9 of those 42 pages), until 1962. Julie Kostitzin had died in 1950. Volterra's approach was strictly adherent to the classical physico-mathematical paradigm, while Lotka's point of view was more eclectic and open-minded as regards the new developments of physics, and somewhat skeptical about formal statements….while Volterra followed the ‘mechanical analogy’ Lotka had an inclination for the ‘thermodynamic analogy’ and had a great interest for the energetic problems in population dynamics. Israel quoted from two of Voltera's letters to Lotka, with English translations in footnotes, and quoted a long passage from Volterra's “Les mathématiques dans les sciences biologiques et sociales,” also with English translation in a footnote. This paper showed Volterra's general interest in using quantitative methods in biological and social sciences before his future son-in-law interested him in population dynamics. Peter Wangersky's “Lotka–Volterra Population Models” (1978) is not an earlier paper on the same subject as Israel's; instead, Wangersky used their names as a general indication of his topic, which was really an explanation of mathematical population models. Israel later published an excellent discussion of how Lotka and Volterra's contributions fit into the larger picture of population dynamics in the 1920s, including a discussion of their priority dispute (1993). Russian zoologist Vladimir Alpatov, who first taught ecology in Moscow (Weiner 1988:66), gained access to several papers by Pearl and wrote to ask if he could come to Johns Hopkins University to study under him (Kingsland 1985:146–148). Pearl obtained a grant from the Rockefeller Foundation which enabled Alpatov to do so, 1927–1929. After returning to Moscow, Alpatov discussed Pearl's work with one of his students, Georgii Gause (or Gauze, 1910–1986), who had already conducted a survey of grasshoppers in the northern Caucasus Mountains and developed a mathematical formula to study the relationship between species abundance and environmental factors, which appeared in Ecology (Gause 1930). I am working in the field of experimental biology, and I am not qualified enough to analyze more closely the mathematical part of the problem and it would be very interesting if it will be possible for you to investigate these equations in the future, and to find their solution in the general form. (a) G. F. Gause. Web. (b) Growth in volume of Saccharomyces cerevisiae, Schizosaccaromyces kephir and mixed population, aerobic (above) and anaerobic (below). Gause 1934:85. Volterra responded in Italian in an undated letter. Volterra did collaborate with a few colleagues, but Gause seems not to have been one of them. In a letter of 12 October 1933, Gause wrote to Volterra telling of his pending book, The Struggle for Existence, and that he had confirmed experimentally some of Volterra's mathematical projections, and requesting any papers he had published since 1931 (Israel and Millán 2002:213–214). Gause had confirmed the results of competition between two similar species for the same resource, leading to the extinction of one species (Gause 1934b:89, 113), now called Gause's axiom, “the competitive exclusion principle.” We expected at the beginning of this chapter to find “classical” oscillations in numbers arising in consequence of the continuous interactions between predators and prey as was assumed by Lotka and by Volterra. But it immediately became apparent that such fluctuations are impossible in the population studied, and that this holds true for more than our special case. Even before the book appeared, Gause published a preview note in Science (1934a:16) with the same mixed message: “Experiments on the competition between two species for a common place in the microcosm agreed completely with Volterra's theoretical equations, but as regards the processes of one species devouring another our results are not concordant with the forecasts of the mathematical theory.” His book became “a foundation stone of ecology” (Hutchinson 1978:120). It is fortunate that Gause had already made arrangements to publish in America, because in January 1934, at an Ecological Conference of the Academy of Sciences' Botanical Institute, a fanatical Stalinist ecologist, I. I. Prezent, denounced Alpatov and Gause for their use of mathematics in their “formalist, mechanist school” (quoted in Kingsland 1985:160–161, Weiner 1988:222). Subsequently, it might have been difficult for Gause to publish that book in the USSR. Gause continued for some years to publish papers in western journals, but he also decided it would be expedient for him to forget about going to America for a few years and instead do research in microbiology. Two ecologically oriented entomologists, Canadian William Thompson and American Royal Chapman, were receptive to mathematical assistance from Lotka and Volterra (Graham 1941, Thorpe 1973, Kingsland 1985:123, 127–131). Thompson (1887–1972), from London, Ontario, earned his B.A. degree at the University of Toronto (1909). In summer 1908, he worked at the USDA Gypsy Moth Parasite Laboratory (discussed above) and in 1909 the USDA sent him to Cornell University to study for a M.S. degree (1911) under Professor Comstock. In 1912, USDA sent him to Professor Filippo Silvestri's Instituto Superiore Agrario, Portici, Italy. In 1913 he resigned from USDA and went to Paris to study under Professor Maurice Caullery. In 1914, he went to the Zoology Department, Cambridge University, to study bacteriology and protozoology. During World War I, he joined the Royal Naval Medical Service and became an assistant in a clinical laboratory. In 1918, he returned to Paris and resumed studies of dipterous parasites, and earned a Ph.D. degree (1921). In 1919, he had married artist Mary Carmody, who became an insect illustrator and illustrated his papers. Leland Howard, at USDA, re-employed Thompson in 1919 and put him in charge of a European Parasite Laboratory, which he established at Hyères and ran for nine years. In 1929 Thompson returned to England to run the Farnham House Laboratory to study insect parasites and predators for the British Empire. During World War II, Farnham Laboratory closed and he returned to Canada as Director of the Imperial Parasite Service, Canadian Department of Agriculture. In 1948–1958 he established biological control laboratories in California, Trinidad, Uruguay, Switzerland, India, and Pakistan. Thompson wrote seven letters, published posthumously, to Volterra, 29 July 1927–23 March 1931, but there is only one undated letter in response from Volterra (Israel and Millán Gasca 2002:368–373). Both wrote in French except for Thompson's last letter in English. With his first letter, Thompson sent a copy of several of his published papers and requested in return a copy of Volterra's longer 1926 paper, in Italian. Volterra responded, thanking him for sending his very interesting works and explained that his copies of that article were all gone, but that more copies were being printed and he would soon send one. On 23 August 1927, Thompson thanked him for a letter of August 16 and stated he hoped to publish in winter a small book on insect parasitology. On September 3, Thompson wrote to thank Volterra for sending him “votre grand travail publie par l'Academie dei Lincei.” In a letter of 5 February 1931, Thompson thanked Volterra for sending him a copy of his Leçons sur la théorie mathématique de la lute pour la vie (1931) and lamented that his own mathematician collaborator, Soper, at the College of Tropical Medicine, had died, though they had a joint paper in press. Culminating Thompson's researches was “Biological Control and the Theories of the Interactions of Populations” (1939), with a 27-page historical survey, beginning with Charles Darwin and including his own researches. He discussed Lotka and Volterra, but was wary of zoologists who substituted mathematical calculation for few field data (1939:381): “mathematical construction is so elegant and impressive that there is already a tendency to overestimate its value and to offer it as a description of what really happens in nature….” He had recently published a polemic, Science and Common Sense: an Aristotelian Excursion (1937), which made this point as well as attacking Darwin's theory of evolution by natural selection (Kingsland 1985:135–141). Howard and Fiske's 1911 concepts of “catastrophic” and “facultative” causes of mortality for insects had been useful, but Thompson (1928) rejected their terms in favor of “general” and “individualized.” Yet, Harry Smith (1935:889) decided that Thompson's terms were not much improvement, and he coined “density-independent” for weather causes and “density-dependent” for biotic causes of mortality. Smith's terms were accepted for some time, but Dempster (1998) suggested population limitation rather than regulation, and Berryman et al. (2002) urged a “requiem for density dependence,” and Price et al. (2011:357) concluded that “population is regulated at some times, and not at others…” (a) William Robin Thompson. Thorpe 1973. (b) Royal Norton Chapman. Web. Minnesotan Royal Chapman (1889–1939) earned his B.A. and M.A. degrees (1914, 1915) at his state university and his Ph.D. (1917) at Cornell University. He returned to the University of Minnesota as a faculty member and developed an ecology course—the first at that university. His course emphasized “controlled experiment and quantitative field studies” (Graham 1941:522). In his Ecology paper, “The Quanti" @default.
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• W2190647748 title "History of Ecological Sciences, Part 55: Animal Population Ecology" @default.
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