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Neither superorganisms nor mere species aggregates: Charles Elton’s sociological analogies and his moderate holism about ecological communities

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Abstract

This paper analyzes community ecologist Charles Elton’s ideas on animal communities, and situates them with respect to the classical opposition between organicist–holistic and individualistic–reductionist ecological views drawn by many historians of ecology. It is argued that Elton espoused a moderate ecological holism, which drew a middle way between the stricter ecological holism advocated by organicist ecologists and the merely aggregationist views advocated by some of their opponents. It is also argued that Elton’s moderate ecological holism resonated with his preference for analogies between ecological communities and human societies over more common ones between communities and individual organisms. I discuss, on the one hand, how the functionalist-interactionist approach to community ecology introduced by Elton entailed a view of ecological communities as more or less self-maintaining functionally organized wholes, and how his ideas on this matter were incorporated into their views by organicist ecologists Frederic Clements, Victor Shelford, and Warder C. Allee et al. On the other hand, I identify some important divergences between Elton’s ecological ideas and those of organicist ecologists. Specifically, I show (1) how Elton’s ideas on species distribution, animal migrations, and ecological succession entailed a view of animal communities as exhibiting a weaker degree of part-whole integration than that attributed to them by Clements and Shelford; and (2) how Elton’s mixed stance on the balance of nature idea and his associated views on community stability attributed to communities a weaker form of self-regulation than that attributed to them by Allee et al.

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Notes

  1. It is noteworthy that the two most detailed treatments of Elton’s ideas on communities (and his related ideas on population regulation) are found in unpublished doctoral dissertations (i.e., Cox 1979; Haak 2000). The below discussion owes a lot to those treatments, as well as to the more succinct ones offered by McIntosh (1985), Sheail (1987), and Hagen (1992).

  2. For Elton’s own account of how he was brought to this shift, see Elton and Miller (1954, pp. 461–462).

  3. As pointed out to me by Kurt Jax, an exception to this general observation is Karl August Möbius (1825–1908), whose picture of ecological communities closely anticipates Elton’s functionalist and interactionist picture. Möbius is best known for his work on oyster banks (Möbius 1883), where he introduced the term “biocenosis” to denote ecological communities. On Möbius’s ideas on ecological communities, see Glaubrecht (2008, pp. 17–18), van der Valk (2017, pp. 115–116), Schwarz and Jax (2011, pp. 242–244), and Potthast (this issue).

  4. For detailed discussions of this earlier classificatory and habitat-focused approach, see Griesemer (1990) and Neumann (2017).

  5. For a detailed history of ideas on food chains and food webs, see Egerton (2007).

  6. The Matamek conference gathered scientists from various fields and was organized by wealthy fisheries trader and nature lover Copley Amory on his private estate (for report and proceedings, see Huntington 1932).

  7. For historical discussions of the niche concept, see Cox (1979, Chapter 4), Schoener (1989), Griesemer (1992), Leibold (1995), and Pocheville (2015).

  8. In this respect, Elton’s niche somewhat anticipated the concept of niche construction (Odling-Smee et al. 2003). Elton, however, in contrast to niche construction theorists, seems to have conceived the niche not so much as what is constructed by organisms (the result of construction), but rather as the constructing activity itself.

  9. A most likely inspiration for Elton’s parallels between animal communities and human societies is sociologist Alexander Carr-Saunders (1886–1966), who, together with Julian Huxley, is considered to have acted as his mentor at Oxford. In the preface of his Animal Ecology, Elton explicitly links his understanding of ecology as “the sociology and economics of animals” to Carr-Saunders’s (1922) study of the “sociology and economics” of humans (Elton 1927, p. vii). On the relationship between Elton and Carr-Saunders, see Sheail (1987, p. 90), Hagen (1992, pp. 56–57), and Anker (2001, Chapter 3). For general discussions of functionalism in sociology, see, e.g., Munch (1976) and Moore (1978).

  10. For detailed discussions of Elton’s ideas on animal communities, see Cox (1979, pp. 51–54, Chapter 3) and Hagen (1992, pp. 51–62).

  11. Interestingly, Elton’s ideas, in this respect, aligned with those of Carr-Saunders, who criticized strict Malthusian views on population regulation (see Carr-Saunders 1922, Chapter 9; and for discussions, see Sheail 1987, p. 90; Angner 2009, p. 81). It should be noted, however, that, in later publications (Elton 1958, p. 131), Elton recognized a more important regulative role to intraspecific competition (see Haak 2000, p. 30). For general discussions of Elton’s ideas on population regulation, see Cox (1979, pp. 50–64), Sheail (1987, sec. 2.5.2), Hagen (1992, pp. 56–60), and Haak (2000, pp. 26–30, 43–44).

  12. Elton restated this idea in later publications (e.g., Elton and Miller 1954, p. 463). The distinction between autecology and synecology invoked by Elton in the latter passage was from Swiss ecologists Schröter and Kirchner (1902) and draws a contrast between ecological studies focusing on single species in relation to their environment (autecology) and studies concerned with ecological communities as a whole (synecology). Elton presumably took this distinction from his Oxford colleague Tansley (1923, p. 20).

  13. For historical discussions of organicist views in early ecology, see Bodenheimer (1957), Acot (1987), McIntosh (1998), and Bergandi (1999).

  14. Additional indirect support may be found in the fact that the idea of communities as superorganisms was espoused by Elton’s mentor Julian Huxley, along with novelist H. G. Wells, in the ecology chapter of their book The Science of Life, a huge book that surveyed life sciences for a popular audience. In this book, Wells and Huxley asserted that “it is not altogether fanciful to compare all [the world’s] various [ecological] communities, each one to a sort of super-organism, and in practice much of the thought and work of the ecologist involves that idea” (Wells et al. 1931, p. 578). According to historian of ecology Anker (2001, p. 112), Wells et al.’s chapter on ecology was entirely reviewed by Elton prior to its publication.

  15. For further discussions of Clements’s ideas on ecological succession, see Whittaker (1974), McIntosh (1980, 1985, pp. 76–82), Hagen (1988, 1992, Chapter 2), and Eliot (2007, 2011).

  16. The concept of homeostasis was coined by Cannon (1932) and denoted organisms’ characteristic ability to maintain their state in response to external perturbations. It was inspired by the work of the French physiologist Bernard (1879).

  17. The “quasi-organism” notion alluded to here was from plant ecologist Tansley (1920, 1935).

  18. Elton and Tansley and are known to have been in relatively close contact with each other (see Anker 2001, Chapter 3). Moreover, Elton’s Animal Ecology cites two books by Tansley, and the discussion of ecological succession in chapter 3 of that book draws heavily on Tansley’s works. Hence, I take it that Elton was likely aware of Tansley’s remarks on the organicist analogy.

  19. For discussions of this aspect of Clements and Shelford’s ecology, see Whittaker (1957, pp. 197–198), and Kirchhoff (this issue).

  20. On Clements’s Lamarckianism, see Hagen (1983).

  21. In the interest of not misrepresenting Gleason’s ideas, it must be remarked that, despite his emphasis on contingency, he did not deny that species interactions were important factors in the organization of communities. He regarded interactions between species as important determinants of their ability to establish and maintain themselves in particular locations. Hence, although, for him, the composition of communities was largely contingent, communities nonetheless remained, to a significant extent, interactive systems. His views, in this respect, were very close to those I am attributing to Elton. On Gleason’s ecological ideas, see McIntosh (1975, 1995), Nicolson (1990), and Nicolson and McIntosh (2002).

  22. Revealingly, in a later work where he reminisces on the development of his views on species distributions, Elton explicitly acknowledges the relevance of Gleason’s “individualistic concept” (Elton and Miller 1954, p. 479; see McIntosh 1995, p. 319).

  23. For discussions of Clements’s ideas on ecological succession, see the references in note 15 above.

  24. For a discussion of the contrast between Clements’s and Tansley’s ideas on succession, see Whittaker (1974) and van der Valk (2014); for further discussion of Elton’s ideas on ecological succession, see Leibold and Wootton (2001, pp. xxv–xxvii).

  25. Historian of ecology Hagen (1992, p. 59) ventures that, living at the time of the 1929 Great Depression and having himself, in that context, lost his job at the Hudson’s Bay Company, Elton could not but be aware that self-regulative processes occurring in human societies were at best imperfect ones compared to those occurring within organisms.

  26. On Elton’s ideas on population regulation and fluctuations, see references in footnote 11.

  27. For a more detailed presentation of Elton’s critique of community selection, see Dussault (forthcoming, sec. 4).

  28. This mode of regulation is commonly referred to by contemporary ecologists as “source-sink dynamics” (see, e.g., Pulliam 1988).

  29. This mode of regulation is commonly called “switching” or “prey switching” by contemporary ecologists (see, e.g., Murdoch 1969).

  30. The contrast between strong and weak regulation I am evoking here is partly inspired by d’Hombres (2012, sec. 4). Unlike d’Hombres, however, I do not restrict strong regulation to regulation involving some kind of centralized control exerted on the parts by the whole. I instead cast the contrast as one between regulation achieved by regulative factors whose behavior has been coordinated by past natural selection, on the one hand, and regulation that arises from the uncoordinated behavior of a system’s freely interacting parts, on the other.

  31. It should also be mentioned that, although I here grant, for the sake of the argument, that Curtis and Whittaker’s studies and Williams’s critique of multilevel selection disqualified the kinds of ecological holism respectively endorsed by Clements and Allee et al., one may question whether this really is the case. For criticisms of Whittaker, see Leibold and Mikkelson (2002) and Wilson et al. (2004); for a partial rehabilitation of community-level selection, see Wilson and Sober (1989).

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Acknowledgements

The author is thankful to Kurt Jax, Roberta Millstein, and two anonymous referees for helpful comments. He also thanks Xander Selene for editing the manuscript. The work for this paper was supported by a research Grant from the Fonds de recherche du Québec—Société et culture (FRQSC, 2018-CH-211053).

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  1. Collège Lionel-Groulx, Sainte-Thérèse, QC, Canada

    Antoine C. Dussault

  2. Centre Interuniversitaire de Recherche sur la Science et la Technologie (CIRST), Université du Québec à Montréal (UQAM), Montréal, QC, Canada

    Antoine C. Dussault

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Dussault, A.C. Neither superorganisms nor mere species aggregates: Charles Elton’s sociological analogies and his moderate holism about ecological communities. HPLS 42, 25 (2020). https://doi.org/10.1007/s40656-020-00316-z

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