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Functionalism without Selectionism: Charles Elton's "Functional" Niche and the Concept of Ecological Function

  • THEMATIC ISSUE ARTICLE: CONCEPTUALIZING THE ENVIRONMENT IN NATURAL SCIENCES
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Abstract

This article offers an analysis of ecologist Charles Elton’s “functional” concept of the niche and of the notion of function implicitly associated with it. It does so in part by situating Elton’s niche concept within the broader context of the “functionalist-interactionist” approach to ecology he introduced, and in relation to his views on the relationship between ecology and evolution. This involves criticizing the common claim that Elton’s idea of species as fulfilling functional roles within ecological communities committed him to an idea of communities as units of selection. While such a claim implicitly attributes to Elton an understanding of function along the lines of the selected-effects theory of function advocated by many biologists and philosophers of biology, Elton’s use of the niche concept, I maintain, involves an understanding of function more along the lines of alternative nonselectionist theories such as the causal-role, goal-contribution, and organizational theories. I also briefly discuss how ecologists after Elton also tend to have typically adopted a nonselectionist understanding of the function concept, similar to his.

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Fig. 1

reproduced from Summerhayes and Elton 1923, p. 232)

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Notes

  1. For historical discussions of the niche concept, see David Cox (1979, chap. 4), Thomas Schoener (1989), James Griesemer (1992), Mathew Leibold (1995), and Arnaud Pocheville (2015).

  2. For reviews of philosophical theories of function, see, e.g., Peter McLaughlin (2001), Arno Wouters (2005), Denis Walsh (2008), Christian Saborido (2014), and Justin Garson (2016).

  3. As Elton himself reports, he and Grinnell “independently started using the term” (see Elton and Miller 1954, p. 477).

  4. For reviews of niche theory, see, e.g., Schoener (1989, pp. 97–102) and Pocheville (2015, sec. 1.4).

  5. His focus on animals reflects only the fact that he developed the concept in the context of animal ecology.

  6. Elton’s analogies between animal communities and human societies were presumably influenced in part by sociologist Alexander Carr-Saunders (1886–1966), who is known to have acted as Elton’s mentor at Oxford (together with zoologist Julian Huxley). Elton’s construal of ecology as “the sociology and economics of animals” is explicitly linked to Carr-Saunders’s (1922) study of the “sociology and economics” of humans in the preface of Animal Ecology (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, Chap. 3). On functionalism in sociology in general, see Munch (1976), Moore (1978), and Bigelow (1998).

  7. Elton, however, acknowledged the importance of intraspecific competition as a regulative factor in later publications (e.g., Elton 1958, p. 131; see Haak 2000, p. 30). On Elton’s ideas on population regulation, see also Cox (1979, pp. 50–64), Sheail (1987, sec. 2.5.2), and Hagen (1992, pp. 56–60).

  8. For the sake of comprehensiveness, it should be mentioned that Elton considered animal populations to be regulated not only through the intracommunity factors just described, but also through climatic factors (Elton 1927, pp. 119, 123), and through the migration of animals from more populated to less populated areas (Elton 1930, pp. 61–62; 1933, pp. 70–72).

  9. Elton’s occasional description of some animals’ niches in relation to abiotic factors evokes the possibility of complementary ecological functions based on species’ impacts on abiotic factors (see Schoener 1989, p. 86; Griesemer 1992, p. 234; Leibold 1995, p. 1373). For instance, he described land crabs living in burrows on coral islands and earthworms elsewhere as occupying the same (soil-burrowing) niche (Elton 1927, p. 67). This suggests the possibility of ecological functions associated with what we would now call ecosystem engineering (Jones et al. 1994; Berke 2010). This idea, however, is not fully developed in Elton’s works.

  10. Kimler bases this reading partly on personal communication with Elton (see Kimler 1986, p. 224).

  11. At variance with Kimler and Hagen, and more in line with what I will maintain below, David Cox (1979, pp. 5–6) interprets Elton as an individual-level selectionist (see also Haak 2000, p. 32).

  12. Historian of ecology Peder Anker (2001, p. 112) notes that Wells et al.’s ecology chapter was entirely reviewed by Elton before publication.

  13. For discussions of Elton’s ideas on the relationship between ecology and evolution, see Scudo (1984, pp. 41–44), Kingsland (1985, pp. 53–55), Hagen (1992, pp. 59–61), Haak (2000, pp. 31–32), Cooper (2003, pp. 47–48), and Holt (2005, pp. 242–245).

  14. Elton also develops some of those ideas in the final chapter of Animal Ecology (Elton 1927, Chap. 9), as well as in papers published before and after Animal Ecology and Evolution (Elton 1924, 1931).

  15. Contemporary ecologists study this latter phenomenon under the term “habitat selection” (see, e.g., Morris 2003), and niche construction theorists refer to it as “relocational niche construction” (see Odling-Smee et al. 2003, pp. 44–47, 123–124).

  16. In a later paper, Elton even drew attention to the phenomenon of cultural evolution, arguing that “among the higher animals we can perceive a method of evolution along a mental plane, unconnected with the spread of new mutations, a method which leads on a small scale to the production of customs, cultures, and gregarious habits, similar to those found in man” (Elton 1931, p. 134; quoted in Hagen 1992, pp. 60–61).

  17. It is important to note that Elton’s rejection of this particular explanation for community stability did not amount to a wholesale rejection of the idea that communities achieved some kind of stability. Elton fully admitted that communities have “some power of regulation, of compensating here for a disturbance there,” and he remarked that, “the species composition of most communities remains very much the same over long periods” (Elton 1930, pp. 38, 25). His main point was that community stability is achieved through other factors, such as the migration of animals from more to less populated areas. This is partly why, above, I say that Elton rejected “a version of” the balance of nature idea. This is meant to leave open the possibility that his views may be ones that other ecologists would interpret as aligning with the balance-of-nature idea. For discussions of Elton’s ideas on community stability and the balance of nature, see Hagen (1992, pp. 56–60), Haak (2000, pp. 26–29), and Dussault (2020, sec. 4.3). For an analysis of the various views that ecologists have associated with the balance-of-nature idea, see Jansen (1972).

  18. This implication may seem to be challenged by Roberta Millstein’s (forthcoming) recent defense of the possibility of selected-effects ecological functions based in coevolution rather than community or ecosystem-level selection. Millstein’s discussion is of particular relevance here, given its focus on Aldo Leopold and given Elton’s known influence on Leopold. Millstein’s discussion, however, seems implicitly focused on a different notion of function than the one I am focusing on here. As she recognizes (footnote 1), her discussion is focused on function as a type of activity achieved by organisms across communities (e.g., predation, parasitism). In contrast, I am here concerned with function as a type of contribution to the capacities or activities of a larger system (e.g., as highlighted above, the transfer of organic matter through the food cycle, the regulation of animal populations involved within the community). Notwithstanding Millstein’s arguments, a selected-effects interpretation of the latter functions would still seem to require community-level selection.

  19. Other philosophers (e.g., Wright 1973, pp. 147–148; Boorse 1976, p. 76) make essentially the same contrast by distinguishing the notion of what the function of x is (Achinstein’s design functions) from the notion of what x functions as (Achinstein’s use and service functions). Wright, however, restricts the legitimate sense of function to the first notion, which, as regards biological functions (as opposed to technical functions), he analyzes in selected-effects terms.

  20. As some historians of ecology have noted, 20th-century ecologists tended to use organicist and mechanistic analogies interchangeably (see Hagen 1992, pp. 11–14; Marshall 2002, Chap. 7). Elton, for his part, at times criticized the mechanistic analogy (see Elton 1930, p. 30) and, as far as I am aware of, never committed to the organicist analogy.

  21. I leave it open whether, from a functionalist sociological perspective, humans can soundly be depicted as fulfilling social functions within their societies. See note 6 above for references on functionalism in sociology.

  22. For a more detailed comparison of Elton’s ideas on ecological communities and those of ecologists who construed communities by close analogy with organisms, see Dussault (2020).

  23. In more contemporary parlance we could say that ecological communities were, for Elton, functionally organized entities by virtue of being niche-construction networks—that is, networks of species that, by their simple presence or their effect on the environment, provide key conditions of existence to each other (see Odling-Smee et al. 2003; Barker and Odling-Smee 2014).

  24. For an application of the organizational theory of function to the context of ecology, see Nunes-Neto et al. (2014).

  25. For the sake of exactness, it should be mentioned that one of the nonselectionist theories mentioned above—namely the organizational theory—is intended to account as much for the teleological (the why) as for the systemic (the how) dimensions of the function concept (see, e.g., Mossio et al. 2009, sec. 4.1). Nevertheless, what has primary relevance for my purposes is how, by accounting for the systemic dimension, the organizational theory, like other nonselectionist theories, can shed light on Elton’s nonselectionist use of the function concept.

  26. On Mayr’s contrast between why and how questions and its relationship to the use of the function concept in ecology in general, see Hagen (1992, pp. 150–151).

  27. It should be noted, however, that, although Dice seems to have shared Elton’s nonselectionist understanding of function, he did not associate the niche concept itself with functions fulfilled by species within communities (see Dice 1952, p. 227). Like most ecologists, he followed Grinnell in adopting a niche concept focused on species’ ecological requirements (in his case, mainly the resources that species require) (for a discussion, see Hurlbert 1981).

  28. See however Millstein (forthcoming) for a dissenting view.

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Acknowledgments

The author is thankful to the audience of the session “L’environnement dans la biologie du XXIe siècle: Ambiguïtés conceptuelles et défis épistémologiques,” held at the 2018 Société de philosophie des sciences conference in Nantes, France, where an earlier version of this paper was presented, as well as to 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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Dussault, A.C. Functionalism without Selectionism: Charles Elton's "Functional" Niche and the Concept of Ecological Function. Biol Theory 17, 52–67 (2022). https://doi.org/10.1007/s13752-020-00360-9

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