Abstract
In the theory and history of ecology, Frederic Clements’s theory of plant communities is usually presented as the historical prototype and a paradigmatic example of synecological organicism, characterised by the assumption that ecological communities are functionally integrated units of mutually dependent species. In this paper, I will object to this standard interpretation of Clements’s theory. Undoubtedly, Clements compares plant communities with organisms and calls them “complex organisms” and “superorganisms”. Further, he can indeed be regarded as a proponent of ecological organicism—provided that one defines ecological organicism as the interpretation of synecological units according to the model of the individual organism. However, Clements’s theory does not include the assumption that mutual dependence is a principle of the organisation of plant communities. Rather, he interprets plant communities as top-down control-hierarchical entities, in which subordinate species depend on dominant species—but not the other way around. Therefore, his theory represents what may be called ‘control-hierarchical organicism’ as against ‘mutualistic organicism’. The erroneous attribution to Clements of ‘mutualistic organicism’ might be due to an unawareness of the existence of different concepts of the organism. This unawareness results in the projection on Clements’s theory of a seemingly self-evident mutualistic concept of organism that Clements himself did not use as a basis for his theory of plant communities.
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Notes
Eliot’s (2007, 2011) diachronic interpretations are insightful and appropriate in many ways. However, he seems not to take sufficient account of the fact that Clements (but not Gleason) imagined an—admittedly ideal—state of self-stabilization in which the reactions of a plant community upon its habitat become “stabilizing or climatic” causes (which counteract further changes) as opposed to “continuing or ecesic” causes (which drive succession forward) (Clements 1916: 5; cf. ibid.: 55, 98, 350 f.). That is, Eliot does not sufficiently acknowledge that Clements’s theory includes the developmental concept of perfection, i.e. of a finite process leading towards a certain predetermined ideal state or goal, while Gleason’s theory refers to the developmental concept of incessant change, i.e. of an infinite process leading permanently away from the current state but never towards a certain ideal state (see Kirchhoff 2014b for a characterisation of these concepts).
Nevertheless, Eliot (2011: 75) develops convincing arguments against the false assertation that Clements assumes a mutualistic organisation of plant communities that exhibits the same degree of specialization and interdependence among its component species as the organs of animals or even much simpler organisms such as paramecia. However, in contrast to my own position, Eliot does not completely rule out ascribing a doctrine of mutualistic organisation to Clements—as long as it does not ascribe species-specific interdependencies but allows for intersubstitutability of functionally equivalent species (ibid.). Accordingly, contrary to my interpretation, Eliot claims that Clements’s “comparison [of plant communities] to [individual] organisms encompasses [… the view that] their component parts demonstrate interdependence acquired through historical adaptation or accommodation to one another” (ibid.: 72 f.), and concludes that “the organism comparison implies a degree of functional integration, even if it is loose” (ibid.: 76).
In this paper, I use the expressions “mutual dependence” and interdependence” as synonyms that both mean (direct or indirect) “reciprocal dependence” as opposed to “one-sided dependence”.
Emphasis of “superorganism” omitted.
Worster does not cite the source of this quotation. It might stem from Spencer (1860), although, given this, the correct quotation would be without “or injure”. I quote the whole passage from Spencer to make more evident his analogy between human societies or communities and single organisms: “Simple communities, like simple creatures, have so little mutual dependence of parts, that mutilation or subdivision causes but little inconvenience; but from complex communities, as from complex creatures, you cannot remove any considerable organ without producing great disturbance or death of the rest.”
Worster does not cite the source of this quotation either, and the same applies to his next quotation from Spencer.
Emphasis of “Clementsian community” omitted.
As Clements regards competition as constitutive of ecological communities (Clements et al. 1929: 327), he does not count pioneer stages of vegetation among communities because they do not exhibit competition.
Emphasis of “Subdominants” omitted.
I will not further discuss this difference because it is irrelevant as regards my critique of the synchronic standard interpretation of Clements’s theory.
For a conceptual distinction between “dependence” and “mutual dependence” or “interdependence” as well as the possibly “facultative” or “obligate”, “beneficial” or “necessary beneficial”, “direct” or “indirect”, “oligophilic” or “polyphilic” character of these ecological relations (see Boucher et al. 1982; Kirchhoff 2007; Eliot 2018).
In nested hierarchies, the elements of the higher level consist of the elements of the lower level, e.g. organisms of organs and organs of cells. In non-nested hierarchies this is not the case, e.g. generals do not consist of (but control) their lieutenants, who do not consist of (but control) their sergeants, and so on. For this distinction between nested and non-nested hierarchies (see Allen and Starr 1982). For the classification of control hierarchies as non-nested hierarchies (cf. Keller and Golley 2000a: 30).
This reference to Hagen is based on a hint by Antoine C. Dussault.
See Sect. 1 for Hagen’s and Eliot’s critique of this ascription.
Clements et al. erroneously indicate “94”.
Some of Warming’s statements, however, indicate an individualistic concept of plant communities rather than an organismic one. For example, he states directly before the passage cited above: “The plant-community […] is merely a congregation of units among which there is no co-operation for the common weal but rather a ceaseless struggle of all against all” (Warming and Vahl 1909: 95). These at first view apparently contradictory statements might be compatible if one assumes that Warming’s first statement does not represent a theory of mutual dependence of properties and of mutual dependence for survival (mutualistic organicism) but a theory of mutual dependence of properties only (holistic individualism; cf. Pettit 1993). For a more detailed discussion of Warming’s theory (see Anker 2011; besides Goodland 1975; Acot 1995).
I refer to the third edition (1971) of the “Fundamentals of Ecology” which was published by Eugene P. Odum alone, while the first (1953) and second (1959) edition were published by E. P. Odum in collaboration with his brother Howard T. Odum.
An encaptic hierarchy is a hierarchy in which the higher level is composed of the lower level. In the case of Odum, organs are composed of cells, organisms of organs, populations of organisms, and so forth.
See https://www.merriam-webster.com/dictionary/organism, accessed 2019-01-22.
My translation. The German-language original reads as follows: “Als das zentrale Moment, das Organismen als eine Einheit mit einem speziellen ontologischen und methodologischen Status kennzeichnet, gilt seit Mitte des 18. Jahrhunderts die Wechselseitigkeit” (Toepfer 2011: 790).
My translation. The German-language original reads as follows: “Ein Organismus ist ein materielles System aus wechselseitig voneinander abhängigen Teilen und Prozessen, das in physischer und funktionaler Hinsicht eine integrierte Einheit bildet und charakteristische Funktionen und Aktivitäten (wie Ernährung, Schutz vor Störungen und Rezeption von Umweltereignissen) aufweist. […] Die physische Einheit eines Organismus besteht in seiner materiellen Verfasstheit in einem kontinuierlich bestehenden kohärenten Körper, dessen Stoffe und Form jedoch einem Wechsel unterliegen können (durch Metabolismus und Metamorphose). Die funktionale Einheit des Systems besteht in seiner Organisation, d.h. in einem Gefüge aus Prozessen, die seine materiellen Teile erzeugen und erhalten und die damit (als Prozesse eines jeweiligen Typs) wechselseitig voneinander abhängen” (Toepfer 2011: 777).
As regards the emergence of this concept of organism in the theories of, especially, Gottfried Wilhelm Leibniz, Immanuel Kant and Georges Cuvier (see Cheung 2000; 2010; Kirchhoff 2002; Weil 2005; Kirchhoff 2007; Toepfer 2011). For the influence of Kant’s mutualistic organicism on biologists (see, e.g., Gilbert and Sarkar 2000: 3).
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Acknowledgements
I dedicate this paper to the memory of Ludwig Trepl, in gratitude for his dedicated teaching and for the thorough and joyful discussions of topics like that of this paper which he made possible in his time as Chair of Landscape Ecology at the Technical University of Munich. I would like to thank the anonymous reviewers and the guest editors for their detailed comments that helped me to significantly improve the manuscript.
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Kirchhoff, T. The myth of Frederic Clements’s mutualistic organicism, or: on the necessity to distinguish different concepts of organicism. HPLS 42, 24 (2020). https://doi.org/10.1007/s40656-020-00317-y
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DOI: https://doi.org/10.1007/s40656-020-00317-y
Keywords
- Theory and history of ecology
- Different concepts of organicism
- Mutualism
- Competition
- Control hierarchy
- Frederic E. Clements