Biological Theory

, Volume 9, Issue 4, pp 382–391 | Cite as

Ecosystem Evolution is About Variation and Persistence, not Populations and Reproduction

  • Frédéric Bouchard
Thematic Issue Article: Ecosystems or Organisms?


Building upon a non-standard understanding of evolutionary process focusing on variation and persistence, I will argue that communities and ecosystems can evolve by natural selection as emergent individuals. Evolutionary biology has relied ever increasingly on the modeling of population dynamics. Most have taken for granted that we all agree on what is a population. Recent work has reexamined this perceived consensus. I will argue that there are good reasons to restrict the term “population” to collections of monophyletically related replicators and interactors, which explains why many existing models in population biology exclude by definition many genuine evolving biological individuals such as communities and ecosystems. By studying a case of community evolution (a symbiotic termite–fungus community), we will see that it is variation that is important to evolutionary processes, not populations. Variation within a population is only one of many types of variation that can lead to evolution by natural selection. The upshot of focusing on variation is that cases of community and ecosystem adaptive change become tractable in evolutionary terms. I will show that complex emergent individuals such as communities and ecosystems cannot be fully accommodated by conventional population/reproduction models but can be accommodated by variation/persistence models.


Biological individuality Community Ecosystem Evolution Fitness Persistence Philosophy of biology 


  1. Bapteste E, Lopez P, Bouchard F et al (2012) Evolutionary analyses of non-genealogical bonds produced by introgressive descent. Proc Natl Acad Sci USA 109:18266–18272CrossRefGoogle Scholar
  2. Bouchard F (2009) Understanding colonial traits using symbiosis research and ecosystem ecology. Biol Theory 4:240–246CrossRefGoogle Scholar
  3. Bouchard F (2010) Symbiosis, lateral function transfer and the (many) saplings of life. Biol Philos 25:623–641CrossRefGoogle Scholar
  4. Bouchard F (2011) Darwinism without populations: a more inclusive understanding of the ‘survival of the fittest.’ Stud Hist Philos Biol Biomed Sci 42:106–114Google Scholar
  5. Bouchard F (2013a) How ecosystem evolution strengthens the case for functional pluralism. In: Huneman P (ed) Functions: selection and mechanism. Springer, Dordrecht, pp 83–95Google Scholar
  6. Bouchard F (2013b) What is a symbiotic superindividual and how do you measure its fitness. In: Bouchard F, Huneman P (eds) From groups to individuals. MIT Press, Cambridge, pp 243–264Google Scholar
  7. Bouchard F, Huneman P (eds) (2013) From groups to individuals: evolution and emerging individuality. MIT Press, CambridgeGoogle Scholar
  8. Bouchard F, Rosenberg A (2004) Fitness, probability and the principles of natural selection. Brit J Philos Sci 55:693–712CrossRefGoogle Scholar
  9. Bronstein JL (1994) Our current understanding of mutualism. Quart Rev Biol 69:31–51CrossRefGoogle Scholar
  10. Buss LW (1987) The evolution of individuality. Princeton University Press, PrincetonGoogle Scholar
  11. Clarke E (2012) Plant individuality: a solution to the demographer’s dilemma. Biol Philos 27:321–361CrossRefGoogle Scholar
  12. Cropp R, Gabric A (2002) Ecosystem adaptation: do ecosystems maximize resilience? Ecology 83:2019–2026CrossRefGoogle Scholar
  13. Dawkins R (1976) The selfish gene. Oxford University Press, New YorkGoogle Scholar
  14. De Sousa R (2005) Biological individuality. Croatian J Philos 5:1–24Google Scholar
  15. Doolittle WF, Zhaxybayeva O (2010) Metagenomics and the units of biological organization. Bioscience 60:102–112CrossRefGoogle Scholar
  16. Dunbar MJ (1960) The evolution of stability in marine environments: natural selection at the level of the ecosystem. Am Nat 94:129–136CrossRefGoogle Scholar
  17. Dupré J (1993) The disorder of things: metaphysical foundations of the disunity of science, New edn. Harvard University Press, CambridgeGoogle Scholar
  18. Dupré J, O’Malley MA (2009) Varieties of living things: life at the intersection of lineage and metabolism. Philosophy & Theory in Biology 1 (December). doi: 10.3998/ptb.6959004.0001.003
  19. Ereshefsky M (ed) (1991) The units of evolution: essays on the nature of species. MIT Press, CambridgeGoogle Scholar
  20. Forber P (2005) On the explanatory roles of natural selection. Biol Philos 20:329–342CrossRefGoogle Scholar
  21. Ghiselin MT (1974) A radical solution to the species problem. Syst Zool 23:536–544CrossRefGoogle Scholar
  22. Ghiselin MT (1997) Metaphysics and the origin of species. State University of New York Press, AlbanyGoogle Scholar
  23. Godfrey-Smith P (2009) Darwinian populations and natural selection. Oxford University Press, New YorkGoogle Scholar
  24. Goodnight CJ (2005) Multilevel selection: the evolution of cooperation in non-kin groups. Popul Ecol 47:3–12CrossRefGoogle Scholar
  25. Hamilton A, Smith NR, Haber MH (2009) Social insects and the individuality thesis: cohesion and the colony as a selectable individual. In: Gadau J, Fewell J (eds) Organization of insect societies: from genome to sociocomplexity. Harvard University Press, Cambridge, pp 572–589Google Scholar
  26. Hoffman A (1979) Community paleoecology as an epiphenomenal science. Paleobiology 5:357–379Google Scholar
  27. Howe HF (1984) Constraints on the evolution of mutualisms. Am Nat 123:764–777CrossRefGoogle Scholar
  28. Hull DL (1976) Are species really individuals? Syst Zool 25:174–191CrossRefGoogle Scholar
  29. Hull DL (1978) A matter of individuality. Philos Sci 45:335–360CrossRefGoogle Scholar
  30. Hull DL (1980) Individuality and selection. Annu Rev Ecol Syst 11:311–332CrossRefGoogle Scholar
  31. Hull DL (2001) Science and selection: essays on biological evolution and the philosophy of science. Cambridge University Press, New YorkGoogle Scholar
  32. Levins R, Lewontin RC (1985) The dialectical biologist. Harvard University Press, CambridgeGoogle Scholar
  33. Lewin RA (1982) Symbiosis and parasitism: definitions and evaluations. Bioscience 32:254–260CrossRefGoogle Scholar
  34. Matthen M, Ariew A (2002) Two ways of thinking about fitness and natural selection. J Philos 99:55–83CrossRefGoogle Scholar
  35. Mayr E (1942) Systematics and the origin of species from the viewpoint of a zoologist. Columbia University Press, New YorkGoogle Scholar
  36. Millstein RL (2006) Natural selection as a population-level causal process. Brit J Phil Sci 57:627–653CrossRefGoogle Scholar
  37. Millstein RL (2009) Populations as individuals. Biol Theory 4:267–273CrossRefGoogle Scholar
  38. Millstein RL, Skipper RA Jr, Dietrich MR (2009) (Mis)interpreting mathematical models: drift as a physical process. Philosophy & Theory in Biology. doi: 10.3998/ptb.6959004.0001.002 Google Scholar
  39. Moran NA (2006) Symbiosis. Curr Biol 16:866–871CrossRefGoogle Scholar
  40. O’Malley MA, Dupré J (2007a) Towards a philosophy of microbiology. Stud Hist Philos Biol Biomed Sci 38:775–779CrossRefGoogle Scholar
  41. O’Malley MA, Dupré J (2007b) Size doesn’t matter: towards a more inclusive philosophy of biology. Biol Philos 22:155–191CrossRefGoogle Scholar
  42. Odenbaugh J (2007) Seeing the forest and the trees: realism about communities and ecosystems. Philos Sci 74:628–641CrossRefGoogle Scholar
  43. Okasha S (2006) Evolution and the levels of selection. Oxford University Press, OxfordCrossRefGoogle Scholar
  44. Ott JA (1981) Adaptive strategies at the ecosystem level: examples from two Benthic marine systems. Mar Ecol 2:113–158CrossRefGoogle Scholar
  45. Paracer S, Ahmadjian V (2000) Symbiosis: an introduction to biological associations, 2nd edn. Oxford University Press, New YorkGoogle Scholar
  46. Pradeu T (2012) The limits of the self: immunology and biological identity. Oxford University Press, OxfordGoogle Scholar
  47. Saffo MB (1988) Symbiosis. BioScience 38:710–711CrossRefGoogle Scholar
  48. Saffo MB (2002) Themes from variation: probing the commonalities of symbiotic associations. Integr Comp Biol 42:291–294CrossRefGoogle Scholar
  49. Sapp J (1994) Evolution by association: a history of symbiosis. Oxford University Press, New YorkGoogle Scholar
  50. Swenson W, Arendt A, Wilson DS (2000a) Artificial selection of microbial ecosystems for 3-chloroaniline biodegradation. Environ Microbiol 2:564–571CrossRefGoogle Scholar
  51. Swenson W, Wilson DS, Elias R (2000b) Artificial ecosystem selection. Proc Natl Acad Sci 97:9110–9114CrossRefGoogle Scholar
  52. Thoday JM (1953) Components of fitness. Symp Soc Exp Biol 7:96–113Google Scholar
  53. Turnbaugh PJ, Hamady M, Yatsunenko T et al (2008) A core gut microbiome in obese and lean twins. Nature 457:480–484CrossRefGoogle Scholar
  54. Turner JS (2000) The extended organism: the physiology of animal-built structures. Harvard University Press, CambridgeGoogle Scholar
  55. Turner JS (2004) Extended phenotypes and extended organisms. Biol Philos 19:327–352CrossRefGoogle Scholar
  56. Turner JS (2013) Superorganisms and superindividuality: the emergence of individuality in a social insect assemblage. In: Bouchard F, Huneman P (eds) From groups to individuals. MIT Press, Cambridge, pp 219–241Google Scholar
  57. Van Valen LM (1989) Three paradigms of evolution. Evol Theory 9:1–17Google Scholar
  58. Van Valen LM (1991) Biotal evolution: a manifesto. Evol Theory 10:1–13Google Scholar
  59. Walsh DM, Ariew A, Lewens T (2002) The trials of life: natural selection and random drift. Philos Sci 69:452–473CrossRefGoogle Scholar
  60. Wilson RA (2004) Boundaries of the mind: the individual in the fragile sciences. Cambridge University Press, CambridgeGoogle Scholar
  61. Wilson J (2007a) Biological individuality: the identity and persistence of living entities. Cambridge University Press, CambridgeGoogle Scholar
  62. Wilson RA (2007b) The biological notion of individuality. In: Zalta N (ed) The Stanford encyclopedia of philosophy.
  63. Wilson DS, Sober E (1989) Reviving the superorganism. J Theor Biol 136:337–356CrossRefGoogle Scholar

Copyright information

© Konrad Lorenz Institute for Evolution and Cognition Research 2014

Authors and Affiliations

  1. 1.Département de philosophie, Centre interuniversitaire de recherche sur la science et la technologieUniversité de MontréalMontrealCanada

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