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Mayr and Tinbergen: disentangling and integrating

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Abstract

Research on animal behavior is typically organized according to a combination of two influential frameworks: Ernst Mayr’s distinction between proximate and ultimate causes, and Niko Tinbergen’s “four questions” (mechanisms, development, survival value, and evolution). My aim is to debunk two common interpretive misconceptions about Mayr’s proximate–ultimate distinction and its relationship to Tinbergen’s four questions, and to offer a new interpretation that avoids both. The first misconception is that the proximate–ultimate distinction maps cleanly onto Tinbergen’s four questions, marking a boundary between Tinbergen’s evolutionary and survival value questions (ultimate) versus developmental and mechanistic questions (proximate). The second is that Mayr’s proximate–ultimate distinction is meant to rule out the relevance of proximate causes to evolutionary explanations. I argue that neither is plausible given the text and Mayr’s philosophical aims, namely, to argue that evolutionary biology cannot be reduced to either the physical sciences or to other areas of biology. Through a reconstruction of Mayr’s anti-reductionist argument, I develop an interpretation according to which the proximate–ultimate distinction marks two ways that teleological reasoning can be naturalistically grounded in biology, corresponding to Mayr’s distinction between teleonomic and adapted systems. Mayr distinguishes reduction, which the proximate–ultimate distinction is meant to block, from analysis, through which he allows that proximate causes, causes that are neither proximate nor ultimate, and chance can all contribute to evolutionary explanations. I conclude by suggesting some ways in which the interpretation defended here reframes our understanding of Mayr’s disagreements with some evolutionary-developmental biologists.

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Notes

  1. The earliest explicit statement of the Standard View I have found occurs in Sherman (1988) and Holekamp and Sherman (1989). As an anonymous referee has noted, the Standard View appears to be presumed in Wilson’s (in)famous “amoeba diagram” in his (1975) Sociobiology. It is worth noting the view met with backlash at the time. I suspect the dominance of this interpretation can largely be attributed to John Alcock’s (1993) inclusion of it, citing Sherman’s and Holekamp’s papers, in the first chapter of the fifth and subsequent editions of his influential Animal Behavior textbook. As of now, the book is in its eleventh edition (Rubenstein and Alcock 2018). The view is usually stated without argument because it is thought of as established textbook knowledge.

  2. Both Gardner (2013) and Scholl and Pigliucci (2015) offer important critiques of Laland et al. (2011). Calcott (2013a, b) and Scholl and Pigliucci (2015) characterize drift and other non-selective evolutionary processes as ultimate causes while Ariew (2003) and Gardner (2013) deny that they are.

  3. Interestingly, Mayr (1974, 1988) uses the term ‘functional analysis’, and Mayr (1992b) cites Cummins (1975), but it is not entirely clear from the context whether the citation is approving, disapproving, or neutral.

  4. However, we should not underestimate Mayr’s awareness of the philosophical literature. Both his citations and his correspondence reveal very active engagement with the philosophy of science community. I suspect that by the end of his life, he did have a well-thought-out account of analysis, but my goal here is not to uncover it.

  5. In the second of Mayr’s four examples at the beginning of “Overview of the two frameworks” section, Mayr (1961) seems to suggest that the possession of a genetic constitution favored by selection is an ultimate cause. He repeats this example, substantially reworded, in a later book (Mayr 1984), explicitly noting that the relevant evolutionary process is selection. However, he still appears to be saying that the genetic program that results from that process is an ultimate cause. Since the example is anomalous in this respect, and originates in his earliest work on the distinction, I am inclined to think it can be dismissed as Mayr being uncareful and conflating process with product.

  6. This claim and the earlier division between the role of narrative in delineating the humanities from the exact sciences echoes the views of neo-Kantians whose work Mayr would likely have encountered, especially Heinrich Rickert (see Staite (2013)). However, I have been unable to find explicit reference to the neo-Kantians in Mayr’s work. It is worth noting that Mayr often cites Kant himself approvingly (see especially Mayr (1982, 1988)).

  7. Mayr’s impression that his developmentalist critics were resurrecting a discredited kind of teleology was further encouraged by the tendency of those critics to label their views Lamarckian (e.g. Ho and Saunders 1984). Laurent Loison (2018) has recently, and I think convincingly, argued that Lamarckism, properly understood, indeed carries the supposition that there is an inherently end-directed vital force in living matter.

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Acknowledgements

Work on this paper was spread over time spent in the departments of philosophy at Cornell University, Virginia Commonwealth University, and Colgate University, and I would like to thank all three for their support. Special thanks go to Richard Boyd, Derk Pereboom, William Starr, Frances Fairbairn, Nicole Lee, Annaliese Beery, Maureen O’Malley and two anonymous reviewers, all of whom read drafts at various stages of development and provided helpful commentary. The paper also benefitted from useful discussions with Darragh Hare, Hudson Kern Reeve, and audiences in the Cornell Sage School of Philosophy, Cornell Neurobiology and Behavior Department, and the 2017 International Society for the History, Philosophy, and Social Studies of Biology. Finally, I wish to dedicate this paper to the memory of Ann Johnson, who provided encouragement and feedback on this and other work.

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Correspondence to Brandon A. Conley.

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Conley, B.A. Mayr and Tinbergen: disentangling and integrating. Biol Philos 35, 4 (2020). https://doi.org/10.1007/s10539-019-9731-x

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