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Adaptationism and the Logic of Research Questions: How to Think Clearly About Evolutionary Causes Biol Theory (2015) 10:343–362 DOI 10.1007/s13752-015-0214-2 ORIGINAL ARTICLE Adaptationism and the Logic of Research Questions: How to Think Clearly About Evolutionary Causes Elisabeth A. Lloyd1 Received: 20 March 2015 / Accepted: 29 May 2015 / Published online: 9 July 2015 Ó Konrad Lorenz Institute for Evolution and Cognition Research 2015 Abstract This article discusses various dangers that popular methods, despite its benign reputation. In ‘‘The accompany the supposedly benign methods in behavioral Spandrels of San Marco and the Panglossian Paradigm,’’ evolutionary biology and evolutionary psychology that fall Gould and Lewontin (1979) drew attention to several under the framework of ‘‘methodological adaptationism.’’ dangers in using this method. In this article, I present a A ‘‘Logic of Research Questions’’ is proposed that aids in framework for analysis that makes their worries clearer. I clarifying the reasoning problems that arise due to the also warn of further risks of this methodological frame- framework under critique. The live, and widely practiced, work, expanding on the dangers it poses to scientific rea- ‘‘evolutionary factors’’ framework is offered as the key soning in evolutionary biology. At the same time, I comparison and alternative. The article goes beyond the emphasize that I am not attacking the notion of looking for traditional critique of Stephen Jay Gould and Richard C. adaptations in evolutionary studies: I am not anti-adapta- Lewontin, to present problems such as the disappearance of tion. The issues concern which framework is most appro- evidence, the mishandling of the null hypothesis, and priate and fruitful. failures in scientific reasoning, exemplified by a case from As evolutionary biology is usually taught and con- human behavioral ecology. In conclusion the paper shows ceived, there are a variety of evolutionary forces or types that ‘‘methodological adaptationism’’ does not deserve its of factors that can influence the form and distribution of a benign reputation. given trait in a population or species (Singh and Krimbas 2000;Futuyma2013). While natural selection may be the Keywords Adaptationism Á Behavioral biology Á most significant factor, we also have sexual selection, Behavioral ecology Á Evolutionary methods Á Evolutionary genetic linkage, phyletic history or ‘‘inertia,’’ develop- psychology Á Evolutionary theory Á Female orgasm mental factors, drift or chance, embryological constraints, and social, environmental, and niche coevolutionary fac- tors (Wright 1931; Odling-Smee et al. 2001; Pigliucci and Introduction Mu¨ller 2010). Traits can also be byproducts, spandrels, or exaptations of any of these processes in a co-related or We do not usually think that the logic of our scientific linked trait, among other causal and explanatory factors methods leads to closed-mindedness, and the inability to (Gould and Lewontin 1979; Gould and Vrba 1982; see alternatives, or evaluate evidence, but that is exactly Futuyma 2013).Letuscallthisbasicapproachthe‘‘evo- what sometimes happens in evolutionary biology of lutionary factors’’ framework of evolutionary theory; its behavioral and morphological traits with one of its most fundamental research question is: ‘‘What evolutionary factors account for the form and distribution of this trait?’’ Often, several of these factors are understood to & Elisabeth A. Lloyd operate simultaneously on a given trait, but only one or [email protected] two are the major factors causing its form and distribution 1 Department of History and Philosophy of Science, Indiana at a given time (e.g., Otsuka 2014; see Newman 1988; University, Bloomington, IN, USA Amundson 1994, 1998, 2005;Griffiths1996;Raff1996; 123 344 E. A. Lloyd Carroll 2005; Newman and Bhat 2008). When we inves- might be able to reach into ant nests with their long tigate the evolutionary origins of a given trait, we usually tongues and eat the most ants, and were thus most able to prioritize the functional factors, natural selection and pursue their food resources. These longer-tongued antea- sexual selection, as the most significant factors in evolu- ter-ancestors—eventually with their 25-inch-long ton- tionary research, and we might start with the question: gues—would represent the best fit—or closest-to-best ‘‘Does this trait have a function?’’1 If the trait, after fit—to their environment. The anteater example thus investigation, does not appear to have a correlation with presents a good instance of a natural selection explanation fitness,2 or does not appear to have evidence of design that reinforces or produces an adaptation. I take it as given (hence, does not appear to have a current or past function), that our living world is filled with examples of such we pursue other possible evolutionary explanations, such adaptations. as whether it might be due to genetic linkage with another Consider a breed of scientist called a ‘‘methodologi- trait, or be an exaptation, or a byproduct of selection (see cal’’ (or ‘‘heuristic’’) adaptationist, an evolutionary biol- Gould and Vrba 1982; Gould 2002; Lloyd and Gould ogist who assumes, at the beginning of investigation, that ([2002]2014). Alternatively, it may be present due to a trait is, indeed, an adaptation.3 Assuming adaptation is developmental or embryological constraints, or due to standard operating procedure among most behavioral phyletic inertia, and so on (Wake 1991, 2009;Newman ecologists, evolutionary psychologists, and human evo- and Bhat 2008, 2011; Linde-Medina 2011; Griesemer lutionists. Many of them cite Ernst Mayr’s (1983) 2015). Pursuit of such explanations would consist of defense of an ‘‘adaptationist research program,’’ written testing them against available evidence and searching for in reaction to Gould and Lewontin’s critical 1979 new evidence specific to those factors, against which they ‘‘Spandrels of San Marco.’’ Mayr sets up the problem so could then be compared. that selection is the only answer to the evolutionary There is another approach, dominant among leading question: animal behaviorists, behavioral ecologists, and many Consequently, when one attempts to explain the human evolutionists and evolutionary psychologists, features of something that is the product of evolution, called ‘‘methodological adaptationism.’’ Under this one must attempt to reconstruct the evolutionary approach, the leading research question is: ‘‘What is the history of this feature…. The most helpful procedure function of this trait?’’ or ‘‘What adaptive explanation in an analysis of historical narratives is to ask ‘‘why’’ can account for this trait?’’ And the research consists of questions; that is, questions (to translate this into an exploration and search for supportive evidence for modern evolutionary language) which ask what is or adaptive hypotheses that can explain the trait’s presence might have been the selective advantage that is in the population. responsible for the presence of a particular feature. (Mayr 1983, p. 325) The Logic of Research Questions: Alternatives Mayr continues on to advocate an adaptationist methodol- ogy for pursuing evolutionary explanations: Issues about method in behavioral ecology and biology When one attempts to determine for a given trait revolve around evolutionary adaptations, one of evolu- whether it is the result of natural selection or of tion’s biggest successes. Evolutionary adaptations are chance (the incidental byproduct of stochastic pro- traits that exist today because they were products of nat- cesses), one is faced by an epistemological dilemma. ural selection acting on a variety of developed phenotypes Almost any change in the course of evolution might in the past history of the species (Burian 1992;West- have resulted by chance. Can one ever prove this? Eberhard 1992;Griffiths1996). In the ancestral popula- Probably never. By contrast, can one deduce the tion of anteaters, for instance, which resembled armadil- probability of causation by selection? Yes, by los, tongue length was likely highly variable, with high showing that possession of the respective feature fitness values accruing to those anteater-ancestors that would be favored by selection. It is this consideration which determines the approach of the evolutionist. He must first attempt to explain biological phenom- 1 See Rose and Lauder (1996) for some examples of the application of this evolutionary factors approach. Or Martins (2000), for some ena and processes as the product of natural selection. methodological details. 2 Symons gives a variety of reasons against using correlation with 3 See Lewens (2009) for an extensive categorization of types of fitness for detecting adaptations, and prefers evidence of design adaptationism. Amundson (2001) and Sansom (2003) have also (1990). Thornhill (1990) provides another perspective of behavioral emphasized the multiple nature of adaptationist questions and adaptationists. answers, but not in the way I do here. 123 Adaptationism and the Logic of Research Questions 345 Only after all attempts to do so have failed, is he ‘‘function’’ in the selective sense. For example, a trait justified in designating the unexplained residue ten- could contribute to fitness in the current population, tatively as a product of chance. (Mayr 1983, p. 326)4 without having been formed by selection to have done so. Gould and Vrba (1982) dubbed such traits ‘‘exapta- Thus, he promotes the key question: ‘‘The adaptationist tions’’; these have current utility but not functions in the question, ‘What is the function of a given structure
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