Are human beings part of the rest of nature?

Unified explanations seek to situate the traits of human beings in a causal framework that also explains the trait values found in nonhuman species. Disunified explanations claim that the traits of human beings are due to causal processes not at work in the rest of nature. This paper outlines a methodology for testing hypotheses of these two types. Implications are drawn concerning evolutionary psychology, adaptationism, and anti-adaptationism. This revised version was published online in July 2006 with corrections to the Cover Date.     

A Philosophical Evaluation of Adaptationism as a Heuristic Strategy

Adaptationism has prompted many a debate in philosophy of biology but the focus is usually on empirical and explanatory issues rather than methodological adaptationism (MA). Likewise, the context of evolutionary biology has provided the grounding for most discussions of the heuristic role of adaptationism. This paper extends the debate by drawing on case studies from physiology and systems biology to discuss the productive and problematic aspects of adaptationism in functional as well as evolutionary studies at different levels of biological organization. Gould and Lewontin’s Spandrels-paper famously criticized adaptationist methodology for implying a risk of generating ‘blind spots’ with respect to non-selective effects on evolution. Some have claimed that this bias can be accommodated through the testing of evolutionary hypotheses. Although this is an important aspect of overcoming the pitfalls of adaptationism, I argue that the issue of methodological biases is broader than the question of testability. I demonstrate the productivity of adaptationist heuristics but also discuss the deeper problematic aspects associated with the imperialistic tendencies of the strong account of MA.     

Mechanical explanation of nature and its limits in Kant's Critique of judgment

In this paper I discuss two questions. What does Kant understand by mechanical explanation in the Critique of judgment? And why does he think that mechanical explanation is the only type of the explanation of nature available to us? According to the interpretation proposed, mechanical explanations in the Critique of judgment refer to a particular species of empirical causal laws. Mechanical laws aim to explain nature by reference to the causal interaction between the forces of the parts of matter and the way in which they form into complex material wholes. Just like any other empirical causal law, however, mechanical laws can never be known with full certainty. The conception according to which we can explain all of nature by means of mechanical laws, it turns out, is based on what Kant calls 'regulative' or 'reflective' considerations about nature. Nothing in Kant's Critique of judgment suggests that these considerations can ever be justified by reference to how the natural world really is. I suggest that what, upon first consideration, appears to be a thoroughly mechanistic conception of nature in Kant is much more limited than one might have expected.     

Explanatory unification and natural selection explanations

The debate between the dynamical and the statistical interpretations of natural selection is centred on the question of whether all explanations that employ the concepts of natural selection and drift are reducible to causal explanations. The proponents of the statistical interpretation answer negatively, but insist on the fact that selection/drift arguments are explanatory. However, they remain unclear on where the explanatory power comes from. The proponents of the dynamical interpretation answer positively and try to reduce selection/drift arguments to some of the most prominent accounts of causal explanation. In turn, they face the criticism raised by statisticalists that current accounts of causation have to be violated in some of their core conditions or otherwise used in a very loose manner in order to account for selection/drift explanations. We propose a reconciliation of both interpretations by conveying evolutionary explanations within the unificationist model of scientific explanation. Therefore, we argue that the explanatory power in natural selection arguments is a result of successful unification of individual- and population-level facts. A short case study based on research on sympatric speciation will be presented as an example of how population- and individual-level facts are unified to explain the morphological mosaic of bill shape in island scrub jays (Aphelocoma insularis).     

How-possibly explanations as genuine explanations and helpful heuristics: A comment on Forber

Recently, Forber introduced a distinction between two kinds of how-possibly explanation, global and local how-possibly explanation, and argued that both play genuinely explanatory roles in evolutionary biology. In this paper I examine the nature of these two kinds of how-possibly explanations, focusing on the question whether they indeed constitute genuine explanations. I will conclude that one of Forber's kinds of how-possibly explanation may be thought of as a kind of genuine explanation but not as a kind of how-possibly explanation, while the other kind plays a heuristic role and should not be conceived of as a kind of explanation at all.     

Mechanical explanation of nature and its limits in Kant’s Critique of judgment

In this paper I discuss two questions. What does Kant understand by mechanical explanation in the Critique of judgment? And why does he think that mechanical explanation is the only type of the explanation of nature available to us? According to the interpretation proposed, mechanical explanations in the Critique of judgment refer to a particular species of empirical causal laws. Mechanical laws aim to explain nature by reference to the causal interaction between the forces of the parts of matter and the way in which they form into complex material wholes. Just like any other empirical causal law, however, mechanical laws can never be known with full certainty. The conception according to which we can explain all of nature by means of mechanical laws, it turns out, is based on what Kant calls ‘regulative’ or ‘reflective’ considerations about nature. Nothing in Kant’s Critique of judgment suggests that these considerations can ever be justified by reference to how the natural world really is. I suggest that what, upon first consideration, appears to be a thoroughly mechanistic conception of nature in Kant is much more limited than one might have expected.     

The evolution of virulence in vector-borne and directly transmitted parasites

Ewald (1994) has suggested that vector-borne parasites are expected to evolve a higher level of host exploitation than directly transmitted parasites, and this should thereby result in them being more virulent. Indeed, some data do conform to this general pattern. Nevertheless, his hypothesis has generated some debate about the extent to which it is valid. I explore this issue quantitatively within the framework of mathematical epidemiology. In particular, I present a dynamic optimization model for the evolution of parasite replication strategies that explicitly explores the validity of this hypothesis. A few different model assumptions are explored and it is found that Ewald's hypothesis has only qualified support as a general explanation for why vector-borne parasites are more virulent than those that are directly transmitted. I conclude by suggesting that an alternative explanation might lie in differences in inoculum size between these two types of transmission.     

Conjecture and explanation: A reply to Reydon

Reydon (2012) comments on my account of how-possibly explanation (Forber, 2010). I distinguish between three types of explanation (global how-possibly, local how-possibly, and how actually) and argue that these distinctions track various roles explanations play in evolutionary biology. While Reydon accepts the distinctions, he questions whether the two different types of how-possibly explanation count as genuine explanations. He summarizes his analysis with a slogan: “global how-possibly explanations are explanations but not how-possibly; local explanations are how-possibly but not explanations.” Reydon’s commentary raises a number of insightful points, and I will not be able to address them all. Instead, after clarifying certain points in my original paper (4 1), I will respond to Reydon’s slogan by addressing whether global how-possibly explanations should count as explaining how possible (4 2), and what (so-called) local how-possibly explanations are, if not explanations (4 3).     

Biological Levers and Extended Adaptationism

Two critiques of simple adaptationism are distinguished: anti-adaptationism and extended adaptationism. Adaptationists and anti-adaptationists share the presumption that an evolutionary explanation should identify the dominant simple cause of the evolutionary outcome to be explained. A consideration of extended-adaptationist models such as coevolution, niche construction and extended phenotypes reveals the inappropriateness of this presumption in explaining the evolution of certain important kinds of features—those that play particular roles in the regulation of organic processes, especially behavior. These biological or behavioral ‘levers’ are distinctively available for adaptation and exaptation by their possessors and for co-optation by other organisms. As a result they are likely to result from a distinctive and complex type of evolutionary process that conforms neither to simple adaptationist nor to anti-adaptationist styles of explanation. Many of the human features whose evolutionary explanation is most controversial belong to this category, including the female orgasm.

Causal and Mechanistic Explanations in Ecology

How are scientific explanations possible in ecology, given that there do not appear to be many-if any-ecological laws? To answer this question, I present and defend an account of scientific causal explanation in which ecological generalizations are explanatory if they are invariant rather than lawlike. An invariant generalization continues to hold or be valid under a special change-called an intervention-that changes the value of its variables. According to this account, causes are difference-makers that can be intervened upon to manipulate or control their effects. I apply the account to ecological generalizations to show that invariance under interventions as a criterion of explanatory relevance provides interesting interpretations for the explanatory status of many ecological generalizations. Thus, I argue that there could be causal explanations in ecology by generalizations that are not, in a strict sense, laws. I also address the issue of mechanistic explanations in ecology by arguing that invariance and modularity constitute such explanations.     

Rationality Wars and the War on Terror: Explaining Terrorism and Social Unrest

Terrorism is problematic at multiple levels. Social scientists debate its cause; policymakers debate what to do about it; many debate the meaning and political use of the term; and many live in fear of it. Current explanations of terrorism hinge on competing models of decision making. Anthropologists are increasingly influential in decision theory as issues of rationality, culture, and evolutionary psychology are invoked to explain patterns in human decision making. In this article, I review and critique current explanations of terrorism, I relate these explanations to larger debates in decision theory and anthropology, and I present an example of how current schisms may be transcended.     

The phylogeny fallacy and the ontogeny fallacy

In 1990 Robert Lickliter and Thomas Berry identified the phylogeny fallacy, an empirically untenable dichotomy between proximate and evolutionary causation, which locates proximate causes in the decoding of ‘genetic programs’, and evolutionary causes in the historical events that shaped these programs. More recently, Lickliter and Hunter Honeycutt (Psychol Bull 129:819–835, 2003a) argued that Evolutionary Psychologists commit this fallacy, and they proposed an alternative research program for evolutionary psychology. For these authors the phylogeny fallacy is the proximate/evolutionary distinction itself, which they argue constitutes a misunderstanding of development, and its role in the evolutionary process. In this article I argue that the phylogeny fallacy should be relocated to an error of reasoning that this causal framework sustains: the conflation of proximate and evolutionary explanation. Having identified this empirically neutral form of the phylogeny fallacy, I identify its mirror image, the ontogeny fallacy. Through the lens of these fallacies I attempt to solve several outstanding problems in the debate that ensued from Lickliter and Honeycutt’s provocative article.     

The Structure of Causal Explanations in Population Biology

The scope of this paper can be clarified by means of a well-known phenomenon that is usually called ‘industrial melanism’: the fact that the melanic form of the peppered moth became dominant in industrial areas in England in the second half of the nineteenth century. Such changes in relative phenotype frequencies are important explananda for population biologists. Apart from trying to explain such changes over time, population biologists also often try to explain differences between populations, e.g. why yellow shell colour is dominant in certain colonies of land snails and almost absent in other colonies. The causal explanations that are given to address such explananda are the objects of analysis in this paper. Our primary aim is to explicate their structure: we want to capture the typical ingredients of causal explanations in population biology, and their organisation. Based on this explication, we discuss how natural selection fits into recent mechanical philosophy of science, and engage in the debate on the nature of evolutionary theory.

     

The Mesoamerican Urban Tradition

The process of preindustrial urbanization is an important archeological issue because of its association with the emergence of early complex societies. There is considerable debate among Mesoamerican archeologists concerning both the evolutionary nature of the process itself and the configurations and functions of the centers that might be called "urban." Part of this debate is caused by the variation that clearly exists among such Mesoamerican centers, as well as the distinctive nature of urbanization in the culture area, which differs in important respects from similar processes in the Old World. Components of a model proposed by Richard Fox are used to set the Mesoamerican urban tradition into a wider, multilineal evolutionary framework. Copán, Teoti-huacan, Tenochtitlan, and other Mesoamerican centers provide comparative examples.     

Which humans behave adaptively,and why does it matter?

There has long been debate about the relevance of evolutionary theory to the study of humans. To many of us, however, the debate has shifted from whether to proceed with an evolutionary approach to how to proceed. Increasingly, it has been argued that studies of the current reproductive function of human traits make little or no contribution to the understanding of the psyche (e.g., Symons 1989). Here, on the basis of arguments about the relationship between an adaptation and an adaptive outcome, and a review of studies that assess current adaptiveness, I argue to the contrary that knowledge of the contexts in which people do or do not behave adaptively provides important information about the nature of the mechanisms that comprise the human psyche. In particular, studies that indicate that people behave adaptively in at least some contemporary environments cast doubt on many nonevolutionary constructions of human nature, and can be used now to distinguish alternative evolutionary constructions that are at odds over many issues pentaining to the human psyche's ontogeny and evolutionary background, especially the extent to which the human psyche is general purpose.     

Habitat suitability and the constraints of migration in New World warblers

Identifying the factors that influence geographic range limits can illustrate the various ecological, physiological, and evolutionary constraints imposed on a species. The range limits of migratory birds are particularly challenging to study as they occur in disjunct regions at different times of the year, which can impose different constraints. Travel between breeding and wintering regions poses a significant energetic and navigational challenge to birds, although it is not clear how these movements influence breeding dispersal and range expansion. Here I ask whether the possible costs of migration limit the breeding ranges of wood warblers, a group of birds with an extensive history of ecological and evolutionary studies. I used occurrence records for multiple wood warbler species, breeding primarily in the boreal forest of North America, to generate environmental niche models. I tested for over‐prediction of habitat suitability into the western boreal forest, where most these species do not have occurrence records but where there is presumably suitable habitat. I found that some of these vagile taxa, primarily found east of the Rocky Mountains, also have predicted habitat suitability that extends into the north and west, where they have little to no occurrence records. I discuss several possible explanations for this discordance. In particular, the patterns are consistent with the costs of a long‐distance migration limiting the benefits of range expansion, as migration may become too onerous as the distance between breeding and wintering areas increases. These results speak to the process of niche filling more generally and call attention to an under‐appreciated explanation for why migratory species may not fully occupy their fundamental niche.     

Optimality modeling in a suboptimal world

The fate of optimality modeling is typically linked to that of adaptationism: the two are thought to stand or fall together (Gould and Lewontin, Proc Relig Soc Lond 205:581–598, 1979; Orzack and Sober, Am Nat 143(3):361–380, 1994). I argue here that this is mistaken. The debate over adaptationism has tended to focus on one particular use of optimality models, which I refer to here as their strong use. The strong use of an optimality model involves the claim that selection is the only important influence on the evolutionary outcome in question and is thus linked to adaptationism. However, biologists seldom intend this strong use of optimality models. One common alternative that I term the weak use simply involves the claim that an optimality model accurately represents the role of selection in bringing about the outcome. This and other weaker uses of optimality models insulate the optimality approach from criticisms of adaptationism, and they account for the prominence of optimality modeling (broadly construed) in population biology. The centrality of these uses of optimality models ensures a continuing role for the optimality approach, regardless of the fate of adaptationism.

Searching for constraints by cross‐species comparison: reaction norms for age and size at maturity in insects

An evolutionary explanation should consider the balance between environmentally‐based selective pressures, and the resistance of the organism's phenotype to adaptive evolution, with the latter being captured by the concept of constraint. The limited attention to non‐adaptive explanations in evolutionary ecology is at least partly caused by methodological difficulties with respect to identifying and quantifying constraints. As an example of an experimental approach evaluating a constraint‐based explanation, we present a cross‐species comparison of the shape of reaction norms for size and age at maturity. Instar‐ and sex‐specific development times and final sizes were recorded for two distantly‐related species of insects (Lepidoptera), with larval growth rates being manipulated by means of refined starvation treatments. We found that (1) the ‘classical’ L‐shaped reaction norms for final size and development time are characteristic also of individual larval instars; (2) these responses show a high degree of quantitative similarity across the species, different larval instars, and sexes within species; and (3) the similarity among species and sexes is higher for the penultimate than for the final instar. The high degree of similarity suggests that some physiological mechanisms determining such reaction norms are evolutionarily conservative. An alternative explanation (i.e. quantitative similarity of ecologically based selective pressures) appears less likely. The results of a previous study on a third lepidopteran species not only support our general conclusions, but also provide a clear case of adaptive evolution in some aspects of such reaction norms. The present study shows one way how the data required to measure evolutionary conservatism in reaction norms for body size can be obtained empirically. © 2014 The Linnean Society of London, Biological Journal of the Linnean Society, 2015, 114 , 296–307.     

Evolutionary medicine at twenty: rethinking adaptationism and disease

Two decades ago, the eminent evolutionary biologist George C. Williams and his physician coauthor, Randolph Nesse, formulated the evolutionary medicine research program. Williams and Nesse explicitly made adaptationism a core component of the new program, which has served to undermine the program ever since, distorting its practitioners’ perceptions of evidentiary burdens and in extreme cases has served to warp practitioner’s understandings of the relationship between evolutionary benefits/detriments and medical ones. I show that the Williams and Nesse program more particularly embraces the panselectionist variety of adaptationism (the empirical assumption that non-adaptive evolutionary processes are causally unimportant compared to natural selection), and argue that this has harmed the field. Panselectionism serves to conceal the enormous evidentiary hurdles that evolutionary medicine hypotheses face, making them appear stronger than they are. I use two examples of evolutionary medicine texts, on neonatal jaundice and on asthma, to show that some evolutionary medicine practitioners have allowed their fervent panselectionism to directly shape their recommendations for clinical practice. I argue that this escalation of panselectionism’s influence is inappropriate under Williams’ and Nesse original stated standards, despite being inspired by their program. I also show that the examples’ conflation of clinical and evolutionary considerations is inappropriate even under Christopher Boorse’s controversial evolution-rooted concepts of disease and health.     

Viability explanation

This article deals with a type of functional explanation, viability explanation, that has been overlooked in recent philosophy of science. Viability explanations relate traits of organisms and their environments in terms of what an individual needs to survive and reproduce. I show that viability explanations are neither causal nor historical and that, therefore, they should be accounted for as a distinct type of explanation.The investigations for this paper were supported by the Foundation for Philosophical Research (SWON), which is subsidized by the Netherlands Organization for Scientific Research (NWO). I am grateful to Theo Kuipers and Ton Derksen for their useful comments on early versions, to Josje Lodder for improving my logic, and to Jaap van Brakel for his help in structuring this article.