This paper develops a critical response to John Beatty’s recent (2006) engagement with Stephen Jay Gould’s claim that evolutionary history is contingent. Beatty identifies two senses of contingency in Gould’s work: an unpredictability sense and a causal dependence sense. He denies that Gould associates contingency with stochastic phenomena, such as drift. In reply to Beatty, this paper develops two main claims. The first is an interpretive claim: Gould really thinks of contingency has having to do with stochastic effects at the (...) level of macroevolution, and in particular with unbiased species sorting. This notion of contingency as macro-level stochasticity incorporates both the causal dependence and the unpredictability senses of contingency. The second claim is more substantive: Recent attempts by other scientists to put Gould’s claim to the test fail to engage with the hypothesis that species sorting sometimes resembles a lottery. Gould’s claim that random sorting is a significant macroevolutionary phenomenon remains an intriguing and largely untested live hypothesis about evolution. (shrink)

One of the first questions that paleontologists ask when they identify a large-scale trend in the fossil record (e.g., size increase, complexity increase) is whether it is passive or driven. In this article, I explore two questions about driven trends: (1) what is the underlying cause or source of the directional bias? and (2) has the strength of the directional bias changed over time? I identify two underdetermination problems that prevent scientists from giving complete answers to these two questions.

The latter half of the twentieth century has been marked by debates in evolutionary biology over the relative significance of natural selection and random drift: the so-called “neutralist/selectionist” debates. Yet John Beatty has argued that it is difficult, if not impossible, to distinguish the concept of random drift from the concept of natural selection, a claim that has been accepted by many philosophers of biology. If this claim is correct, then the neutralist/selectionist debates seem at best futile, and at worst, (...) meaningless. I reexamine the issues that Beatty raises, and argue that random drift and natural selection, conceived as processes, can be distinguished from one another. (shrink)

Recent discussions in the philosophy of biology have brought into question some fundamental assumptions regarding evolutionary processes, natural selection in particular. Some authors argue that natural selection is nothing but a population-level, statistical consequence of lower-level events (Matthen and Ariew [2002]; Walsh et al. [2002]). On this view, natural selection itself does not involve forces. Other authors reject this purely statistical, population-level account for an individual-level, causal account of natural selection (Bouchard and Rosenberg [2004]). I argue that each of these (...) positions is right in one way, but wrong in another; natural selection indeed takes place at the level of populations, but it is a causal process nonetheless. (shrink)

Evolution in its contemporary meaning in biology typically refers to the changes in the proportions of biological types in a population over time (see the entry on the concept of evolution to 1872 for earlier meanings). As evolution is too large of a topic to address thoroughly in one entry, the primary goal of this entry is to serve as a broad overview of contemporary issues in evolution with links to other entries where more in-depth discussion can be found. The (...) entry begins with a brief survey of definitions of evolution, followed by a discussion of the different modes of evolution and related philosophical issues, and ends with a summary of other topics in the philosophy of evolution focusing particularly on topics covered in this encyclopedia. (shrink)

Behavioural flexibility is often treated as the gold standard of evidence for more sophisticated or complex forms of animal cognition, such as planning, metacognition and mindreading. However, the evidential link between behavioural flexibility and complex cognition has not been explicitly or systematically defended. Such a defence is particularly pressing because observed flexible behaviours can frequently be explained by putatively simpler cognitive mechanisms. This leaves complex cognition hypotheses open to ‘deflationary’ challenges that are accorded greater evidential weight precisely because they offer (...) putatively simpler explanations of equal explanatory power. This paper challenges the blanket preference for simpler explanations, and shows that once this preference is dispensed with, and the full spectrum of evidence—including evolutionary, ecological and phylogenetic data—is accorded its proper weight, an argument in support of the prevailing assumption that behavioural flexibility can serve as evidence for complex cognitive mechanisms may begin to take shape. An adaptive model of cognitive-behavioural evolution is proposed, according to which the existence of convergent trait–environment clusters in phylogenetically disparate lineages may serve as evidence for the same trait–environment clusters in other lineages. This, in turn, could permit inferences of cognitive complexity in cases of experimental underdetermination, thereby placing the common view that behavioural flexibility can serve as evidence for complex cognition on firmer grounds. (shrink)

The ecologist today finds scarce ground safe from controversy. Decisions must be made about what combination of data, goals, methods, and theories offers them the foundations and tools they need to construct and defend their research. When push comes to shove, ecologists often turn to philosophy to justify why it is their approach that is scientific. Karl Popper’s image of science as bold conjectures and heroic refutations is routinely enlisted to justify testing hypotheses over merely confirming them. One of the (...) most controversial theories in contemporary science is the Neutral Theory of Ecology. Its chief developer and proponent, Stephen Hubbell, presents the neutral theory as a bold conjecture that has so far escaped refutation. Critics of the neutral theory claim that it already stands refuted, despite what the dogmatic neutralists say. We see the controversy through a Popperian lens. But Popper’s is an impoverished philosophy of science that distorts contemporary ecology. -/- The controversy surrounding the neutral theory actually rests on a methodological fault. There is a strong but messy historical link between the concepts of being neutral and being null in biology, and Hubbell perpetuates this when he claims that the neutral theory is supplies the appropriate null for testing alternative theories. What method is being followed here? There are three contenders: -/- Null hypothesis testing tests for whether a there is a pattern to be explained. Null modeling tests for whether a process is causally relevant to a pattern. Baseline modeling apportions relative responsibility to multiple processes each relevant to a pattern. -/- Whether the neutral theory supplies an appropriate “null” depends upon whether null hypothesis, null modeling, or baseline model is intended. These methods prescribe distinct inference patterns. If they are null hypothesis testing or null modeling, the neutralists’s reasoning is invalid. If they are baseline modeling, the justification of a crucial assumption remains opaque. Either way, the neutral-null connection is being exploited rhetorically to privilege the neutral theory over its rivals. Clarifying the reasoning immunizes us against the rhetoric and foregrounds the underlying virtues of the neutralist approach to ecology. -/- The Popperian lens distorts theoretical development as dogmatism. Lakatos’s view of science as the development of research programmes clarifies the epistemology of the neutral theory. Focusing philosophical attention on the neutralist research programme illuminates (1) the synchronic uses of the neutral theory to make predictions and give descriptions and explanations; (2) its diachronic development in response to theoretical innovation and confrontation with data; (3) its complex relationships to alternative theories. For example, baseline modeling is now seen to be its primary explanatory heuristic. The justification for baseline modeling with the neutral theory, previously hidden from view, is seen in the logic of in the neutralist research programme. (shrink)

The distinction between idiographic science, which aims to reconstruct sequences of particular events, and nomothetic science, which aims to discover laws and regularities, is crucial for understanding the paleobiological revolution of the 1970s and 1980s. Stephen Jay Gould at times seemed conflicted about whether to say (a) that idiographic science is fine as it is or (b) that paleontology would have more credibility if it were more nomothetic. Ironically, one of the lasting results of the paleobiological revolution was a new (...) way of doing historical science that defies categorization as idiographic or nomothetic. Yet the tension between (a) and (b) persists in some recent work in the philosophy of science, with Carol Cleland defending a version of (a) and Dan McShea and Robert Brandon defending a version of (b). In placing Cleland's work into conversation with that of McShea and Brandon, this paper defends a third (c) synthetic approach that emphasizes the blending of idiographic and nomothetic work. (shrink)

Proceedings of the Pittsburgh Workshop in History and Philosophy of Biology, Center for Philosophy of Science, University of Pittsburgh, March 23-24 2001 Session 4: Evolutionary Indeterminism.

The four case studies on chance in evolution provide a rich source for further philosophical analysis. Among the issues raised are the following: Are there different conceptions of chance at work, or is there a common underlying conception? How can a given concept of chance be distinguished from other chance concepts and from nonchance concepts? How can the occurrence of a given chance process be distinguished empirically from nonchance processes or other chance processes? What role does chance play in evolutionary (...) theory? I argue that in order to answer these questions, a careful distinction between process and outcome must be made; however, the purpose of this essay is not to answer these questions definitively, but rather to elaborate on them and to provide a starting point for further discussion. (shrink)

‘Gouldian arguments’ appeal to the contingency of a scientific domain to establish that domain’s autonomy from some body of theory. For instance, pointing to evolutionary contingency, Stephen Jay Gould suggested that natural selection alone is insufficient to explain life on the macroevolutionary scale. In analysing contingency, philosophers have provided source-independent accounts, understanding how events and processes structure history without attending to the nature of those events and processes. But Gouldian Arguments require source-dependent notions of contingency. An account of contingency is (...) source-dependent when it is indexed to some pattern and some process. Positions like Gould’s do not turn on the mere fact of life’s contingency—that life’s shape could have been different due to its sensitivity to initial conditions, path-dependence or stochasticity. Rather, Gouldian arguments require that the contingency is due to particular kinds of processes: in this case, those which microevolutionary theory cannot account for. This source-dependent perspective clarifies both debates about the nature and importance of contingency, and empirical routes for testing Gould’s thesis. (shrink)

Historically, one of the most controversial aspects of Darwinian evolution has been the prominent role that randomness and random change play in it. Most biologists agree that mutations in DNA have random effects on fitness. However, fitness is a highly simplified scalar representation of an enormously complex phenotype. Challenges to Darwinian thinking have focused on such complex phenotypes. Whether mutations affect such complex phenotypes randomly is ill understood. Here I discuss three very different classes of well-studied molecular phenotypes in which (...) mutations cause nonrandom changes, based on our current knowledge. What is more, this nonrandomness facilitates evolutionary adaptation. Thus, living beings may translate DNA change into nonrandom phenotypic change that facilitates Darwinian evolution. (shrink)

Stephen Jay Gould argued that replaying the “tape of life” would result in a radically different evolutionary outcome. Some biologists and philosophers, however, have pointed to convergent evolution as evidence for robust replicability in macroevolution. These authors interpret homoplasy, or the independent origination of similar biological forms, as evidence for the power of natural selection to guide form toward certain morphological attractors, notwithstanding the diversionary tendencies of drift and the constraints of phylogenetic inertia. In this paper, I consider the implications (...) of homoplasy for the debate over the nature of macroevolution. I argue that once the concepts of contingency and convergence are fleshed out, it becomes clear that many instances of homoplasy fail to negate Gould’s overarching thesis, and may in fact support a Gouldian view of life. My argument rests on the distinction between parallelism and convergence, which I defend against a recent challenge from developmental biology. I conclude that despite the difficulties in defining and identifying parallelism, the concept remains useful and relevant to the contingency controversy insofar as it underscores the common developmental origins of iterated evolution. (shrink)

Darwinism designates a distinctive form of evolutionary explanation for the history and diversity of life on earth. Its original formulation is provided in the first edition of On the Origin of Species in 1859. This entry first formulates ‘Darwin's Darwinism’ in terms of five philosophically distinctive themes: (i) probability and chance, (ii) the nature, power and scope of selection, (iii) adaptation and teleology, (iv) nominalism vs. essentialism about species and (v) the tempo and mode of evolutionary change. Both Darwin and (...) his critics recognized that his approach to evolution was distinctive on each of these topics, and it remains true that, though Darwinism has developed in many ways unforeseen by Darwin, its proponents and critics continue to differentiate it from other approaches in evolutionary biology by focusing on these themes. This point is illustrated in the second half of the entry by looking at current debates in the philosophy of evolutionary biology on these five themes, with a special focus on Stephen Jay Gould's The Structure of Evolutionary Theory. (shrink)

The Gouldian argument for evolutionary contingency found in Wonderful Life can be dissected into three premises: palaeontological, macro-evolutionary, and developmental. Discussions of evolutionary contingency have revolved primarily around the developmental. However, a shift in methodological practice and new palaeontological evidence subsequent to the book’s publication appears to threaten the palaeontological premise that asserts high Cambrian disparity, or, roughly, that morphological differences between the Cambrian species were high. This presents a prima facie problem: Did the Cambrian consist of enough anatomical variety (...) to fuel a different outcome upon a replay of the tape of life? I investigate the role that disparity plays in the Gouldian argument for evolutionary contingency and conclude that the reduced disparity view does not undermine the argument, per se. As for the macro-evolutionary premise, Gould never offers any explicit justification for the idea that species sorting was indiscriminate or ‘lottery-like’ (though he certainly gestures at it). Putting aside the exegetical, I consider certain empirical avenues for underwriting the macro-evolutionary premise and argue that the most plausible route is to appeal to the special status of mass extinctions, a move that is not entirely unfamiliar to macro-evolutionists. That is, mass extinctions can be good sources of contingency due to their being ‘chancy’ in at least three respects. In sum, I update the palaeontological and macro-evolutionary premises as to reconstrue the Gouldian argument for evolutionary contingency in its strongest form. (shrink)

Work throughout the history and philosophy of biology frequently employs ‘chance’, ‘unpredictability’, ‘probability’, and many similar terms. One common way of understanding how these concepts were introduced in evolution focuses on two central issues: the first use of statistical methods in evolution (Galton), and the first use of the concept of “objective chance” in evolution (Wright). I argue that while this approach has merit, it fails to fully capture interesting philosophical reflections on the role of chance expounded by two of (...) Galton's students, Karl Pearson and W.F.R. Weldon. Considering a question more familiar from contemporary philosophy of biology—the relationship between our statistical theories of evolution and the processes in the world those theories describe—is, I claim, a more fruitful way to approach both these two historical actors and the broader development of chance in evolution. (shrink)

There is still a great deal of debate over what counts as a constraint and about how to assess experimentally the relative importance of constraints and selection in evolutionary history. I will argue that the notion of a constraint on variation, and thus the selection-constraint distinction, depends on two specifications: (1) what counts as a variant -- constraints limit or bias the production of what? and (2) range of assessment -- over what range of times or conditions is the variation (...) assessed? Specifications 1 and 2 help us to understand empirical work on the relative importance of constraint and selection in evolution. (shrink)