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)

Much philosophical progress has been made in elucidating the idea of evolutionary contingency in a recent re-burgeoning of the debate. However, additional progress has been impaired on three fronts. The first relates to its characterisation: the under-specification of various contingency claims has made it difficult to conceptually pinpoint the scope to which ‘contingency’ allegedly extends, as well as which biological forms are in contention. That is—there appears to be no systematic means with which to fully specify contingency claims which has (...) led to a tendency for authors to talk past each other. Secondly, on the matter of evidence, recent research has focused on the evidential import of convergent evolution which is taken to disconfirm the evolutionary contingency thesis. However, there has been a neglect of convergent evolution’s converse: ‘evolutionary idiosyncrasies’ or the singular evolution of certain forms, which I argue is evidentially supportive of evolutionary contingency. Thirdly, evolutionary contingency has often been claimed to vary in degrees and that the debate, itself, is a matter of ‘relative significance’. However, there has been no formal method of evaluating the strength of contingency and its relative significance in a particular domain. In this paper, I address all three issues by proposing a systematic means of fully specifying contingency theses with the concept of the modal range. Secondly, I propose an account of evolutionary idiosyncrasies, investigate the explanations for their occurrences, and, subsequently, spell out their significance with respect to the evolutionary contingency thesis. Finally, having been equipped with the evidential counterpart to convergent evolution, I shall sketch a likelihood framework for evaluating, precisely on the basis of a sequence of opposing data, the strength and relative significance of evolutionary contingency in a particular domain. With this in hand, the relative observations of idiosyncrasies and convergences can be informative of the strength and relative significance of contingency in any particular domain. (shrink)

A number of authors have pointed to “convergent evolution” as evidence for the central role of natural selection in shaping predictable trajectories of macroevolution. However, there are numerous conceptual and empirical difficulties that arise in broadly appealing to the frequency of homoplasy as evidence for a non-contingently constrained adaptational design space. Most important is the need to distinguish between convergent (externally constrained) and parallel (internally constrained) evolution, and to consider how the respective frequencies of these significantly different sources of homoplasy (...) affect a strong adaptationist view of life. In this paper, I critically evaluate Simon Conway Morris’s use of the homoplasy literature to support his argument for a non-contingent, counterfactually stable account of macroevolutionary pattern. In so doing, I offer a conception of parallelism which avoids the charge that it differs from convergence merely in degree and not in kind. I argue that although organisms sharing a homoplastic trait will also share varying degrees of homology, it is the underlying developmental homology with respect to the generators directly causally responsible for the homoplastic event that defines parallel evolution and non-arbitrarily distinguishes it from convergence. The notion of “screening-off” is used to distinguish the proximal generators of a homoplastic trait from its more distal genetic causes (such as a master control gene). (shrink)

The comparative method grants epistemic access to the biological past. Comparing lineages provides empirical traction on both hypotheses about particular lineages and models of trait evolution. Understanding this evidential role is important. Although philosophers have recently turned their attention to relations of descent, little work exists exploring the status of evidence from convergences. I argue that, where they exist, convergences play a central role in the confirmation of adaptive hypotheses. I focus on ‘analogous inferences’, show how such inferences ought to (...) be analysed and suggest three methods for strengthening their evidential weight. 1 Introduction2 Analogous Inferences2.1 Adaptive explanations and analogies2.2 Analogous inferences2.3 Scope, grain, and specificity3 Parallel Modelling, Integrated Explanations, and Convergent Modelling3.1 Parallel modelling3.2 Integrated explanations3.1 Convergent modelling4 Conclusion. (shrink)

Abstract Starting with Ben-Menahem's definition of historical contingency as sensitivity to variations in initial conditions, we suggest that historical events and processes can be thought of as forming a complex landscape of contingency and necessity. We suggest three different ways of extending and elaborating Ben-Menahem's concepts: (1) By supplementing them with a notion of historical disturbance; (2) by pointing out that contingency and necessity are subject to scaling effects; (3) by showing how degrees of contingency/necessity can change over time. We (...) also argue that further development of Sterelny's notion of conditional inevitability leads to our conclusion that the topography of historical contingency is something that can change over time. (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)

George McGhee’s book “Convergent Evolution: limited forms most beautiful” provides an extensive survey of biological convergence. This paper has two main aims. First, it examines the theoretical claims McGhee makes about convergent evolution—specifically criticizing his use of a total morphospace to understand contingency and his assumption that functional constraints are non-contingent. Second, it sketches a group of important conceptual challenges facing researchers interested in convergence.