Biological lineages move through time, space, and each other. As they do, they diversify, diverge, and grade away from and into one another. One result of this is genealogical discordance; i.e., the lineages of a biological entity may have different histories. We see this on numerous levels, from microbial networks, to holobionts, to population-level lineages. This paper considers how genealogical discordance impacts our study of species. More specifically, I consider this in the context of three framing questions: (1) How, if (...) at all, does genealogical discordance challenge, modify, or revise how we conceive of species? (2) How has growing appreciation of genealogical discordance impacted scientific practice? Of systematics in particular? (3) How do lineages at different levels diverge and diversify? All told, genealogical discordance enriches and complicates our taxonomic ontology, as well as the practice of reconstructing phylogenies. This presents both challenges and opportunities for the study of divergence and diversification. (shrink)

The article defends the doctrine that Linnaean taxa, including species, have essences that are, at least partly, underlying intrinsic, mostly genetic, properties. The consensus among philosophers of biology is that such essentialism is deeply wrong, indeed incompatible with Darwinism. I argue that biological generalizations about the morphology, physiology, and behavior of species require structural explanations that must advert to these essential properties. The objection that, according to current “species concepts,” species are relational is rejected. These concepts are primarily concerned with (...) what it is for a kind to be a species and throw little light on the essentialist issue of what it is for an organism to be a member of a particular kind. Finally, the article argues that this essentialism can accommodate features of Darwinism associated with variation and change. (shrink)

This paper surveys recent philosophy of biology. It aims to introduce outsiders to the field to the recent literature (which is reviewed in the footnotes) and the main recent debates. I concentrate on three of these: recent critiques of the replicator/vehicle distinction and its application to the idea of the gene as the unit of section; the recent defences of group selection and the idea that standard alternatives to group selection are in fact no more than a disguised form of (...) group selection; and recent ideas on the role of selection in evolution, especially the role of selection in structuring the large-scale history of life. The paper connects philosophy of biology to some more general problems in the philosophy of science, and concludes with a few suggestions about unfinished business. (shrink)

I spell out and update the individuality thesis, that species are individuals, and not classes, sets, or kinds. I offer three complementary presentations of this thesis. First, as a way of resolving an inconsistent triad about natural kinds; second, as a phylogenetic systematics theoretical perspective; and, finally, as a novel recursive account of an evolved character. These approaches do different sorts of work, serving different interests. Presenting them together produces a taxonomy of the debates over the thesis, and isolates ways (...) it has been productive. This goes to the larger point of this paper: a defense of the individuality thesis in terms of its utility, and an update of it in light of recent theoretical developments and empirical work in biology. (shrink)

The project of this paper is to understand what a phylogenetic tree represents and to discuss some of the implications that this has for the practice of systematics. At least the first part of this task, if not both parts, might appear trivial—or perhaps better suited for a single page in a textbook rather than a scholarly research paper. But this would be a mistake. While the task of interpreting phylogenetic trees is often treated in a trivial way, their interpretation (...) is tied to foundational conceptual questions at the heart of systematics—questions whose answers are hotly disputed. I have previously argued that widely shared ideas about the meaning and interpretation of phylogenetic trees are inconsistent with species concepts other than some genealogical version of a phylogenetic species concept (Velasco 2008). Here I rely on a similar approach and concentrate on the implications of the necessary conditions underlying the inferences that we make using phylogenetic trees. I argue that common practices for the interpretation and use of trees are in conflict and that unacceptable principles about species as units of phylogeny must be given up. According to the view that I will develop, all phylogenetic trees depict the history of populations. The branches on trees represent collections of population lineages through time and the splits represent population lineage splits. This is true regardless of whether the tips of the trees are themselves populations, or are species or higher taxa. Although this conclusion might be paired naturally with a view that species must be monophyletic groups, this population-centric view of trees is independent of that view of species. If we still want to have species that are paraphyletic groups of populations, this is permissible as long as we also do not treat species as the units of phylogeny. This population-centric view opposes a species-centric view of phylogeny and might be called a “rank-free” approach since it entails that we do not need to determine which groups are species (which is partly a ranking question) in order to build a tree. This conclusion and the argument for it are meant to be consistent with, but not require, acceptance of the conclusions of Velasco (2008) regarding species. (shrink)

The Developmental Systems approach to evolution is defended against the alternative extended replicator approach of Sterelny, Smith and Dickison (1996). A precise definition is provided of the spatial and temporal boundaries of the life-cycle that DST claims is the unit of evolution. Pacé Sterelny et al., the extended replicator theory is not a bulwark against excessive holism. Everything which DST claims is replicated in evolution can be shown to be an extended replicator on Sterelny et al.s definition. Reasons are given (...) for scepticism about the heuristic value claimed for the extended replicator concept. For every competitive, individualistic insight the replicator theorist has a cooperative, systematic blindspot. (shrink)

The variation and selection form of explanationcan be prescinded from the evolutionary biologyhome ground in which it was discovered and forwhich it has been most developed. When this isdone, variation and selection explanations arefound to have potential application to a widerange of phenomena, far beyond the classicalbiological ground and the contemporaryextensions into epistemological domains. Itappears as the form of explanation most suitedto phenomena of fit. It is also found toparticipate in multiple interestingrelationships with other forms of explanation. We proceed with (...) an examination of multiplekinds of phenomena, interrelationships withother members of the family of forms ofexplanation, and some novel applications evenwithin the home ground of evolutionary biology. (shrink)

This paper clarifies the historical nature of species by showing that species are path-dependent entities. A species’ identity is not determined by its intrinsic properties or its origin, but by its unique evolutionary path. Seeing that species are path-dependent entities has three implications: it shows that origin essentialism is mistaken, it rebuts two challenges to the species-are-historical-entities thesis, and it demonstrates that the identity of a species during speciation depends on future events.

The two great modern naturalists, Linnaeus and Darwin, expressed their intuition about how best to visualize patterns of affinities, that is, morphological similarities and divergences between taxa. Linnaeus suggested that “all plants show affinities on all sides, like a territory on a geographical map,” while Darwin thought that it was virtually impossible to understand the affinities between living and extinct species without a genealogical tree. Genealogical trees follow the diachronic, evolving logic of a timeline, whereas maps depict a synchronous pattern (...) of extant taxa. Although the two seem unrelated, various naturalists made attempts to combine them. Surprisingly, these resulted in three-dimensional images that, in order to be observed, had to be projected on paper. The naturalists Max Fürbringer and Richard Bowdler Sharpe were aware of this fact, but even Darwin himself twice combined the basic intuitions underlying the two modes of representation to produce three-dimensional images. This article is a brief history of the efforts to merge genealogical trees and map-like cross sections of affinities into one three-dimensional image. (shrink)

Different concepts define species at the pattern-level grouping of organisms into discrete clusters, the level of the processes operating within and between populations leading to the formation and maintenance of these clusters, or the level of the inner-organismic genetic and molecular mechanisms that contribute to species cohesion or promote speciation. I argue that, unlike single-level approaches, a multi-level framework takes into account the complex sequences of cause-effect reinforcements leading to the formation and maintenance of various patterns, and allows for revisions (...) and refinements of pattern-based characterizations in light of the gradual elucidation of the causes and mechanisms contributing to pattern formation and maintenance. (shrink)