At least 22 concepts of species are in use today and many of these are notably incompatible in their accounts of biological diversity. Much of the traditional turmoil embodied in the species problem ultimately derives from the packaging of inappropriate criteria for species into a single concept. This results from a traditional conflation of function of concepts with their applications, definitions with concepts, taxonomic categories with groups, and the ontological status of real species with teleological approaches to recover them. Analogous (...) to classifications of supraspecific taxa, our forging inappropriate and ambiguous information relating to theoretical and operational discussions of species ultimately results in a trade-off between convenience, accuracy, precision, and the successful recovery of natural biological diversity. Hence, none of these expectations or intentions of species or classifications is attainable through composite, and possibly discordant, concepts of biological diversity or its descent. Reviewing and evaluating the concepts of species for their theoretical and operational qualities illustrates that a monistic, primary concept of species, applicable to the various entities believed to be species, is essential. This evaluation reveals only one theoretical concept as appropriate for species, the Evolutionary Species Concept. This conceptualization functions as a primary concept and is essential in structuring our ideas and perceptions of real species in the natural world. The remaining concepts are secondary, forming a hierarchy of definitional guidelines subordinate to the primary concept, and are essential to the study of species in practice. Secondary concepts should be used as operational tools, where appropriate, across the variance in natural diversity to discover entities in accord with the primary concept. Without this theoretical and empirical structuring of concepts of species our mission to achieve reconciliation and understanding of pattern and process of the natural world will fail. (shrink)

Species concepts aim to define the species category. Many of these rely on defining species in terms of natural lineages and groupings. A dominant gene-centred metaconception has shaped notions of what constitutes both a natural lineage and a natural grouping. I suggest that relying on this metaconception provides an incomplete understanding of what constitute natural lineages and groupings. If we take seriously the role of epigenetic, behavioural, cultural, and ecological inheritance systems, rather than exclusively genetic inheritance, a broader notion of (...) what constitutes a natural grouping or lineage may be required. I conclude by outlining an alternative metaconception that is a de-centred metaschema for species. (shrink)

When developing phylogenetic systematics, the entomologist Willi Hennig adopted elements from Nicolai Hartmann’s ontology. In this historical essay I take on the task of documenting this adoption. I argue that in order to build a metaphysical foundation for phylogenetic systematics, Hennig adopted from Hartmann four main metaphysical theses. These are (1) that what is real is what is temporal; (2) that the criterion of individuality is to have duration; (3) that species are supra-individuals; and (4) that there are levels of (...) reality, each of which may be subject to different kinds of law. Reliance on Hartmann’s metaphysics allowed Hennig to ground some of the main theoretical principles of phylogenetic systematics, namely that the biological categories—from the semaphoront to the highest rank—have reality and individuality despite not being universals, and that they form a hierarchy of levels, each of which may require different kinds of explanation. Hartmann’s metaphysics thereby provided a philosophical justification for Hennig’s phylogenetic systematics, both as a theory and as a method of classification. (shrink)

de Queiroz (1995), Griffiths (1999) and LaPorte (2004) offer a new version of essentialism called "historical essentialism". According to this version of essentialism, relations of common ancestry are essential features of biological taxa. The main type of argument for this essentialism proposed by Griffiths (1999) and LaPorte (2004) is that the dominant school of classification, cladism, defines biological taxa in terms of common ancestry. The goal of this paper is to show that this argument for historical essentialism is unsatisfactory: cladism (...) does not assume that relations of common ancestry are essential attributes of biological taxa. Therefore, historical essentialism is not justified by cladism. (shrink)

Many phylogenetic systematists have criticized the Biological Species Concept (BSC) because it distorts evolutionary history. While defenses against this particular criticism have been attempted, I argue that these responses are unsuccessful. In addition, I argue that the source of this problem leads to previously unappreciated, and deeper, fatal objections. These objections to the BSC also straightforwardly apply to other species concepts that are not defined by genealogical history. What is missing from many previous discussions is the fact that the Tree (...) of Life, which represents phylogenetic history, is independent of our choice of species concept. Some species concepts are consistent with species having unique positions on the Tree while others, including the BSC, are not. Since representing history is of primary importance in evolutionary biology, these problems lead to the conclusion that the BSC, along with many other species concepts, are unacceptable. If species are to be taxa used in phylogenetic inferences, we need a history-based species concept. (shrink)

Biologists and philosophers that debate the existence of the species category fall into two camps. Some believe that the species category does not exist and the term 'species' should be eliminated from biology. Others believe that with new biological insights or the application of philosophical ideas, we can be confident that the species category exists. This paper offers a different approach to the species problem. We should be skeptical of the species category, but not skeptical of the existence of those (...) taxa biologists call 'species.' And despite skepticism over the species category, there are pragmatic reasons for keeping the word 'species.' This approach to the species problem is not new. Darwin employed a similar strategy to the species problem 150 years ago. (shrink)

The concept of individuality as applied to species, an important advance in the philosophy of evolutionary biology, is nevertheless in need of refinement. Four important subparts of this concept must be recognized: spatial boundaries, temporal boundaries, integration, and cohesion. Not all species necessarily meet all of these. Two very different types of pluralism have been advocated with respect to species, only one of which is satisfactory. An often unrecognized distinction between grouping and ranking components of any species concept is necessary. (...) A phylogenetic species concept is advocated that uses a grouping criterion of monophyly in a cladistic sense, and a ranking criterion based on those causal processes that are most important in producing and maintaining lineages in a particular case. Such causal processes can include actual interbreeding, selective constraints, and developmental canalization. The widespread use of the biological species concept is flawed for two reasons: because of a failure to distinguish grouping from ranking criteria and because of an unwarranted emphasis on the importance of interbreeding as a universal causal factor controlling evolutionary diversification. The potential to interbreed is not in itself a process; it is instead a result of a diversity of processes which result in shared selective environments and common developmental programs. These types of processes act in both sexual and asexual organisms, thus the phylogenetic species concept can reflect an underlying unity that the biological species concept can not. (shrink)

Integrating concepts of maintenance and of origins is essential to explaining biological diversity. The unified theory of evolution attempts to find a common theme linking production rules inherent in biological systems, explaining the origin of biological order as a manifestation of the flow of energy and the flow of information on various spatial and temporal scales, with the recognition that natural selection is an evolutionarily relevant process. Biological systems persist in space and time by transfor ming energy from one state (...) to another in a manner that generates structures which allows the system to continue to persist. Two classes of energetic transformations allow this; heat-generating transformations, resulting in a net loss of energy from the system, and conservative transformations, changing unusable energy into states that can be stored and used subsequently. All conservative transformations in biological systems are coupled with heat-generating transformations; hence, inherent biological production, or genealogical proesses, is positively entropic. There is a self-organizing phenomenology common to genealogical phenomena, which imparts an arrow of time to biological systems. Natural selection, which by itself is time-reversible, contributes to the organization of the self-organized genealogical trajectories. The interplay of genealogical (diversity-promoting) and selective (diversity-limiting) processes produces biological order to which the primary contribution is genealogical history. Dynamic changes occuring on times scales shorter than speciation rates are microevolutionary; those occuring on time scales longer than speciation rates are macroevolutionary. Macroevolutionary processes are neither redicible to, nor autonomous from, microevolutionary processes. (shrink)

The theory and practice of contemporary comparative biology and phylogeny reconstruction (systematics) emphasizes algorithmic aspects but neglects a concern for the evidence. The character data used in systematics to formulate hypotheses of relationships in many ways constitute a black box, subject to uncritical assessment and social influence. Concerned that such a state of affairs leaves systematics and the phylogenetic theories it generates severely underdetermined, we investigate the nature of the criteria of homology and their application to character conceptualization in the (...) context of transformationist and generative paradigms. Noting the potential for indeterminacy in character conceptualization, we conclude that character congruence (the coherence of character statements) relative to a hierarchy is a necessary, but not a sufficient, condition for phylogeny reconstruction. Specifically, it is insufficient due to the lack of causal grounding of character hypotheses. Conceptualizing characters as homeostatic property cluster natural kinds is in accordance with the empirical practice of systematists. It also accounts for the lack of sharpness in character conceptualization, yet requires character identification and re-identification to be tied to causal processes. (shrink)

Academia is subdivided into separate disciplines, most of which are quite discrete. In this review I trace the interactions between two of these disciplines: biology and philosophy of biology. I concentrate on those topics that have the most extensive biological content: function, species, systematics, selection, reduction and development. In the final section of this paper I touch briefly on those issues that biologists and philosophers have addressed that do not have much in the way of biological content.

Stephen Jay Gould argued that replaying the ‘tape of life’ would result in radically different evolutionary outcomes. Recently, biologists and philosophers of science have paid increasing attention to the theoretical importance of convergent evolution—the independent origination of similar biological forms and functions—which many interpret as evidence against Gould’s thesis. In this paper, we examine the evidentiary relevance of convergent evolution for the radical contingency debate. We show that under the right conditions, episodes of convergent evolution can constitute valid natural experiments (...) that support inferences regarding the deep counterfactual stability of macroevolutionary outcomes. However, we argue that proponents of convergence have problematically lumped causally heterogeneous phenomena into a single evidentiary basket, in effect treating all convergent events as if they are of equivalent theoretical import. As a result, the ‘critique from convergent evolution’ fails to engage with key claims of the radical contingency thesis. To remedy this, we develop ways to break down the heterogeneous set of convergent events based on the nature of the generalizations they support. Adopting this more nuanced approach to convergent evolution allows us to differentiate iterated evolutionary outcomes that are probably common among alternative evolutionary histories and subject to law-like generalizations, from those that do little to undermine and may even support, the Gouldian view of life. (shrink)

A number of influential biologists are currently pursuing efforts to restore previously extinct species. But for decades, philosophers of biology have regarded “de-extinction” as conceptually incoherent. Once a species is gone, it is gone forever. We argue that a range of metaphysical, biological, and ethical grounds for opposing de-extinction are at best inconclusive and that a pragmatic stance that allows for its possibility is more appealing.

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)

Plant and animal species are information-processing entities of such complexity, integration, and adaptive competence that it may be scientifically fruitful to consider them intelligent. The possibility arises from the analogy between learning and evolution, and from recent developments in evolutionary science, psychology and cognitive science. Species are now described as spatiotemporally localized individuals in an expanded hierarchy of biological entities. Intentional and cognitive abilities are now ascribed to animal, human, and artificial intelligence systems that process information adaptively, and that manifest (...) problem-solving abilities. The structural and functional similarities between such systems and species are extensive, although they “are usually obscured by population-genetic metaphors that have nonetheless contributed much to our understanding of evolution.In this target article, I use Sewall Wright's notion of the “adaptive landscape” to compare the performances of evolving species to those of intelligent organisms. With regard to their adaptive achievements and the kinds of processes by which they are achieved, biological species compare very favorably to intelligent animals by virtue of interactions between populations and their environments, between ontogeny and phylogeny, and between natural, interdemic, organic, and species selection. Addressing the question of whether species are intelligent could help to refine our ideas about species, evolution, and intelligence, and could open new lines of empirical and theoretical inquiry in many disciplines. (shrink)

We tend to think that there are different types of biological taxa: some taxa are species, others are genera, while others are families. Linnaeus gave us his ranks in 1731. Biological theory has changed since Linnaeus's time. Nevertheless, the vast majority of biologists still assign Linnaean ranks to taxa, even though that practice is at odds with evolutionary theory and even though it causes a number of practical problems. The Linnaean ranks should be abandoned and alternative methods for displaying the (...) hierarchical relations of taxa should be adopted. (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)

A natural starting place for developing a phylogenetic species concept is to examine monophyletic groups of organisms. Proponents of “the” Phylogenetic Species Concept fall into one of two camps. The first camp denies that species even could be monophyletic and groups organisms using character traits. The second groups organisms using common ancestry and requires that species must be monophyletic. I argue that neither view is entirely correct. While monophyletic groups of organisms exist, they should not be equated with species. Instead, (...) species must meet the more restrictive criterion of being genealogically exclusive groups where the members are more closely related to each other than to anything outside the group. I carefully spell out different versions of what this might mean and arrive at a working definition of exclusivity that forms groups that can function within phylogenetic theory. I conclude by arguing that while a phylogenetic species concept must use exclusivity as a grouping criterion, a variety of ranking criteria are consistent with the requirement that species can be placed on phylogenetic trees. (shrink)

Marc Ereshefsky argues that pluralism about species suggests that the species concept is not theoretically useful. It is to be abandoned in favor of several concrete species concepts that denote real categories. While accepting species pluralism, the present paper rejects eliminativism about the species category. It is argued that the species concept is important and that it is possible to make sense of a general species concept despite the existence of different concrete species concepts.

The observational-theoretical distinction has played a central role in the philosophy of science from logical empiricism onwards. Developments within the discipline have shown that such a distinction hides two non-identical distinctions: observational vs. non-observational and theoretical vs. non-theoretical. Likewise, metatheoretical structuralism proposed a more sophisticated treatment of theoricity in terms of the operational dependence or independence of concepts from the theories in which they appear. Despite this remarkable sophistication, it is still generally accepted that the distinction between theoretical and non-theoretical (...) concepts coincides with the explanatory vs. non-explanatory and testing vs. non-testing distinctions. Ginnobili and Carman argued for the independence between theoricity and explanation. In this paper we will focus on testing and argue for its independence from both theoricity and explanation. To do so, we propose an account of theory testing based on the framework of metatheoretical structuralism and apply it to natural selection theory and cladistics. (shrink)

According to contemporary understanding of the universal tree of life, the traditionally recognized kingdoms of eukaryotic organisms—Protista, Fungi, Animalia and Plantae—are irregularly interspersed in a vast phylogenetic tree. There are numerous groups that in any Linnaean classification advised by phylogenetic relationships would form sister groups to those kingdoms, therefore requiring us to admit them the same rank. In practice, this would lead to the creation of ca. 25-30 new kingdoms that would now be listed among animals and plants as “major (...) types of life”. This poses problems of an aesthetic and educational nature. There are, broadly speaking, two ways to deal with that issue: a) ignore the aesthetic and educational arguments and propose classification systems that are fully consistent with the Hennigian principles of phylogenetic classification, i.e. are only composed of monophyletic taxa; b) ignore Hennigian principles and bunch small, relatively uncharacteristic groups into paraphyletic taxa, creating systems that are more convenient. In the paper, I present the debate and analyze the pros and cons of both options, briefly commenting on the deeper, third resolution, which would be to abandon classification systems entirely. Recent advances in eukaryotic classification and phylogeny are commented in the light of the philosophical question of the purpose and design principles of biological classification systems. (shrink)

Species are central to biology, but there is currently no agreement on what the adequate species concept should be, and many have adopted a pluralist stance: different species concepts will be required for different purposes. This thesis is a multidimensional analysis of species pluralism. First I explicate how pluralism differs monism and relativism. I then consider the history of species pluralism. I argue that we must re-frame the species problem, and that re-evaluating Aristotle's role in the histories of systematics can (...) shed light on pluralism. Next I consider different forms of pluralism: evolutionary and extra-evolutionary species pluralism, which differ in their stance on evolutionary theory. I show that pluralism is more than a debate about the species category, but a debate about which concepts are legitimate and a claim about how they interact with one another. Following that, I consider what sort of ontology is required for different forms of species pluralism. I argue that pluralists who deny the unity of biology will require a further plurality of frameworks, while those that ground their pluralism in evolution need only one framework. Finally, I consider what pluralism means for biological practice. I argue that species concepts are tools, and reflect on how pluralism can illuminate the way systematists approach the discovery of new species of yeast. Pluralism can make sense of the way species concepts are used, and can be developed to aid researchers in thinking about how to use the right tools for the right jobs. (shrink)

According to the species-as-individuals thesis(hereafter S-A-I), species are cohesive entities. Barker and Wilson recently pointed out that the type of cohesion exhibited by species is fundamentally different from that of organisms (paradigmatic individuals), suggesting that species are homeostatic property cluster kinds. In this article, I propose a shift in how to approach cohesion in the context of S-A-I: instead of analyzing the different types of cohesion and questioning whether species have them, I focus on the role played by cohesion in (...) the identity of individuals. This shift allows us to recognize why cohesion matters to S-A-I, as well as to reconceive the analogy between species and organisms (paradigmatic individuals), and also allows us to highlight the context sensitivity of both ‘‘cohesion’’ and ‘‘individuals.’’ From this perspective, I identify two problems in Barker and Wilson’s argumentation. Firstly, the authors fail to recognize that species are individuals even if they do not have the same type of cohesion that organisms have. Secondly, their argument relies on a misinterpretation of S-A-I. I conclude that species cohesion is still best framed as a feature of species individuality rather than a feature of species as homeostatic property cluster kinds. The arguments presented here contribute to the re-articulation and reevaluation of S-A-I in the face of contemporary discussions. (shrink)

The longstanding species problem in biology has a history that suggests a solution, and that history is not the received history found in many texts written by biologists or philosophers. The notion of species as the division into subordinate groups of any generic predicate was the staple of logic from Aristotle through the middle ages until quite recently. However, the biological species concept during the same period was at first subtly and then overtly different. Unlike the logic sense, which relied (...) on definitions of the essence of both genus and species, biological species from the time of Epicurus were consistently considered to involve a reproductive element: in short, living species relied not on essential definitions, but on the generative cause, which might not be definable. I term this the generative conception of species: species were the generation by reproduction of form. This undercuts the claim that species before Darwin were essentialist, and divorces the notion of a type from that of essence. In fact, as late as the end of the nineteenth century, logicians explicitly treated biological “species” as a homonym only of logical essentialist species, and permitted considerable deviation from the type or form. At every point, species in logic were thought to be a subset, in effect, of some more general notion. I sketch a history of both philosophical and biological traditions of the species concept, before turning to the current conceptions. These are reconsidered in the light of this history, and in particular Mayr’s changing views are shown to be somewhat Whiggish, historiographically. Of the many touted biological species concepts, only one of which (Mayr’s) is called _the_ Biological Species Concept, none appears to capture all the relevant facts, intuitions, and operational requirements of biology. Cladistic conceptions, however, have much in common with the older philosophical literature, in that the natural group of cladism is the clade, or monophyletic group. After considering the Individuality Thesis, and the metaphysics of species, we see that species are the most particular terminal taxa in a clade, and that they are “defined” in terms of the particular synapomorphies, or evolved characters, that are causally responsible for keeping the lineages that organisms form distinct from one another. In this way, we can remain within the generative conception of species that has been in play for over two millennia, and yet avoid the pitfalls of prior attempts to find a universal conception of species. (shrink)

In this article, I examine the issue of the alleged circularity in the determination of homologies within cladistic analysis. More specifically, I focus on the claims made by the proponents of the dynamic homology approach, regarding the distinction (sometimes made in the literature) between primary and secondary homology. This distinction is sometimes invoked to dissolve the circularity issue, by upholding that characters in a cladistic data matrix have to be only primarily homologous, and thus can be determined independently of phylogenetic (...) hypotheses, by using the classical Owenian criteria (for morphological characters) or via multiple sequence alignment (for sequence data). However, since in the dynamic approach, sequence data can be analyzed without being pre-aligned, proponents have claimed that the distinction between primary and secondary homology has no place within cladistics. I will argue that this is not the case, since cladistic practice within the dynamic framework does presuppose primary homology statements at a higher level. (shrink)

Many attempts have been made at supporting either one of the allegedly complementary divergence models Phyletic Gradualism and Punctuated Equilibria by patterns found in specific fossil sequences. However, assessing each model's connection with reality via such “individual case histories” appears not to constitute a relevant approach. Instead, in order to correctly establish the possible merits of both concepts, the claims of each have to be verified against general evolutionary theory. This is being pointed out herein by analyzing cladogenesis at the (...) specific level as an example. [Phyletic Gradualism; Punctuated Equilibria; evolutionary theory; divergence models; case histories; additive speciation.]. (shrink)

A common view is that species occupy a unique position on the Tree of Life. Evaluating this claim requires an understanding of what the Tree of Life represents. The Tree represents history, but there are at least three biological levels that are often said to have genealogies: species, organisms, and genes. Here I focus on defending the plausibility of a gene-based account of the Tree. This leads to an account of species that are determined by gene genealogies. On this view, (...) an exclusive group is a group of organisms that forms a clade for a higher proportion of the genome than any conflicting clade. Taxa occupy a unique position in what can be called the ‘primary concordance tree’. But each gene has its own historical ‘Tree of Life’. I conclude by arguing that both organismal pedigrees with their corresponding Tree as well as gene genealogies and their trees are objectively real and play important, but different, roles in biological practice. (shrink)

The dangers of character reification for cladistic inference are explored. The identification and analysis of characters always involves theory-laden abstraction—there is no theory-free “view from nowhere.” Given theory-ladenness, and given a real world with actual objects and processes, how can we separate robustly real biological characters from uncritically reified characters? One way to avoid reification is through the employment of objectivity criteria that give us good methods for identifying robust primary homology statements. I identify six such criteria and explore each (...) with examples. Ultimately, it is important to minimize character reification, because poor character analysis leads to dismal cladograms, even when proper phylogenetic analysis is employed. Given the deep and systemic problems associated with character reification, it is ironic that philosophers have focused almost entirely on phylogenetic analysis and neglected character analysis. (shrink)

The Linnaean system of classification is a threefold system of theoretical assumptions, sorting rules, and rules of nomenclature. Over time, that system has lost its theoretical assumptions as well as its sorting rules. Cladistic revisions have left it less and less Linnaean. And what remains of the system is flawed on pragmatic grounds. Taking all of this into account, it is time to consider alternative systems of classification.

Contrary to the common-sense view and positivist aspirations, scientific concepts are often imprecise. Many of these concepts are ambiguous, vague, or have an under-specified meaning. In this paper, I discuss how imprecise concepts promote integration in biology and thus benefit science. Previous discussions of this issue focus on the concepts of molecular gene and evolutionary novelty. The concept of molecular gene helps biologists integrate explanatory practices, while the notion of evolutionary novelty helps them integrate research questions into an interdisciplinary problem (...) New directions in the philosophy of science, Springer, Dordrecht, 2014). In what follows, I compare molecular gene and evolutionary novelty to another imprecise concept, namely biological lineage. This concept promotes two other types of scientific integration: it helps biologists integrate theoretical principles and methodologies into different areas of biology. The concept of biological lineage facilitates these types of integration because it is broad and under-specified in ways that the concepts of molecular gene and evolutionary novelty are not. Hence, I use the concept of biological lineage as a case study to reveal types of integration that have been overlooked by philosophers. This case study also shows that even very imprecise concepts can be beneficial to scientific practice. (shrink)

: According to Kuhn, theory choice is not governed by algorithms, but by values, which influence yet do not determine theory choice. Cladistic hypotheses, however, seem to be evaluated relative to a parsimony algorithm, which asserts that the best phylogenetic hypothesis is the one that requires the fewest character changes. While this seems to be an unequivocal evaluative rule, it is not. The application of the parsimony principle is ultimately indeterminate because the choice and individuation of characters that figure in (...) parsimony computations are indeterminate. The cladistic approach is Kuhnian because the application of parsimony depends on persuasion, background, training and tradition. (shrink)

The Darwinian theory of evolution is itself evolving and this book presents the details of the core of modern Darwinism and its latest developmental directions. The authors present current scientific work addressing theoretical problems and challenges in four sections, beginning with the concepts of evolution theory, its processes of variation, heredity, selection, adaptation and function, and its patterns of character, species, descent and life. The second part of this book scrutinizes Darwinism in the philosophy of science and its usefulness in (...) understanding ecosystems, whilst the third section deals with its application in disciplines beyond the biological sciences, including evolutionary psychology and evolutionary economics, Darwinian morality and phylolinguistics. The final section addresses anti-Darwinism, the creationist view and issues around teaching evolution in secondary schools. The reader learns how current experimental biology is opening important perspectives on the sources of variation, and thus of the very power of natural selection. This work examines numerous examples of the extension of the principle of natural selection and provides the opportunity to critically reflect on a rich theory, on the methodological rigour that presides in its extensions and exportations, and on the necessity to measure its advantages and also its limits. Scholars interested in modern Darwinism and scientific research, its concepts, research programs and controversies will find this book an excellent read, and those considering how Darwinism might evolve, how it can apply to the human sciences and other disciplines beyond its origins will find it particularly valuable. Originally produced in French (Les Mondes Darwiniens), the scope and usefulness of the book have led to the production of this English text, to reach a wider audience. This book is a milestone in the impressive penetration by Francophone scholars into the world of Darwinian science, its historiography and philosophy over the last two decades. Alex Rosenberg, R. Taylor Cole Professor of Philosophy, Duke University Until now this useful and comprehensive handbook has only been available to francophones. Thanks to this invaluable new translation, this collection of insightful and original essays can reach the global audience it deserves. Tim Lewens, University of Cambridge. (shrink)

A few philosophers of biology have recently explicitly rejected Essential Membership, the doctrine that if an individual organism belongs to a taxon, particularly a species, it does so essentially. But philosophers of biology have not addressed the broader issue, much discussed by metaphysicians on the basis of modal intuitions, of what is essential to the organism. In this paper, I address that issue from a biological basis, arguing for the Kripkean view that an organism has a partly intrinsic, partly historical, (...) essence. The arguments appeal to the demands of biological explanation and are analogous to arguments that I have given elsewhere that a taxon has a partly intrinsic, partly historical, essence. These conclusions about the essences of individuals and taxa yield an argument for Essential Membership. Finally, I cast doubt on LaPorte’s objection to that doctrine arising from the view that a species cannot survive having a daughter. (shrink)

Quinn offered seven definitions of “cladist” and discussed the context in which they are used in relation to historical and current debates in systematics. As a member of her study taxon, I offer some contextual color commentary, clarifications on the views of “pattern cladists” regarding monophyly, ancestors, synapomorphy and other concepts, a definition of “syncretist”, and some thoughts on cladistics and philosophy in the twenty first century.

Following Mayr (1961) evolutionary biologists often maintain that the hallmark of biology is its evolutionary perspective. In this view, biologists distinguish themselves from other natural scientists by their emphasis on why-questions. Why-questions are legitimate in biology but not in other natural sciences because of the selective character of the process by means of which living objects acquire their characteristics. For that reason, why-questions should be answered in terms of natural selection. Functional biology is seen as a reductionist science that applies (...) physics and chemistry to answer how-questions but lacks a biological point of view of its own. In this paper I dispute this image of functional biology. A close look at the kinds of issues studied in biology and at the way in which these issues are studied shows that functional biology employs a distinctive biological perspective that is not rooted in selection. This functional perspective is characterized by its concern with the requirements of the life-state and the way in which these are met. (shrink)

Biological type specimens are a particular kind of voucher specimen stored in natural history collections. Their special status and practical use are discussed in relation to the description and naming of taxonomic zoological diversity. Our current system, known as Linnaean nomenclature, is governed by the International Code of Zoological Nomenclature. The name of a species is fixed by its name-bearing type specimen, linking the scientific name of a species to the type specimen first designated for that species. The name-bearing type (...) specimen is not necessarily a typical example of the species, while establishment of the boundaries of a species requires empirical taxonomic studies. The International Code of Zoological Nomenclature allows for the naming of new species in the absence of preserved specimens. However, photos and DNA sequences should not function as primary type material, while new species should not be described and named without deposition of at least one type specimen in a collection. Philosophically, species are individuals, spatiotemporally restricted entities. Therefore, Linnaean species names are proper names, which do not define the taxon but serve as a label, providing an ostensive definition of a species. Paratypes have no name-bearing function but, nevertheless, are highly valued specimens in natural history collections. Paratypes should be restricted to those specimens originating from the same sample as the holotype. Diagnosis of a species taxon involves establishment of a connection between a Linnaean name and determination of the boundaries of the species. A first step in this process is the choice of an appropriate species concept. It is not the examination of holotypes and paratypes that necessarily provides the best estimate of the taxonomic boundaries of a species, but this is facilitated by a set of voucher specimens known as the hypodigm. Dissatisfaction with the present nomenclatural code led some researchers to propose emendations. Other taxonomists suggested abandoning Linnaean nomenclature and proposed the alternative PhyloCode, albeit that it relegates the naming of species taxa to the traditional nomenclatural codes. (shrink)

What are biological species? Aristotelians and Lockeans agree that they are natural kinds; but, evolutionary theory shows that neither traditional philosophical approach is truly adequate. Recently, Michael Ghiselin and David Hull have argued that species are individuals. This claim is shown to be against the spirit of much modern biology. It is concluded that species are natural kinds of a sort, and that any 'objectivity' they possess comes from their being at the focus of a consilience of inductions.

Although the term taxon is one of the most common concepts in biology, a range of its meanings cannot be comprehended by an universal definition. Usually, biologists construe their knowledge of “the same” taxon by substantially different interpretations, so they find themselves in need either to justify this “multiplication of taxon essences”, or to surmount their plurality unifying its interpretations into a single explanation of what a taxon is. In both cases, an ontological status (“reality”) of that taxon is questioned. (...) Therefore, discrepancy between universality of the taxon concept in biology and unavoidable plurality of its interpretations can be regarded as a source of problem of the taxon ontology. The present work aims to clarify the premises of this discrepancy using phenomenological approach. Three ways of the taxon positing (as a class, as a place in the world, and as a individualized body) have been distinguished. Taxon as a class is established by common essence that is shared by a set of living beings. These living beings are regarded as speculative objects beyond an idea of the world, i.e. as objects of the experimental science. A question about ontology of taxon as a class refers to the scholastic problem of universalia; its status can be defined within the scope of the nominalism/realism opposition. Taxon as a place of common appearance of the specimens is regarded in the context of the etiological relations unifying various entities into the entire world. Taxon as a place refers to a certain position in the Natural System that is construed as an etiological map of the world. In this case a specimen of a living being is known as a curiosity, i.e. representant of its relationships as well as of the place of its origin. Ontological status of a taxon as a place is to be clarified within the framework of the natural/artificial opposition. The positing of a taxon as a collective body marked off by limits of joint survival of living beings is characteristic for biology in the strict sense which arose in the very beginning of the 19 century. A taxon as a body established by the techniques of disciplinary power sensu M. Foucault extended from the human bodies to bodies of other living beings. The ontological status of a taxon as a collective body can be defined within the scope of the wild/domesticated opposition. Therefore, the discrepancy between the universality of the taxon concept and the plurality of its interpretations is underlayed by interpenetration of three distinct modi of taxon establishing. Distinguishing between these three modi can clarify sources of ontological problems concerned with the term taxon in each case when they arise. (shrink)

This is an essay about ethics and environmental responsibility. The thesis is that biologic species qua species--not only as collections of individuals or as elements of ecosystems--deserve moral regard. The argument establishes moral considerability on powers and freedoms of relative self-determination and autonomy. It is argued that species are living beings in their own right with their own projects and interests which deserve special regard. The essay draws from the arguments of Plato, Aristotle, Plotinus, Boethius, Avicenna, Maimonides, Leibniz, Spinoza, Kant, (...) and others, concerning the moral value of natural beings, but situates itself in the contemporary dialogue of environmental ethics. Lengthy consideration is given to species concepts defended by evolution theorists, biological systematists, and ecologists. ;The argument claims that natural beings, including species, have moral value because they have projects and interests which make a legitimate, but not universal, claim on the moral agent. Beings make moral claims when they are integral and integrating, individual, and concreative--conditions comparable to those with which we describe the moral status of a person. We may call such beings rational by analogy to the integration and singularity of intellect, and confer upon them analogous moral consideration. They are due regard in proportion to their projects in the same sense that moral subjects are due regard for having the projects of moral subjects. (shrink)

Taking its clues from Popperian philosophy of science, cladistics adopted a number of assumptions of the empiricist tradition. These include the identification of a dichotomy between observation reports and theoretical statements and its subsequent abandonment on the basis of the insight that all observation reports are theory-laden. The neglect of the ‘context of discovery’, which is the step of theory (hypothesis) generation. The emphasis on coherentism in the ‘context of justification’, which is the step of evaluation of the relative merits (...) of alternative theories. The appeal to a total evidence approach in phylogenetic inference. And finally, a silence about causation, which results in an instrumentalist approach to phylogeny reconstruction. This paper explores how these empiricist assumptions are embedded in phylogenetic systematics, and why these assumptions are problematic for cladists (or any taxonomists). (shrink)

Recent work on inheritance systems can be divided into inclusive conceptions, according to which genetic and non-genetic inheritance are both involved in the development and transmission of nearly all animal behavioral traits, and more demanding conceptions of what it takes for non-genetic resources involved in development to qualify as a distinct inheritance system. It might be thought that, if a more stringent conception is adopted, homologies could not subsist across two distinct inheritance systems. Indeed, it is commonly assumed that homology (...) relations cannot survive a shift between genetic and cultural inheritance systems, and substantial reliance has been placed on that assumption in debates over the phylogenetic origins of hominin behavioral traits, such as male-initiated intergroup aggression. However, in the homology literature it is widely accepted that a trait can be homologous—that is, inherited continuously in two different lineages from a single common ancestor—despite divergence in the mechanisms involved in the trait’s development in the two lineages. In this paper, we argue that even on an extremely stringent understanding of what it takes for developmental resources to form a separate inheritance system, homologies can nonetheless subsist across shifts between distinct inheritance systems. We argue that this result is a merit of this way of characterizing what it is to be an inheritance system, that it has implications for adjudicating between alternative accounts of homology, and that it offers an important cautionary lesson about how to reason with the homology concept, particularly in the context of cultural species. (shrink)

For the past twenty years, taxonomy has been in a state of turmoil. This confusion brings along with it four distinct schools of thought, each of which offers a different concept of biological species. The thesis will show that these concepts are purely operational and have only a weak theoretical force. In turn, it will be argued that a sound definition of species uses the notion of natural kinds, which is itself defined in term of non-causal nomological regularities.

Despite the amount of work that has been produced on the subject over the years, the ‘transformation of cladistics’ is still a misunderstood episode in the history of comparative biology. Here, I analyze two outstanding, highly contrasting historiographic accounts on the matter, under the perspective of an influential dichotomy in the philosophy of science: the opposition between Scientific Realism and Empiricism. Placing special emphasis on the notion of ‘causal grounding’ of morphological characters in modern developmental biology’s theories, I arrive at (...) the conclusion that a ‘new transformation of cladistics’ is philosophically plausible. This ‘reformed’ understanding of ‘pattern cladistics’ entails retaining the interpretation of cladograms as ‘schemes of synapomorphies’, but in association to construing cladogram nodes as ‘developmental-genetic taxic homologies’, instead of ‘standard Darwinian ancestors’. The reinterpretation of pattern cladistics presented here additionally proposes to take Bas Van Fraassen’s ‘constructive empiricism’ as a philosophical stance that could properly support such analysis of developmental-genetic data for systematic purposes. The latter suggestion is justified through a reappraisal of previous ideas developed by prominent pattern cladists , which concerned a scientifically efficient ‘observable/non-observable distinction’ linked to the conceptual pair ‘ontogeny and phylogeny’. Finally, I argue that a robust articulation of Antirealist alternatives in systematics may provide a rational basis for its disciplinary separation from evolutionary biology, as well as for a critical reconsideration of the proper role of certain Scientific Realist positions, currently popular in comparative biology. (shrink)

The philosophy of pattern cladism has been variously explained by reference to the work of Louis Agassiz. The present study analyzes Agassiz's attempt to combine an empirical approach to the study of nature with an idealistic philosophy. From this emerges the problem of empiricism and of the isomorphy between the order of nature and human thinking. The analysis of the writings of Louis Agassiz serves as the basis for discussion of the reality of natural groups as postulated by pattern cladists.

In this paper, I show how idealizations contribute to social activities in science, such as the recruitment of experts to a research project. These contributions have not been explicitly discussed by recent philosophical accounts of scientific idealization. These accounts have focused on how idealizations influence activities like scientific theorization, explanation, and modeling. Other accounts focus on how idealizations influence policy-making and science communication. I expand these accounts by exploring the uses of idealized phylogenetic trees in science. Trees are not only (...) useful for improving our understanding and public communication of evolutionary history, but they also help with the organization of laboratories and collaboration among scientists. Attending to the relation between idealizations and these social practices in science matters. It can help us understand why idealized models become entrenched and why certain social practices change over time. (shrink)

An examination of the post-Darwinian history of biological taxonomy reveals an implicit assumption that the definitions of taxon names consist of lists of organismal traits. That assumption represents a failure to grant the concept of evolution a central role in taxonomy, and it causes conflicts between traditional methods of defining taxon names and evolutionary concepts of taxa. Phylogenetic definitions of taxon names (de Queiroz and Gauthier 1990) grant the concept of common ancestry a central role in the definitions of taxon (...) names and thus constitute an important step in the development of phylogenetic taxonomy. By treating phylogenetic relationships rather than organismal traits as necessary and sufficient properties, phylogenetic definitions remove conflicts between the definitions of taxon names and evolutionary concepts of taxa. The general method of definition represented by phylogenetic definitions of clade names can be applied to the names of other kinds of composite wholes, including populations and biological species. That the names of individuals (composite wholes) can be defined in terms of necessary and sufficient properties provides the foundation for a synthesis of seemingly incompatible positions held by contemporary individualists and essentialists concerning the nature of taxa and the definitions of taxon names. (shrink)

College of Medicine, University of South Alabama Mobile, AL 36688-0002, USA wbp501{at}jaguar1.usouthal.edu ' + u + '@' + d + ' '//--> Abstract Recent work in viral genomics has shown that bacteriophages exhibit a high degree of mosaicism, which is most likely due to a long history of prolific horizontal gene transfer (HGT). Given these findings, we argue that each of the most plausible attempts to properly classify bacteriophages into distinct species fail. Mayr's biological species concept fails because there is (...) no useful viral analog to sexual reproduction. Phenetic species concepts fail because they obscure the mosaicism and the rich reticulated viral histories. Phylogenetic species concepts, even when extended to take into account reticulation, fail because there is no non-arbitrary distinction between recombination events that create a new viral species and those that do not. There is good reason to think that bacteriophages, arguably the Earth's most abundant biological agent, evolve without forming species. Introduction The Biology of Viruses 2.1 Bacteriophage life cycles 2.2 Mechanisms of HGT The Species Problem and Species Concepts 3.1 Phenetic species concepts 3.2 The biological species concept 3.3 Phylogenetic species concepts 3.4 The ecological species concept 3.5 Homeostatic property cluster species Viruses and Species Taxonomy Reticular Phylogenies Conclusion CiteULike Connotea Del.icio.us What's this? (shrink)

Species serve as both the basic units of macroevolutionary studies and as the basic units of taxonomic classification. In this paper I argue that the taxa identified as species by the Phylogenetic Species Concept (Mishler and Brandon 1987) are the units of biological organization most causally relevant to the evolutionary process but that such units exist at multiple levels within the hierarchy of any phylogenetic lineage. The PSC gives us no way of identifying one of these levels as the privileged (...) level on which taxonomic classifications can be based. (shrink)

The term “cladist” has distinct meanings in distinct contexts. Communication between philosophers, historians, and biologists has been hindered by different understandings of the term in various contexts. In this paper I trace historical and conceptual connections between several broadly distinct senses of the term “cladist”. I propose seven specific definitions that capture distinct contemporary uses. This serves to disambiguate some cases where the meaning is unclear, and will help resolve apparent disagreements that in fact result from conflicting understandings of the (...) term. (shrink)