It is often said that there is just one “objective” tree of life: a single accurate branching hierarchy of species reflecting order of descent. For any two species there is a single correct answer as to whether one is a “daughter” of the other, whether the two are “sister species” by virtue of their descent from a common parental species, whether they belong to a family line that excludes any given third species, and so on. This position is not right. (...) We may whittle a tree of life, paring troublesome branches, in order to portray an ordering that admits of no legitimate dissent. But the history of life can sustain many legitimate arrangements of the same branches of species. The same can be said about other taxonomically relevant groups besides species, such as “Least Inclusive Taxonomic Units” (LITUs), so the basic claim survives even if we abandon traditional species. Similarly, the claim survives even if we distinguish between synchronic and diachronic groups, even if we consider polytomies, even if we distinguish between models and the world modeled, and even if we recognize an objective world. Nor is the claim merely an epistemic one. (shrink)

The species category is defined as thesmallest historical individual within which there is a parental pattern of ancestry and descent. The use of historical individual in this definition is consistent with the prevailing notion that speciesper se are not involved in processes — they are effects, not effectors. Reproductive isolation distinguishes biparental historical species from their parts, and it provides a basis for understanding the nature of the evidence used to discover historical individuals.

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)

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)

Weak links, in the form of inadequacies in both reasoning and supporting evidence, exist at several critical steps in the derivation of an hierarchical concept of evolution from punctuated equilibria. Punctuation itself is predicated on a distorted reading of phyletic change as phyletic gradualism, and of allopatric speciation as the instantaneous formation of unchanging typological taxa. The concept of punctuation is further confounded by the indescriminate employment of the same term to denote both a causal explanation for evolutionary change and (...) an outcome of substantiated evolutionary processes. Even when the intended usage for the term is specified, each denotation of punctuation entails respective drawbacks. As a causal explanation, punctuation clearly belongs to the class of quantum theories with all their attendant impedimenta, including special salsatory non-adaptive mechanisms of evolutionary change. Redefinition of punctuation as a pattern of morphologic change reduces it to one possible outcome of known microevolutioanry processes, thus obviating any need for an hierarchical explanation of macroevolution. While vacillation between usages has preserved the term in the literature, the end result of this obfuscation has been a circle of faulty reasoning in which the pattern of punctuation is invoked as its own proof. Widespread confusion concerning what constitutes an adequate test of punctuation is directly attributable to imprecision in both the original and revised formulations of the concept.The argument for species-level selection is based on the typological and philosphically flawed premise of species as individuals, and further requires the hypothesis of heritable emergent properties, for which empirical evidence is lacking. (shrink)

Many authors, including paleobiologists, cladists and so on, adopt a nested hierarchical viewpoint to examine the relationships among different levels of biological organization. Furthermore, species are often considered to be unique entities in functioning evolutionary processes and one of the individuals forming a nested hierarchy.I have attempted to show that such a hierarchical view is inadequate in evolutionary biology. We should define units depending on what we are trying to explain. Units that play an important role in evolution and ecology (...) do not necessarily form a nested hierarchy. Also the relationships among genealogies at different levels are not simply nested. I have attempted to distinguish the different characteristics of passages when they are used for different purposes of explanation. In my analysis, species and monophyletic taxa cannot be uniquely defined as single units that function in ecological and evolutionary processes. (shrink)

The Empiricist or Lockean view says natural kinds do not exist objectively in nature but are practical categories reflecting use of words. The Modern, Ostensive view says they do exist, and one can refer to such a kind by ostention and recursion, assuming his designation of it is related causally to the kind itself. However, this leads to a problem: Kinds are abstract repeatables, and it seems impossible that abstractions could have causal force. In defence of the Modern view, I (...) suggest we can think of kinds as — or as like — ecological niches existing in nature, which are causally effective by virtue of the fact that they predictively determine (some) properties of the things that happen to occupy them. (shrink)

This article describes developments in the use of information technology in the biological discipline of taxonomy, using both a historical overview and a detailed case study of a particular information systems project. Taxonomy has experienced problems with both its scientific legitimacy and its utility to other biologists. IT has been introduced into the discipline m response to these perceived problems. The information systems project described here served as a means of managing the tensions between scientific legitimacy and utility. It is (...) argued that this project represents an example of the use of a technological development m an attempt to re-engineer a discipline. The development of the information system is analyzed as an attempt to develop a scientific instrument that will embody a particular model of the discipline. The concerns of taxonomy with status and legitimacy make it appropriate that this new technology should be introduced at the interface between the discipline and the rest of biology as a means of disseminating results, and thus come to represent the discipline and the plants described to outsiders, just as the system represents outsiders to taxonomists. (shrink)

The relation between method, concept and theory in science is complicated. I seek to shed light on that relation by considering an instance of it in systematics: The additional challenges phylogeneticists face when reconstructing phylogeny not at a single level, but simultaneously at multiple levels of the hierarchy. How does this complicate the task of phylogenetic inference, and how might it inform and shape the conceptual foundations of phylogenetics? This offers a lens through which the interplay of method, theory and (...) concepts may be understood in systematics, which, in turn, provides data for a more general account. (shrink)

The term adaptive radiation has been recurrently used to describe evolutionary patterns of several lineages, and has been proposed as the main driver of biological diversification. Different definitions and criteria have been proposed to distinguish an adaptive radiation, and the current literature shows disagreements as to how radiating lineages should be circumscribed. Inconsistencies increase when authors try to differentiate a clade under adaptive radiation from clades evolving under ‘regular’ speciation with adaptation, a pattern anticipated and predicted by the evolutionary theory (...) in any lineage. The most important disagreement is as to which evolutionary rate authors analyze to characterize a radiation; a discussion embedded in a prevailing inability to provide mechanistic explanations of the relationship among evolutionary rates. The union of pattern and process in the same term, the inadequacy of reported null hypotheses, and the frequent use of ad hoc comparisons between lineages have also contributed to the lack of consensus. A rigorous use of available terms and the articulation of solid criteria with objective methodologies in distinguishing evolutionary patterns are imperative. Given the difficulties in detecting adaptation, the use of the ‘adaptive’ term to qualify a radiation should be avoided unless methodologically tested. As an unambiguous method to distinguish radiating lineages, the statistical detection of significant increases in taxonomic diversification rates on monophyletic lineages can be considered a distinctive signature of a radiation. After recognizing this pattern, causal hypotheses explaining them can be stated, as well as correlates with other rates of evolution. (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)

Since the 1970s, there has been a tremendous amount of literature on Ghiselin's proposal that species are individuals. After recalling the origins and stakes of this thesis in contemporary evolutionary theory, I show that it can also be found in the writings of the French naturalist Buffon in the 18th Century. Although Buffon did not have the conception that one species could be derived from another, there is an interesting similarity between the modern argument and that of Buffon regarding the (...) individuality of species. The analogy is strong enough to force us to recognize that genuine evolutionary (or Darwinian) questions might be of secondary importance in the discussion. In consequence, the third section of the paper proposes an alternative schema for the logical structure of the Darwinian concept of species. Darwin distinguished the problem of the designation of a concrete species, and the problem of its signification of species within his theory of descent? The resulting notion of species involves a logical structure based on the fusion of the logical operations of classification and ordering. The difficulty — and interest — is that this interpretation of species does not entail any precise operational definition of species; it can only tell us what the ultimate signification of classification is within the theory of descent with modification through natural selection. (shrink)

Prototypes and fuzziness are regarded in this paper as fundamental phenomena in the inherent logic of concepts whose relationship, however, has not been sufficiently clarified. Therefore, modifications are proposed in the definition of both. Prototypes are defined as the elements possessing maximal degree of membership in the given category such thatthis membership has maximal cognitive efficiency in representing theelement. A modified fuzzy set (m-fuzzy set) is defined on aclass (possibly self-contradictory collection) such that its core (the collection of elements with (...) full membership) is aset (self-consistent collection) comprising the finite set of the prototypes. In this scheme of recursive representation the possibility of contradictions, which has been proven to be inevitable in the logic of concepts, islocalized. Finally a related model of the brain''s pattern recognition mechanism is briefly summarized. (shrink)

Philosophical discussions of species have focused on multicellular, sexual animals and have often neglected to consider unicellular organisms like bacteria. This article begins to fill this gap by considering what species concepts, if any, apply neatly to the bacterial world. First, I argue that the biological species concept cannot be applied to bacteria because of the variable rates of genetic transfer between populations, depending in part on which gene type is prioritized. Second, I present a critique of phylogenetic bacterial species, (...) arguing that phylogenetic bacterial classification requires a questionable metaphysical commitment to the existence of essential genes. I conclude by considering how microbiologists have dealt with these biological complexities by using more pragmatic and not exclusively evolutionary accounts of species. I argue that this pragmatism is not borne of laziness but rather of the substantial conceptual problems in classifying bacteria based on any evolutionary standard. (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.

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)

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)

This paper argues that typical biological species are natural kinds, on a familiar realist understanding of natural kinds—classes of individuals across which certain properties cluster together, in virtue of the causal workings of the world. But the clustering is far from exceptionless. Virtually no properties, or property-combinations, characterize every last member of a typical species—unless they can also appear outside the species. This motivates some to hold that what ties together the members of a species is the ability to interbreed, (...) others that it is common descent. Yet others hold that species are scattered individuals,of which organisms are parts rather than members. But not one of these views absolves us of the need to posit a typical phenotypic profile. Vagueness is here to stay. Some seek to explain the vagueness by saying species are united by “homeostatic property clusters”; but this view collapses into the more familiar realist picture. (shrink)

The notion of the ‘division of physiological labour’ is today an outdated relic in the history of science. This contrasts with the fate of another notion, which was so frequently paired with the division of physiological labour, which is the concept of ‘morphological differentiation.’ This is one of the elementary modal concepts of ontogenesis. In this paper, we intend to target the problems and causes that gradually led biologists to combine these two notions during the 19th century, and to progressively (...) dissociate them, retaining only the concept of differentiation by the early 20th century. We shall adhere to the following: 1. The primitive economic concept of the division of labour is not a descriptive notion denoting a type of organisation of labour, but an etiological one: the idea of a causal relationship between this type of organization and the improvement of the whole. 2. This concept rapidly interested naturalists such as Henri Milne-Edwards, who were keen to find a rational ground for hierarchizing living forms based on anatomical complexity. 3. The validation of this notion in the realms of biology was subject to at least two conditions which were far from being fully satisfied. This did not prevent, however, the initial success of the concept of the division of physiological labour during the second half of the 19th century. 4. Finally, the gradual disqualification, within the Darwinian theoretical context, of the conception of an intrinsic hierarchical rank of organisms, led to a lack of interest in the concept of the physiological division of labour, at least in its non-Darwinian and non-ecological variant (the link between the division of labour within an organism and organic perfection). (shrink)

The notion of the ‘division of physiological labour’ is today an outdated relic in the history of science. This contrasts with the fate of another notion, which was so frequently paired with the division of physiological labour, which is the concept of ‘morphological differentiation.’ This is one of the elementary modal concepts of ontogenesis. In this paper, we intend to target the problems and causes that gradually led biologists to combine these two notions during the 19th century, and to progressively (...) dissociate them, retaining only the concept of differentiation by the early 20th century. We shall adhere to the following: 1. The primitive economic concept of the division of labour is not a descriptive notion denoting a type of organisation of labour, but an etiological one: the idea of a causal relationship between this type of organization and the improvement of the whole. 2. This concept rapidly interested naturalists such as Henri Milne-Edwards, who were keen to find a rational ground for hierarchizing living forms based on anatomical complexity. 3. The validation of this notion in the realms of biology was subject to at least two conditions which were far from being fully satisfied. This did not prevent, however, the initial success of the concept of the division of physiological labour during the second half of the 19th century. 4. Finally, the gradual disqualification, within the Darwinian theoretical context, of the conception of an intrinsic hierarchical rank of organisms, led to a lack of interest in the concept of the physiological division of labour, at least in its non-Darwinian and non-ecological variant. (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)

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)

Biologists and philosophers have long recognized the importance of species, yet species concepts serve two masters, evolutionary theory on the one hand and taxonomy on the other. Much of present-day evolutionary and systematic biology has confounded these two roles primarily through use of the biological species concept. Theories require entities that are real, discrete, irreducible, and comparable. Within the neo-Darwinian synthesis, however, biological species have been treated as real or subjectively delimited entities, discrete or nondiscrete, and they are often capable (...) of being decomposed into other, smaller units. Because of this, biological species are generally not comparable across different groups of organisms, which implies that the ontological structure of evolutionary theory requires modification. Some biologists, including proponents of the biological species concept, have argued that no species concept is universally applicable across all organisms. Such a view means, however, that the history of life cannot be embraced by a common theory of ancestry and descent if that theory uses species as its entities.These ontological and biological difficulties can be alleviated if species are defined in terms of evolutionary units. The latter are irreducible clusters of reproductively cohesive organisms that are diagnosably distinct from other such clusters. Unlike biological species, which can include two or more evolutionary units, these phylogenetic species are discrete entities in space and time and capable of being compared from one group to the next. (shrink)

This paper explains the metaphysical implications of the view that species are individuals (SAI). I first clarify SAI in light of the separate distinctions between individuals and classes, particulars and universals, and abstract and concrete things. I then show why the standard arguments given in defense of SAI are not compelling. Nonetheless, the ontological status of species is linked to the traditional "species problem," in that certain species concepts do entail that species are individuals. I develop the idea that species (...) names are rigid designators and show how this provides additional motivation for SAI. (shrink)

After decades of debates about species concepts, there is broad agreement that species are evolving lineages. However, species classification is still in a state of disorder: different methods of delimitation lead to competing outcomes for the same organisms, and the groups recognised as species are of widely different kinds. This paper considers whether this problem can be resolved by developing a unitary scale for evolutionary independence. Such a scale would show clearly when groups are comparable and allow taxonomists to choose (...) a conventional threshold of independence for species status. Existing measurement approaches to species delimitation are typically shot down by what I call the heterogeneity objection, according to which independently evolving groups are too heterogeneous to be captured by a single scale. I draw a parallel with the measurement of temperature to argue that this objection does not provide sufficient reasons to abandon the measurement approach, and that such an approach may even help to make the vague notion of evolutionary independence more precise. (shrink)

The recognition of species proceeds by two fairly distinct phases: (1) the sorting of individuals into groups or basic taxa (‘discovery’) (2) the checking of those taxa as candidates for species-hood (‘justification’). The target here is a rational reconstruction of phase 1, beginning with a discussion of key terms. The transmission of ‘meaning’ is regarded as bimodal: definition states the intension of the term, and diagnosis provides a disjunction of criteria for recognition of its extension. The two are connected by (...) a spectrum, with purely theoretical definition at one pole and purely ‘operational’ diagnosis at the other, with the more operational elements explained by the more theoretical. The current plethora of species concepts provides a good example. Accepting the Ghiselin–Hull thesis, that a species is an individual, a basic taxon is therefore also an individual with organisms as its parts. In a generalised synchronic individual its parts are conceptually integrated by an integrating principle (IP), which consists of a relation applying within a plan or rule. Fully developed, such an IP ensures the maximisation of the information content of the individual. A diachronic individual is then the set of its component synchronic parts, and its IP is provided by near-identity in an appropriate space-time (not necessarily physical). The integration of parts of an individual is illuminated by Gasking's concept of a proper group (in this case a chain-group), whose members are related by the relation serially fitting together with, the IP completed by an appropriate plan or rule. Gasking also applied the term cluster to a proper group persisting over a substantial period of time, individuated by any member acting as focus. A basic taxon is therefore a cluster of individuals, integrated by the relation significantly taxonomically similar and the rule in character-space-time. The nature of those concepts is discussed and defended. A species will inherit certain of the attributes of the preceding basic taxon (taxa). In at least the synchronic version its parts (individuals) will resemble each other significantly, providing the intuitive applicability of ‘members of’ and ‘instances of’ a species. Also, the notion that a species’ name is given by pure ostension, via a name-bearer (holotype) is empirically incoherent: the name cannot be applied in practice without an appropriate set of diagnostic traits. (shrink)

Current issues concerning the nature of ancestry and homology are discussed with reference to the evolutionary origin of the tetrapod limb. Homologies are argued to be complex conjectural inferences dependant upon a pre-existing phylogenetic analysisand a theoretical model of the evolutionary development of ontogenetic information. Ancestral conditions are inferred primarily from character (synapomorphy/homology) distributions within phylogeny, because of the deficiencies of palaeontological data. Recent analyses of tetrapod limb ontogeny, and the diverse, earliest morphologies known from the fossil record, are inconsistent (...) with typological concepts such as fixed ancestral patterns or bauplans, emphasising the incompatibility of these with evolutionary continuity. The evolutionary origin of the tetrapod limb is also examined in the light of its recent discussion in developmental genetics. While this field promises to reveal more of the fundamental ontogenetic content of homology (identity), at present it is concerned mostly with the abstraction of a new set of types, rather than investigating diversity and change. (shrink)

Richard Owen has been condemned by Darwinians as an anti-evolutionist and an essentialist. In recent years he has been the object of a revisionist analysis intended to uncover evolutionary elements in his scientific enterprise. In this paper I will examine Owen's evolutionary hypothesis and its connections with von Baer's idea of divergent development. To give appropriate importance to Owen's evolutionism is the first condition to develop an up-to-date understanding of his scientific enterprise, that is to disentagle Owen's contribution to the (...) modernization of typology and morphology. I will argue that Owen's Platonic essentialism is rhetorical and incongruous. On the contrary, an interpretation of the archetype based on Aristotle's biological works makes possible a new conception of type, based on a homeostatic mechanism of stability. The renewal of morphology hinges on homological correspondences and a homeostatic process is also the origin of serial and special homology. I will argue that special homology shows an evolutionary orientation insofar as it is a typically inter-specific character while serial homology is determined through an elementary usage of the categories of developmental morphology. (shrink)

Ideas about the natural world are intertwined with the personalities, practices, and the workplaces of scientists. The relationships between these categories are explored in the life of the taxonomist William Steel Creighton. Creighton studied taxonomy under William Morton Wheeler at Harvard University. He took the rules he learned from Wheeler out of the museum and into the field. In testing the rules against a new situation, Creighton found them wanting. He sought a new set of taxonomic principles, one he eventually (...) found in Ernst Mayr's "Systematics and the Origin of Species". Mayr's ideas tied together a number of themes running through Creighton's life: the need for a revised taxonomy, the emphasis on fieldwork, and the search for a new power center for ant taxonomy after Wheeler died. Creighton's adoption of Mayr's ideas as part of his professional identity also had very real implications for his career path: field studies required long and intensive studies, and Creighton would always be a slow worker. His method of taxonomy contrasted sharply not only with Wheeler's but also with two of his younger colleagues, William L. Brown and E. O. Wilson, who took over Wheeler's spot at Harvard in 1950. The disputes between these men over ant taxonomy involved, in addition to questions of technical interest, questions about where and how best to do taxonomy and who could speak with the most authority. Creighton's story reveals how these questions are interrelated. The story also reveals the importance of Mayr's book for changes occurring in taxonomy in the middle of the twentieth century. (shrink)

Günter Wagner’s Homology, Genes, and Evolutionary Innovation is a compelling, and empirically well-supported account of the evolution of character identity and character origination which emphasizes the importance of homology and novelty as central explananda for 21st century evolutionary biology. In this essay review, I focus on the similarities and differences between the structuralist picture of evolutionary biology advocated by Wagner, and that presented by standard evolutionary theory. First, I outline the ways in which Wagner’s genetic theory of homology diverges from (...) the account of homology offered by standard evolutionary theory. Then, I consider the motivations for these divergences. Lastly, I discuss a number of concerns with Wagner’s view, and offer some concluding thoughts on the relationship between structuralism and adaptationism. (shrink)

By linking the concepts of homology and morphological organization to evolvability, this paper attempts to (1) bridge the gap between developmental and phylogenetic approaches to homology and to (2) show that developmental constraints and natural selection are compatible and in fact complementary. I conceive of a homologue as a unit of morphological evolvability, i.e., as a part of an organism that can exhibit heritable phenotypic variation independently of the organism’s other homologues. An account of homology therefore consists in explaining how (...) an organism’s developmental constitution results in different homologues/characters as units that can evolve independently of each other. The explanans of an account of homology is developmental, yet the very explanandum is an evolutionary phenomenon: evolvability in a character-by-character fashion, which manifests itself in phylogenetic patterns as recognized by phylogenetic approaches to homology. While developmental constraints and selection have often been viewed as antagonistic forces, I argue that both are complementary as they concern different parts of the evolutionary process. Developmental constraints, conceived of as the presence of the same set of homologues across phenotypic change, pertain to how heritable variation can be generated in the first place (evolvability), while natural selection operates subsequently on the produced variation. (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.

Homology is a natural kind term and a precise account of what homology is has to come out of theories about the role of homologues in evolution and development. Definitions of homology are discussed with respect to the question as to whether they are able to give a non-circular account of the correspondence or sameness referred to by homology. It is argued that standard accounts tie homology to operational criteria or specific research projects, but are not yet able to offer (...) a concept of homology that does not presuppose a version of homology or a comparable notion of sameness. This is the case for phylogenetic definitions that trace structures back to the common ancestor as well as for developmental approaches such as Wagner's biological homology concept. In contrast, molecular homology is able to offer a definition of homology in genes and proteins that explicates homology by reference to more basic notions. Molecular correspondence originates by means of specific features of causal processes. It is speculated that further understanding of morphogenesis might enable biologists to give a theoretically deeper definition of homology along similar lines: an account which makes reference to the concrete mechanisms that operate in organisms. (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)

Cladism, today the dominant school of systematics in biology, includes a classification component – the view that classification ought to reflect phylogeny only, such that all and only taxa are monophyletic (i.e. consist of an ancestor and all its descendants) - and a metaphysical component – the view that all and only real groups or kinds of organisms are monophyletic. For the most part these are seen as amounting to much the same thing, but I argue they can and should (...) be distinguished, in particular that cladists about classification need not accept the typically cladist view about real groups or kinds. Cladists about classification can and should adopt an explanatory criterion for the reality of groups or kinds, on which being monophyletic is neither necessary nor sufficient for being real or natural. Thus the line of reasoning that has rightly led to cladism becoming dominant within systematics, and the attractive line of reasoning in the philosophical literature that advocates a more liberal approach to natural kinds, are seen to be, contrary to appearances, compatible. (shrink)

Over the decades, there has been substantial empirical evidence showing that the unity of species cannot be maintained by gene flow. The biological species concept is inconclusive on this point. The suggestion is made that the unity of species is maintained rather by selection constantly spreading new alleles throughout the species, or bygene circulation. There is a lack in conceptual distinction between gene flow and gene circulation which lies at the heart of the problem. The concept of gene circulation also (...) sheds some new light on the problem of typology and on such a broad concept as evolution. A new species definition is proposed. (shrink)

-/- Because species names play an important role in scientific communication, it is more important that species be understood to be taxa than that they be equated with functional ecological or evolutionary entities. Although most biologists would agree that taxa are composed of organisms that share a unique common history, 2 major challenges remain in developing a species-as-taxa concept. First, grouping: in the face of genealogical discordance at all levels in the taxonomic hierarchy, how can we understand the nature of (...) taxa? Second, ranking: what criteria should be used to designate certain taxa in a nested series as being species? The grouping problem can be solved by viewing taxa as exclusive groups of organisms— sets of organisms that form a clade for a plurality of the genome (more than any conflicting set). However, no single objective criterion of species rank can be proposed. Instead, the species rank should be assigned by practitioners based on the semisubjective application of a set of species-ranking criteria. Although these criteria can be designed to yield species taxa that approximately match the ecological, evolutionary, and morphological entities that taxonomists have traditionally associated with the species rank, such a correspondence cannot be enforced without undermining the assumption that species are taxa. The challenge and art of monography is to use genealogical and other kinds of data to assign all organisms to one and only one species-ranked taxon. Various implications of the species-as-ranked-taxa view are discussed, including the synchronic nature of taxa, fossil species, the treatment of hybrids, and species nomenclature. I conclude that, although challenges remain, adopting the view that species are ranked taxa will facilitate a much-needed revolution in taxonomy that will allow it to better serve the biodiversity informatic needs of the 21st century. (shrink)

Pluralisms of various sorts are popular in philosophy of science, including those that imply some scientific concept x should be eliminated from science in favour of a plurality of concepts x1, x2, … xn. This article focuses on influential and representative arguments for such eliminative pluralism about the concept species. The main conclusions are that these arguments fail, that all other extant arguments also fail, and that this reveals a quite general dilemma, one that poses a defeasible presumption against many (...) eliminative pluralisms about various scientific concepts. The article ends by outlining a novel integrative alternative in defence of species. 1) Introduction 2) The species Concept, the Category ‘Species’, and the ‘Species’ Category Problem 3) What Are Eliminative Pluralism about species, and the Arguments for It? 4) Evaluation of Arguments 4.1 Splitting? 4.2 Lumping? 4.3 The eliminative pluralist’s dilemma 5) More General Lessons6Species Cohesion: An Integrative Alternative7Conclusion. (shrink)

The circumscription of taxa and classification of organisms are fundamental tasks in the systematization of biological diversity. Their success depends on a unified idea concerning the species concept, evolution, and taxonomy; paradoxically, however, it requires a complete distinction between taxa and evolutionary units. To justify this view, I discuss these three topics of systematics. Species concepts are examined, and I propose a redefinition for the Taxonomic Species Concept based on nomenclatural properties, in which species are classes conventionally represented by a (...) binomial. Speciation is subsequently discussed, to demonstrate that concepts on the evolutionary process are damaged when species are considered as evolutionary units. Speciation should be considered a transition among patterns of a population and, consequently, always a sympatric process. This view contrasts with the majority of speciation models, which analyze cladogeneses and classify speciation geographically, usually considering allopatry the essential condition for the differentiation process. Finally, taxonomy is considered, to show that the equivalence between evolutionary and taxonomic units may also damage the practice of biological systematization. Principles and rules of nomenclature and classification should offer freedom to accommodate divergent opinions about systematics and to incorporate the evolution of knowledge; therefore, they should remain independent of biological theories. (shrink)

The present paper analyzes the use and understanding of the homology concept across different biological disciplines. It is argued that in its history, the homology concept underwent a sort of adaptive radiation. Once it migrated from comparative anatomy into new biological fields, the homology concept changed in accordance with the theoretical aims and interests of these disciplines. The paper gives a case study of the theoretical role that homology plays in comparative and evolutionary biology, in molecular biology, and in evolutionary (...) developmental biology. It is shown that the concept or variant of homology preferred by a particular biological field is used to bring about items of biological knowledge that are characteristic for this field. A particular branch of biology uses its homology concept to pursue its specific theoretical goals. (shrink)

The essentialism story is a version of the history of biological classification that was fabricated between 1953 and 1968 by Ernst Mayr, who combined contributions from Arthur Cain and David Hull with his own grudge against Plato. It portrays pre-Darwinian taxonomists as caught in the grip of an ancient philosophy called essentialism, from which they were not released until Charles Darwin's 1859 Origin of Species. Mayr's motive was to promote the Modern Synthesis in opposition to the typology of idealist morphologists; (...) demonizing Plato served this end. Arthur Cain's picture of Linnaeus as a follower of 'Aristotelian' (scholastic) logic was woven into the story, along with David Hull's application of Karl Popper's term, 'essentialism', which Mayr accepted in 1968 as a synonym for what he had called 'typological thinking'. Although Mayr also pointed out the importance of empiricism in the history of taxonomy, the essentialism story still dominates the secondary literature. The history of the first telling of the essentialism story exposes its scant basis in fact. (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)

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)

The genealogy-based classificatory programs of Kant and Darwin are briefly discussed for context. It is detailed how in biology there is no unambiguous term to reference infraspecific-level descent-based divisions. The term lineage population is introduced and defined for analytic purposes as one of a set of inter-fertile divisions of organisms into which members are arranged by propinquity of descent. It is argued that the lineage population concept avoids the ambiguities associated with related biological and anthropological concepts and polysemes such as (...) population, ethnicity, and race. Other terms and concepts, such as form, cline, cluster, geographic population, breeding population, genetic population, breed, species, subspecies, ancestry, geographic ancestry, biogeographic ancestry, ancestral population, ancestry population, natural division, and population lineage, are discussed in relation to this concept. It is concluded that the lineage population concept is a useful analytic tool which describes, in line with the Kantian/Darwinian perspective, an interesting class of biological variation. (shrink)