Frank E. Zachos offers a comprehensive review of one of today's most important and contentious issues in biology: the species problem. After setting the stage with key background information on the topic, the book provides a brief history of species concepts from antiquity to the Modern Synthesis, followed by a discussion of the ontological status of species with a focus on the individuality thesis and potential means of reconciling it with other philosophical approaches. More than 30 (...) different species concepts found in the literature are presented in an annotated list, and the most important ones, including the Biological, Genetic, Evolutionary and different versions of the Phylogenetic Species Concept, are discussed in more detail. Specific questions addressed include the problem of asexual and prokaryotic species, intraspecific categories like subspecies and Evolutionarily Significant Units, and a potential solution to the species problem based on a hierarchical approach that distinguishes between ontological and operational species concepts. A full chapter is dedicated to the challenge of delimiting species by means of a discrete taxonomy in a continuous world of inherently fuzzy boundaries. Further, the book outlines the practical ramifications for ecology and evolutionary biology of how we define the species category, highlighting the danger of an apples and oranges problem if what we subsume under the same name ("species") is in actuality a variety of different entities. A succinct summary chapter, glossary and annotated list of references round out the coverage, making the book essential reading for all biologists looking for an accessible introduction to the historical, philosophical and practical dimensions of the species problem. (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)

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)

This chapter defends a pluralist understanding of species on which a normative species concept is viable and can support natural goodness evaluations. The central question here is thus: Since organisms are to be evaluated as members of their species, how does a proper understanding of species affect the feasibility of natural goodness evaluations? Philippa Foot has argued for a form of natural goodness evaluation in which living things are evaluated by how well fitted they are for (...) flourishing as members of their species, in ways characteristic of their species. She has further argued that assessments of moral goodness in humans are of the same evaluative form. However, natural goodness evaluations and, by extension, the natural goodness approach, do not garner justification in virtue of employing a scientifically privileged conception of species. The natural goodness approach is only justified given particular metaethical and normative commitments that are independent of naturalism, since the approach does not depend upon naturalism alone. (shrink)

The species concept is one of the oldest and most fundamental in biology. And yet it is almost universally conceded that no satisfactory definition of what constitutes a species has ever been proposed. The present article is devoted to an attempt to review the status of the problem from a methodological point of view. Since the species is one of the many taxonomic categories, the question of the nature of these categories in general needs to be entered (...) into. (shrink)

Students of later medieval semantics are familiar with the controversy that developed at the end of the thirteenth century over the signification of names. The debate focused on the signification of common nouns such as ‘man’ and ‘animal’: Do they signify an extramental thing or a mental representation of an extramental thing?Some authors at the end of the thirteenth century also discussed another question concerning what names signify, that is, whether they signify the composite of matter and form or only (...) the form of the composite. This second debate will not be considered in this article. Duns Scotus is commonly recognized as having played an important role in this debate.See Costantino Marmo, “Ontology and Semantics in the Logic of Duns Scotus,” in Umberto Eco and Costantino Marmo, ed., On Medieval Theory of Signs, (Amsterdam and Philadelphia: John Benjamins, 1989), pp. 161–63; and esp. Dominik Perler, “Duns Scotus on Signification,” Medieval Philosophy and Theology 3 (1993): 97–120. On the topic in general, see Paul Vincent Spade, “The Semantics of Terms,” in N. Kretzmann, A. Kenny, J. Pinborg, eds., The Cambridge History of Later Medieval Philosophy (Cambridge: Cambridge U.P., 1982), pp. 188–90; Jan Pinborg, “Bezeichnung in der Logik des XIII. Jahrhunderts,” in Albert Zimmermann, ed., Der Begriff der Repraesentatio im Mittelalter, Miscellanea Mediaevalia 8 (Berlin and New York: de Gruyter, 1971): 238–81; Elizabeth J. Ashworth, “Signification and Modes of Signifying in Thirteenth-Century Logic: A Preface to Aquinas on Analogy,” Medieval Philosophy and Theology 1 (1991): 43–53; Claude Panaccio, “From Mental Word to Mental Language,” Philosophical Topics 20 (1992): 125–47. In his Ordinatio, he alludes to a magna altercatio among his contemporaries concerning signification.John Duns Scotus, Ordinatio 1.27.1–3 n.83, in Commissio Scotistica, ed, Opera omnia 6 (Vatican City: Typis Polyglottis, 1963), p. 97. What is more, he gives, in his two commentaries on Aristotle’s Peri hermeneias, a detailed and fair analysis of the two contrasting positions on this issue.John Duns Scotus, Super Peri hermeneias 1.2, and Super Peri hermeneias 2.1, in Opera omnia 1 (Paris: L. Vivès, 1891), pp. 540–44, 582–85. Scotus’s logical commentaries are usually thought to have been composed before his theological writings, in the last decade of the thirteenth century. (shrink)

The nature of science is a primary goal in school science. Most teachers are not well-prepared for teaching NOS, but a sophisticated and in-depth understanding of NOS is necessary for effective teaching. Some authors emphasize the need for teaching NOS in context. Species, a central concept in biology, is proposed in this article as a concrete example of a means for achieving increased understanding of NOS. Although species are commonly presented in textbooks as fixed entities with a single (...) definition, the concept of species is a highly discussed one in the science and the philosophy of biology. A multitude of species concepts exist, reflecting both the views and interests of researchers and their utility in different organism groups. The present study serves to address the following questions: How do textbooks in Norwegian primary and lower secondary schools present the concept of species? Can inquiries into the concept of “species” serve to highlight aspects of NOS? A review of the available literature on species and species concepts in school is also performed. In the schoolbooks, the biological species concept is commonly used as the main definition, whereas the morphological species concept is represented by additional remarks of similarity. The potential and pitfalls of using the species concept for teaching NOS are discussed, with NOS being discussed both as a family resemblance concept and as a consensus list. Teacher education is proposed as a starting point for inducing a more sophisticated view of biology into schools. (shrink)

Species are the basis of the taxonomic scheme. They are the lowest taxonomic category that are used as units for describing biodiversity and evolution. In this contribution we discuss the current species concept for prokaryotes. Such organisms are considered to represent the widest diversity among living organisms. Species is currently circumscribed as follows: A prokaryotic species is a category that circumscribes a (preferably) genomically coherent group of individual isolates/strains sharing a high degree of similarity in (many) (...) independent features, comparatively tested under highly standardized conditions. Although the number of described prokaryotic species is underrepresented in the living world, this phylo-phenetic or polythetic species concept currently in use is considered to be pragmatic, operational and universally applicable and successfully used for identification processes. (shrink)

Species are the basis of the taxonomic scheme. They are the lowest taxonomic category that are used as units for describing biodiversity and evolution. In this contribution we discuss the current species concept for prokaryotes. Such organisms are considered to represent the widest diversity among living organisms. Species is currently circumscribed as follows: A prokaryotic species is a category that circumscribes a (preferably) genomically coherent group of individual isolates/strains sharing a high degree of similarity in (many) (...) independent features, comparatively tested under highly standardized conditions. Although the number of described prokaryotic species is underrepresented in the living world, this phylo-phenetic or polythetic species concept currently in use is considered to be pragmatic, operational and universally applicable and successfully used for identification processes. (shrink)

Summary The common view has been that Darwin regarded species as artificial and arbitrary constructions of taxonomists, not as distinct natural units. However, in his transmutation notebooks he clearly subscribed to the reality of species, on the basis of the criterion of non-interbreeding. A consequence of this biological species concept was his identification of the acquisition of reproductive isolation as the mark of the completion of speciation. He developed in the notebooks a theory of geographic speciation on (...) the grounds of (a) an empirical correlation between barriers and the geographical distributions of allied species, (b) the analogy of artificial isolation in the formation of new domesticated varieties, and (c) the logical consequence of blending inheritance. A brief discussion of Darwin's species concept in later writings concludes the paper. (shrink)

Integration (interaction among parts of an entity) is suggested to be necessary for individuality (contra, Metaphysics and the Origin of Species). A synchronic species is an integrated individual that can evolve as a unified whole; a diachronic lineage is a non-integrated historical entity that cannot evolve. Synchronic species and diachronic lineages are consequently suggested to be ontologically distinct entities, rather than alternative perspectives of the same underlying entity (contra Baum (1998), Syst. Biol. 47, 641–653; de Queiroz (1995), (...) Endless Forms: Species and Speciation, pp. 57–75; Genes, Categories and Species). Species concepts usually refer to either one or the other entity; for instance, the Biological Species Concept refers to synchronic species, whereas the Cladistic Species Concept refers to diachronic lineages. The debate over species concepts has often failed to recognise this distinction, resulting in invalid comparisons between definitions that attempt to delineate fundamentally different entities. (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 (...) class='Hi'>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)

The biological species (biospecies) concept applies only to sexually reproducing species, which means that until sexual reproduction evolved, there were no biospecies. On the universal tree of life, biospecies concepts therefore apply only to a relatively small number of clades, notably plants andanimals. I argue that it is useful to treat the various ways of being a species (species modes) as traits of clades. By extension from biospecies to the other concepts intended to capture (...) the natural realities of what keeps taxa distinct, we can treat other modes as traits also, and so come to understand that theplurality of species concepts reflects the biological realities of monophyletic groups.We should expect that specialists in different organisms will tend to favour those concepts that best represent the intrinsic mechanisms that keep taxa distinct in their clades. I will address the question whether modes ofreproduction such as asexual and sexual reproduction are natural classes, given that they are paraphyletic in most clades. (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)

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 debate over the ethics of radically, technologically altering the capacities and traditional form of the human body is rife with appeals to and dismissals of the importance of the integrity of the human species. Species-integrist arguments can be found in authors as varied as Annas, Fukuyama, Habermas, and Agar. However, the ethical salience of species integrity is widely contested by authors such as Buchanan, Daniels, Fenton, and Juengst. This article proposes a Phenomenological approach to the question (...) of species-integrity, arguing in favor of a phenomenon of species-recognition that carries an ethical pull. Building on Husserl’s Phenomenological account of empathy and the lived-body, as well as Schopenhauer’s concept of compassion as an ethical urphenomenon, I develop a “Phenomenological species concept” , which I argue has the ethical significance that biological species concepts do not. The PSC reorients the debate over human alteration and species integrity. (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)

The phylogenetic species concept is applied for the first time to a major radiation of birds, the birds-of-paradise (Paradisaeidae) of Australasia. Using the biological species concept, previous workers have postulated approximately 40–42 species in the family. Of these, approximately 13 are monotypic and 27 are polytypic with about 100 subspecies. Phylogenetic species are irreducible (basal) clusters of organisms (terminal taxa) that are diagnosably distinct from other such clusters. Within the context of this concept, approximately 90 (...) class='Hi'>species of paradisaeids are postulated to have diversified within Australasia. The phylogenetic species concept more accurately describes evolutionary diversity within the family and provides a better theoretical and empirical framework for analysing speciation, historical biogeography and patterns of morphological, behavioral and ecological diversification within this group than does the biological species concept. (shrink)

RESUMEN: En el actual contexto científico que forma la concepción darwiniana de las especies aún persisten las interpretaciones esencialistas de los conceptos de especie. ¿Se trata aquí sólo de la ignorancia de la teoría biológica? O, más bien, ¿es posible comprender la persistencia de los enfoques esencialistas sobre la base de la potencialidad de estos enfoques para explicar el logro de ciertos valores epistémicos de los actuales conceptos de especie? Me propongo responder afirmativamente a esta última pregunta. En la sección (...) 1 argumento que Samir Okasha (2002) no logra mostrar que hay otras razones, distintas a la de la ignorancia de la biología, que motivan el error de Kripke y de Putnam acerca de las especies. En la sección 2 propongo mi respuesta a la pregunta inicial en términos de algunos valores epistémicos que comparten las ideas esencialistas acerca de las especies y algunos de los actuales conceptos de especie.ABSTRACT: In the current scientific context that forms Darwinian conception of species, essentialist interpretations of the species concept persist. Is this only because of an ignorance of the biological theory? Or rather, is it possible to understand the persistence of essentialist approaches on the basis of the potentiality of these approaches to account for the achievement of certain epistemic values of the current species concepts? I intend to give a positive answer to this second question. In Section 1, I argue that Samir Okasha (2002) does not succeed in demonstrating that there are other reasons, which are different from that one regarding the ignorance about biology, which cause Kripke and Putnam to be mistaken about the species. In Section 2, I put forward my answer to the initial question in terms of some epistemic values sharing the essentialist ideas about the species and some of the current species concepts. (shrink)

The conceptual dilemma that species entail has divided, since its formulation, biologists and philosophers in two spheres: those who believe in the existence of a unified category of species and those who defend the unyielding plurality of equally legitimate concepts. The aim of this paper is to comprise the analysis of the problems that revolve around the species category with the only purpose being to determine the existence of only one univocal and unrestricted definition of (...) class='Hi'>species. For this reason, the paper will be divided into two sections. The first section will analyse the extent to which essentialism amounts to an antithetical theory to the modern biological theory. In the second section a detailed critique will be carried out on existing attempts to devise a definition of species. Two conclusions can be drawn from the previous statements. First and due to the fall of essentialism, that there is not only one single category of species but an uncompromising plurality of concepts. Secondly and following previous assertion, it can be stated that the most consistent viewpoint in the evolutionary theory is the one in which an ontological pluralism is embraced and, consequently, a taxonomical pluralism. (shrink)

The conceptual dilemma that species entail has divided, since its formulation, biologists and philosophers in two spheres: those who believe in the existence of a unified category of species and those who defend the unyielding plurality of equally legitimate concepts. The aim of this paper is to comprise the analysis of the problems that revolve around the species category with the only purpose being to determine the existence of only one univocal and unrestricted definition of (...) class='Hi'>species. For this reason, the paper will be divided into two sections. The first section will analyse the extent to which essentialism amounts to an antithetical theory to the modern biological theory. In the second section a detailed critique will be carried out on existing attempts to devise a definition of species. Two conclusions can be drawn from the previous statements. First and due to the fall of essentialism, that there is not only one single category of species but an uncompromising plurality of concepts. Secondly and following previous assertion, it can be stated that the most consistent viewpoint in the evolutionary theory is the one in which an ontological pluralism is embraced and, consequently, a taxonomical pluralism. (shrink)

The so-called ‘‘species problem’’ has plagued evolution- ary biology since before Darwin’s publication of the aptly titled Origin of Species. Many biologists think the problem is just a matter of semantics; others complain that it will not be solved until we have more empirical data. Yet, we don’t seem to be able to escape discussing it and teaching seminars about it. In this paper, I briefly examine the main themes of the biological and philosophical liter- atures on the (...) species problem, focusing on identifying common threads as well as relevant differences. I then argue two fundamental points. First, the species problem is not primarily an empirical one, but it is rather fraught with philosophical questions that require—but cannot be settled by—empirical evidence. Second, the (dis-)solution lies in explicitly adopting Wittgenstein’s idea of ‘‘family resemblance’’ or cluster concepts, and to consider spe- cies as an example of such concepts. This solution has several attractive features, including bringing together apparently diverging themes of discussion among bio- logists and philosophers. The current proposal is con- ceptually independent (though not incompatible) with the pluralist approach to the species problem advocated by Mishler, Donoghue, Kitcher and Dupre ́, which implies that distinct aspects of the species question need to be emphasized depending on the goals of the researcher. From the biological literature, the concept of species that most closely matches the philosophical discussion pre- sented here is Templeton’s cohesion idea. (shrink)

RESUMEN: La biología evolucionista no ha logrado definir un concepto de especie que satisfaga a todos sus colaboradores. El presente panorama crítico de las principales propuestas y sus respectivas dificultades apunta, por un lado, a ilustrar los procesos de formación de conceptos en las ciencias empíricas y, por otro, a socavar la visión parateológica del conocimiento y la verdad que inspiró inicialmente a la ciencia moderna y prevalece aún entre muchas personas educadas. El artículo se divide en dos partes. La (...) primera atiende al concepto biológico (o genético) de especie adoptado por Theodosius Dobzhansky y Ernst Mayr alrededor de 1940, así como a las alternativas introducidas para superar sus limitaciones. La segunda parte estudia la tradición “cladista” fundada por Willi Hennig (1950) y sus ramificaciones. Varios conceptos de especie que no era fácil integrar en estos dos grupos se omitieron en aras de la coherencia y la brevedad de la exposición.ABSTRACT: Evolutionary biology has not suceeded in defining a concept of species that will satisfy all researchers. This critical survey of the main proposals and their respective difficulties tends, on the one hand, to throw light on the processes of concept formation in the empirical sciences, and, on the other, to undermine the paratheological vision of knowledge and truth that initially inspired modern science and still prevails among many educated persons. The article is divided into two parts. The first part concerns the biological (or genetical) concept of species which was adopted by Theodosius Dobzhansky and Ernst Mayr ca. 1940 and some alternatives which were subsequently introduced to overcome its limitations. The second part deals with several branches of the cladist tradition founded by Willi Hennig (1950). Various concepts of species that could not be easily integrated in either group were omitted for the sake of coherence and brevity. (shrink)

Species concepts for bacteria and other microbes are contentious, because they are often asexual. There is a Problem of Homogeneity: every mutation in an asexual lineage forms a new strain, of which all descendents are clones until a new mutation occurs. We should expect that asexual organisms would form a smear or continuum. What causes the internal homogeneity of asexual lineages, if they are in fact homogeneous? Is there a natural “species concept” for “microbes”? Two main (...) class='Hi'>concepts devised for metazoans and metaphytes have been applied to bacteria. One is the Recombination Concept, a revised form of the Biological Species Concept in which the homogenizing mechanism is the sharing of genome fragments, somewhat akin to sexual recombination. The other is the Ecological Species Concept, in which the ecological niche is that which maintains lineages as cohesive. In this paper I will discuss these two concepts, and offer an underlying model that conjoins them, and consider the implications for species concepts in general. In short, my argument is that asexual species are instances of the most primitive and underived notion of species, which I will call “quasispecies”, following Eigen, and that sexual species are merely one derived kind of species. Moreover, I will argue that there is a continuum of recombination from simple viral models in which each strain is a clone, through to obligate recombination of 50% of the parents’ genome, and that consequently there is no sharp division between “microbial” and more familiar species. (shrink)