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)

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)

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)

Patricia Williams made a number of claims concerning the methods and practise of cladistic analysis and classification. Her argument rests upon the distinction of two kinds of hierarchy: a divisional hierarchy depicting evolutionary descent and the Linnean hierarchy describing taxonomic groups in a classification. Williams goes on to outline five problems with cladistics that lead her to the conclusion that systematists should eliminate cladism as a school of biological taxonomy and to replace it either with something that is philosophically coherent (...) or to replace it with pure methodology, untainted by theory (Williams 1992, 151). Williams makes a number of points which she feels collectively add up to insurmountable problems for cladistics. We examine Williams' views concerning the two hierarchies and consider what cladists currently understand about the status of ancestors. We will demonstrate that Williams has seriously misunderstood many modern commentators on this subject and all of her five persistent problems are derivable from this misunderstanding. Some persons believe and argue, on grounds approaching faith it seems to me, that phylogeny comes from our knowledge of evolution. Others have found to their surprise, and sometimes dismay, that phylogeny comes from our knowledge of systematics. Nelson (1989, 67). (shrink)

In Phylogenetic Systematics (1966), Willi Hennig conflates the Linnaean hierarchy with what Hennig refers to as the divisional hierarchy. In doing so, he lays the foundations of that school of biological taxonomy known as cladism on a philosophically ambiguous basis. This paper compares and contrasts the two hierarchies and demonstrates that Hennig conflates them. It shows that Hennig's followers also conflate them. Finally, it illuminates five persistent problems in cladism by suggesting that they arise from Hennig's original confusion.

Industrial melanism, according to the traditional explanation, amounts to niche construction since it involves changes in predation pressure. Indeed, it would be difficult to imagine selection without niche construction. This cannot be what Laland, Odling-Smee & Feldman mean. They offer convincing examples, but they should provide a better definition of “niche construction” to indicate how their view supplements traditional evolutionary biology.

John Beatty (1995) and Alexander Rosenberg (1994) have argued against the claim that there are laws in biology. Beatty's main reason is that evolution is a process full of contingency, but he also takes the existence of relative significance controversies in biology and the popularity of pluralistic approaches to a variety of evolutionary questions to be evidence for biology's lawlessness. Rosenberg's main argument appeals to the idea that biological properties supervene on large numbers of physical properties, but he also develops (...) case studies of biological controversies to defend his thesis that biology is best understood as an instrumental discipline. The present paper assesses their arguments. (shrink)

Over the past decade, it has been discovered that disparate aspects of morphology – often of distantly related groups of organisms – are regulated by the same genetic regulatory mechanisms. Those discoveries provide a new perspective on morphological evolutionary change. A conceptual framework for exploring these research findings is termed ‘deep homology’. A comparative framework for morphological relations of homology is provided that distinguishes analogy, homoplasy, plesiomorphy and synapomorphy. Four examples – three from plants and one from animals – demonstrate (...) that homologous developmental mechanisms can regulate a range of morphological relations including analogy, homoplasy and examples of uncertain homology. Deep homology is part of a much wider range of phenomena in which biological (genes, regulatory mechanisms, morphological traits) and phylogenetic levels of homology can both be disassociated. Therefore, to understand homology, precise, comparative, independent statements of both biological and phylogenetic levels of homology are necessary. (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.

The correct explanation of why species, in evolutionary theory, are individuals and not classes is the cladistic species concept. The cladistic species concept defines species as the group of organisms between two speciation events, or between one speciation event and one extinction event, or (for living species) that are descended from a speciation event. It is a theoretical concept, and therefore has the virtue of distinguishing clearly the theoretical nature of species from the practical criteria by which species may be (...) recognized at any one time. Ecological or biological (reproductive) criteria may help in the practical recognition of species. Ecological and biological species concepts are also needed to explain why cladistic species exist as distinct lineages, and to explain what exactly takes place during a speciation event. The ecological and biological species concepts work only as sub-theories of the cladistic species concept and if taken by themselves independently of cladism they are liable to blunder. The biological species concept neither provides a better explanation of species indivudualism than the ecological species concept, nor, taken by itself, can the biological species concept even be reconciled with species individualism. Taking the individuality of species seriously requires subordinating the biological, to the cladistic, species concept. (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)

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)

The metaphysical nature of homologies has been variously characterized as natural kind, individualist, and pluralist-pragmatic. In this essay, I aim to build on the work of proponents of a natural kinds ontology for homologies using Richard Boyd’s influential HPC account of natural kinds. I aim to advance this position by showing the unique fit of extending the HPC account to homologies, deflecting individualist critiques, as well as the pluralist-pragmatic alternative, showing that homologies have a determinate metaphysical character as kinds. As (...) an important extension of this position, I attempt to explain away how the mistaken metaphysics of the individualist, and derivatively the pluralist-pragmatic approach that contextually embraces it, can facilitate certain elements of biological practice. (shrink)

"The Natural System" is the abstract notion of the order in living diversity. The richness and complexity of this notion is revealed by the diversity of representations of the Natural System drawn by ornithologists in the Nineteenth Century. These representations varied in overall form from stars, to circles, to maps, to evolutionary trees and cross-sections through trees. They differed in their depiction of affinity, analogy, continuity, directionality, symmetry, reticulation and branching, evolution, and morphological convergence and divergence. Some representations were two-dimensional, (...) and some were three-dimensional; n-dimensional representations were discussed but never illustrated. The study of diagrammatic representations of the Natural System is made difficult by the frequent failure of authors to discuss them in their texts, and by the consequent problem of distinguishing features which carried meaning from arbitrary features and printing conventions which did not. Many of the systematics controversies of the last thirty years have their roots in the conceptual problems which surrounded the Natural System in the late 1800s, problems which were left unresolved when interest in higher-level systematics declined at the turn of this century. (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)

Many of the current comparisons of taxic phylogenetic and biological homology in the context of morphology focus on what are seen as categorical distinctions between the two concepts. The first, it is claimed, identifies historical patterns of conservation and variation relating taxa; the second provides a causal framework for the explanation of this conservation and variation. This leads to the conclusion that the two need not be placed in conflict and are in fact compatible, having non-competing epistemic purposes or mapping (...) the same extensions in the form of monophyletic groupings (see Roth, The biological basis of homology 1–26, 1988; Sluys, J Zool Syst Evol Res 34:145–152, 1996; Abouheif, Trends Ecol Evol 12:405–408, 1997; Brigandt, J Exp Zool 299:9–17, 2003, Biol Philos 22:709–725, 2007; Assis and Brigandt, Evol Biol 36:248–255, 2009). This article argues that moves in this direction miss the essential disagreement between these concepts as they have been developed in the context of the debate concerning the best concept for evolutionary investigation. We should rather see these concepts employing a common fundamental methodological approach to homology, but disagreeing about how to apply the methodology effectively. Both concepts employ class reasoning, which pursues homologies as units of generalization—more precisely, as sources of reliable and relevant group-bound information in the form of shared underlying causes. The dispute can be better understood by two poles that structure such reasoning: the need for a reliable basis for projections about the causal history of shared structures, and the desire to identify homologous characters with more informative and specific causal information relevant to generalizing about evolutionary processes. Judgments in favor of one or the other in turn have affected the scope or extension of these competing homology concepts. (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.

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)

Karl Popper has been one of the few philosophers of sciences who has influenced scientists. I evaluate Popper's influence on our understanding of evolutionary theory from his earliest publications to the present. Popper concluded that three sorts of statements in evolutionary biology are not genuine laws of nature. I take him to be right on this score. Popper's later distinction between evolutionary theory as a metaphysical research program and as a scientific theory led more than one scientist to misunderstand his (...) position on evolutionary theory as a scientific theory. In his later work Popper also introduced what he took to be improvements of evolutionary theory. Thus far these improvements have had almost no influence on evolutionary biology. I conclude by examining the influence of Popper on the reception of cladistic analysis. (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)

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.

In my dissertation I define historical disciplines as disciplines that aim to give a historical interpretation of the evidence. Phylogenetic systematics is a historical discipline and therefore in my definition phylogenies should be thought of as historical interpretations of relationships between taxa.

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)

A number of areas of biology raise questions about what is of value in the natural environment and how we ought to behave towards it: conservation biology, environmental science, and ecology, to name a few. Based on my experience teaching students from these and similar majors, I argue that the field of environmental ethics has much to teach these students. They come to me with pent-up questions and a feeling that more is needed to fully engage in their subjects, and (...) I believe some exposure to environmental ethics can help focus their interests and goals. I identify three primary areas in which environmental ethics can con- tribute to their education. The first is an examination of who (or what) should be considered to be part of our moral community (i.e., the community to whom we owe direct duties). Is it humans only? Or does it include all sentient life? Or all life? Or ecosystems considered holistically? Often, readings implicitly assume one or more of these answers; the goal is to make the student more sensitive to these implicit claims and to get them to think about the different reasons that support them. The second area, related to the first, is the application of the different answers concerning the extent of the ethical community to real environmental issues and problems. Students need to be aware of how the different answers concerning the moral community can imply conflicting answers for how we should act in certain cases and to think about ways to move toward conflict resolution. The third area in which environmental ethics can contribute is a more conceptual one, focusing on central concepts such as biodiversity, sustainability, species, and ecosystems. Exploring and evaluating various meanings of these terms will make students more reflective and thoughtful citizens and biologists, sensitive to the implications that different conceptual choices make. (shrink)