Argues for the primacy of the phylogenetic system as the general reference system in biology. This book, first published in 1966, generated significant controversy and opened possibilities for evolutionary biology.

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

Systematics based solely on structuralist principles is non-science because it is derived from first principles that are inconsistent in dealing with both synchronic and diachronic aspects of evolution, and its evolutionary models involve hidden causes, and unnameable and unobservable entities. Structuralist phylogenetics emulates axiomatic mathematics through emphasis on deduction, and “hypotheses” and “mapped trait changes” that are actually lemmas and theorems. Sister-group-only evolutionary trees have no caulistic element of scientific realism. This results in a degenerate systematics based on (...) patterns of fact or evidence being treated as so fundamental that all other data may be mapped to the cladogram, resulting in an apparently well-supported classification that is devoid of evolutionary theory. Structuralism in systematics is based on a non-ultrametric analysis of sister-group informative data that cannot detect or model a named taxon giving rise to a named taxon, resulting in classifications that do not reflect macroevolutionary changes unless they are sister lineages. Conservation efforts are negatively affected through epistemological extinction of scientific names. Evolutionary systematics is a viable alternative, involving both deduction and induction, hypothesis and theory, developing trees with both synchronic and diachronic dimensions often inferring nameable ancestral taxa, and resulting in classifications that advance evolutionary theory and explanations for particular groups. (shrink)

The long-awaited revision of the industry standard on phylogenetics Since the publication of the first edition of this landmark volume more than twenty-five years ago, phylogenetic systematics has taken its place as the dominant paradigm of systematic biology. It has profoundly influenced the way scientists study evolution, and has seen many theoretical and technical advances as the field has continued to grow. It goes almost without saying that the next twenty-five years of phylogenetic research will prove as (...) fascinating as the first, with many exciting developments yet to come. This new edition of Phylogenetics captures the very essence of this rapidly evolving discipline. Written for the practicing systematist and phylogeneticist, it addresses both the philosophical and technical issues of the field, as well as surveys general practices in taxonomy. Major sections of the book deal with the nature of species and higher taxa, homology and characters, trees and tree graphs, and biogeography—the purpose being to develop biologically relevant species, character, tree, and biogeographic concepts that can be applied fruitfully to phylogenetics. The book then turns its focus to phylogenetic trees, including an in-depth guide to tree-building algorithms. Additional coverage includes: Parsimony and parsimony analysis Parametric phylogenetics including maximum likelihood and Bayesian approaches Phylogenetic classification Critiques of evolutionary taxonomy, phenetics, and transformed cladistics Specimen selection, field collecting, and curating Systematic publication and the rules of nomenclature Providing a thorough synthesis of the field, this important update to Phylogenetics is essential for students and researchers in the areas of evolutionary biology, molecular evolution, genetics and evolutionary genetics, paleontology, physical anthropology, and zoology. (shrink)

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

The question of whether or not to partition data for the purposes of inferring phylogenetic hypotheses remains controversial. Opinions have been especially divided since Kluge's (1989, Systematic Zoology 38, 7–25) claim that data partitioning violates the requirement of total evidence (RTE). Unfortunately, advocacy for or against the RTE has not been based on accurate portrayals of the requirement. The RTE is a basic maxim for non-deductive inference, stipulating that evidence must be considered if it has relevance to an inference. (...) Evidence is relevant if it has a positive or negative effect on a given conclusion. In the case of ℈partitioned’ phylogenetic inferences, the RTE is violated, and the basis for rational belief in any conclusion is compromised, unless it is shown that the partitions are evidentially irrelevant to one another. The goal of phylogenetic systematics is to hypothesize past causal conditions to account for observed shared similarities among two or more species. Such inferences are non-deductive, necessitating consideration of the RTE. Some phylogeneticists claim the parsimony criterion as justification for the RTE. There is no relation between the two – parsimony is a relation between a hypothesis and causal question(s). Parsimony does not dictate the content of premises prior to an inference. ℈Taxonomic congruence,’ ℈supertrees,’ and ℈conditional combination’ methods violate the RTE. Taxonomic congruence and supertree methods also fail to achieve the intended goal of phylogenetic inference, such that ℈consensus trees’ and ℈supertrees’ lack an empirical basis. ℈Conditional combination’ is problematic because hypotheses derived from partitioned data cannot be compared – a causal hypothesis inferred to account for a set of effects only has relevance to those effects, not any comparative relevance to other causal hypotheses. A similar problem arises in the comparisons of hypotheses derived from different causal theories. (shrink)

I attempt to raise questions regarding elements of systematics—primarily in the realm of phylogenetic reconstruction—in order to provoke discussion on the current state of affairs in this discipline, and also evolutionary biology in general: e.g., conceptions of homology and homoplasy, hypothesis testing, the nature of and objections to Hennigian “phylogenetic systematics”, and the schism between Darwinian descendants of the “modern evolutionary synthesis” and their supposed antagonists, cladists and punctuationalists.

I respond to the bad lot argument in the context of biological systematics. The response relies on the historical nature of biological systematics and on the availability of pattern explanations. The basic assumption of common descent enables systematic methodology to naturally generate candidate explanatory hypotheses. However, systematists face a related challenge in the issue of character analysis. Character analysis is the central problem for contemporary systematics, yet the general problem of which it is a case—what counts as (...) evidence?—has not been adequately discussed by proponents of inference to the best explanation. Facing this problem is the price of adopting abductive methods. I sketch an account of how systematists approach the problem of evidence. (shrink)

Taxonomies of living things and the methods used to produce them changed little with the institutionalization of evolutionary thinking in biology. Instead, the relationships expressed in existing taxonomies were merely reinterpreted as the result of evolution, and evolutionary concepts were developed to justify existing methods. I argue that the delay of the Darwinian Revolution in biological taxonomy has resulted partly from a failure to distinguish between two fundamentally different ways of ordering identified by Griffiths : classification and systematization. Classification consists (...) of ordering entities into classes, groups defined by the attributes of their members; in contrast, systematization consists of ordering entities into systems, more inclusive entities whose existence depends on some natural process through which their parts are related. Evolutionary, or phylogenetic, systematics takes evolutionary descent to be the natural process of interest in biological taxonomy. I outline a general framework for a truly phylogenetic systematics and examine some of its consequences. (shrink)

I address David Hull's theses about the process of science from the perspective of an evolutionary biologist, particularly emphasizing phylogenetic systematics, an area that has figured prominently in Hull's work as a source of both sociological data and metatheory. The goal is to carefully explore analogies and disanalogies between scientific process and comparative biology. There do seem to be remarkable analogies, indeed these lead to important insights that might not otherwise have been made, yet some possible analogies present (...) novel problems: Are "memes" like genes or like traits? What is the nature of replication in science? It is argued that the primary need is for some rigorously worked-out case studies. (shrink)

Although Bayesian methods are widely used in phylogenetic systematics today, the foundations of this methodology are still debated among both biologists and philosophers. The Bayesian approach to phylogenetic inference requires the assignment of prior probabilities to phylogenetic trees. As in other applications of Bayesian epistemology, the question of whether there is an objective way to assign these prior probabilities is a contested issue. This paper discusses the strategy of constraining the prior probabilities of phylogenetic trees (...) by means of the Principal Principle. In particular, I discuss a proposal due to Velasco (Biol Philos 23:455–473, 2008) of assigning prior probabilities to tree topologies based on the Yule process. By invoking the Principal Principle I argue that prior probabilities of tree topologies should rather be assigned a weighted mixture of probability distributions based on Pinelis’ (P Roy Soc Lond B Bio 270:1425–1431, 2003) multi-rate branching process including both the Yule distribution and the uniform distribution. However, I argue that this solves the problem of the priors of phylogenetic trees only in a weak form. (shrink)

This chapter contains sections titled: Introduction From Art to Science: An Introduction to Schools of Thought How to Infer Phylogeny, Or, Why Some Cladists Aren't “Cladists” Summary and Synthesis Acknowledgment References.

Framing systematics as a field consistent with scientific inquiry entails that inferences of phylogenetic hypotheses have the goal of producing accounts of past causal events that explain differentially shared characters among organisms. Linking observations of characters to inferences occurs by way of why-questions implied by data matrices. Because of their form, why-questions require the use of common-cause theories. Such theories in phylogenetic inferences include natural selection and genetic drift. Selection or drift can explain ‘morphological’ characters but selection (...) cannot be causally applied to sequences since fitness differences cannot be directly associated with individual nucleotides or amino acids. The relation of selection to sequence data is by way of downward or top-down causation from those phenotypes upon which selection occurs. The application of phylogenetic inference to explain sequence data is thus restricted to instances where drift is the relevant theory; those nucleotides or amino acids that can be explained via downward causation are precluded from inclusion in the data matrix. The restrictions on the inclusion of sequence data in phylogenetic inferences equally apply to species hypotheses, precluding the more restrictive approach known as DNA barcoding. Not being able to discern drift and selection as relevant causal mechanisms can severely constrain the inclusion and explanations of sequence data. Implications of such exclusion are discussed in relation to the requirement of total evidence. (shrink)

Two new modes of thinking have spread through systematics in the twentieth century. Both have deep historical roots, but they have been widely accepted only during this century. Population thinking overtook the field in the early part of the century, culminating in the full development of population systematics in the 1930s and 1940s, and the subsequent growth of the entire field of population biology. Population thinking rejects the idea that each species has a natural type (as the earlier (...) essentialist view had assumed), and instead sees every species as a varying population of interbreeding individuals. Tree thinking has spread through the field since the 1960s with the development of phylogenetic systematics. Tree thinking recognizes that species are not independent replicates within a class (as earlier group thinkers had tended to see them), but are instead interconnected parts of an evolutionary tree. It lays emphasis on the explanation of evolutionary events in the context of a tree, rather than on the states exhibited by collections of species, and it sees evolutionary history as a story of divergence rather than a story of development. Just as population thinking gave rise to the new field of population biology, so tree thinking is giving rise to the new field of phylogenetic biology. (shrink)

The author proposes a system of identification, positional, and phyletic numbers for taxa that makes possible a significant relationship between numerical classification and phylogeny.

Ontological questions in biology have typically focused on the nature of species: what are species; how are they identified and individuated? There is an analogous, but much neglected concern: what are characters; how are they identified and individuated? Character individuation is significant because biological systematics relies on a parsimony principle to determine phylogeny and classify taxa, and the parsimony principle is usually interpreted to favor the phylogenetic hypothesis that requires the fewest changes in characters. But no character individuation (...) principle identified so far is adequate. For biological systematics we need a better way of conceiving characters. (shrink)

The reduced costs of DNA sequencing and the use of such data for HIV‐1 clinical management and phylogenetic analysis have led to a massive increase of HIV‐1 sequences in the last few years. Phylogenetic analysis has shed light on the origin, spread and characteristics of HIV‐1 epidemics and outbreaks. Phylogenetic analysis is now also being used to advance our knowledge of the drivers of HIV‐1 transmission in order to design effective interventions. However, HIV phylogenetic analysis presents (...) unique ethical challenges, which have not been fully explored. This review presents an analysis of what appear to be key ethical issues in HIV phylogenetics in the hope of stimulating further conceptual and empirical work in this rapidly emerging area. We structure the review using the Emanuel Framework, a systematic, holistic framework, which has been adapted for use in developing countries, which bear the brunt of the HIV‐1 pandemic. (shrink)

While Bayesian methods have become very popular in phylogenetic systematics, the foundations of this approach remain controversial. The star-tree paradox in Bayesian phylogenetics refers to the phenomenon that a particular binary phylogenetic tree sometimes has a very high posterior probability even though a star tree generates the data. I argue that this phenomenon reveals an unattractive feature of the Bayesian approach to scientific inference and discuss two proposals for how to address the star-tree paradox. In particular, I (...) defend the polytomy prior as a solution of the paradox and argue that it is preferable to a data-size dependent branch lengths prior from a methodological perspective. However, while this reply dissolves the star-tree paradox, the general challenge to Bayesian confirmation theory remains unmet. (shrink)

Deduction leads to causal explanation in phylogenetic inference when the evidence, the systematic character, is conceptualized as a transformation series. Also, the deductive entailment of modus tollens is satisfied when those kinds of events are operationalized as patristic difference. Arguments to the contrary are based largely on the premise that character-states are defined intensionally as objects, in terms of similarity relations. However, such relations leave biologists without epistemological access to the causal explanation and explanatory power of historical statements. Moreover, (...) the prediction-making to which those kinds of relations are limited in practice can lead to a category error—the mental conversion of an abstraction into a thing . The latter practices and problems characterize pattern cladistics, taxa being interpreted as homeostatic property cluster natural kinds, and other instrumentalist research programs. (shrink)

138 titles across a wide range of scholarly publications illustrate the conceptual affinities that connect the palaetiological sciences of biological systematics, historical linguistics, and stemmatics. These three fields all have as their central objective the reconstruction of evolutionary "trees of history" that depict phylogenetic patterns of descent with modification among species, languages, and manuscripts. All three fields flourished in the nineteenth century, underwent parallel periods of quiescence in the early twentieth century, and in recent decades have seen widespread (...) parallel revivals. "Tree thinking" (O'Hara, 1988) is now standard within evolutionary biology, and the unity of what William Whewell called palaetiology is being once again appreciated by scholars and scientists in many fields. (shrink)

This paper explores the use of Popper's philosophy of science by cladists in their battle against evolutionary and numerical taxonomy. Three schools of biological systematics fiercely debated each other from the late 1960s: evolutionary taxonomy, phenetics or numerical taxonomy, and phylogenetic systematics or cladistics. The outcome of that debate was the victory of phylogenetic systematics/cladistics over the competing schools of thought. To bring about this "cladistic turn" in systematics, the cladists drew heavily on the (...) philosopher K.R. Popper in order to dress up phylogenetic systematics as a hypothetico-deductivist, indeed falsificationist, research program that would put an end to authoritarianism. As the case of the "cladistic revolution" demonstrates, scientists who turn to philosophy in defense of a research program read philosophers with an agenda in mind. That agenda is likely to distort the philosophical picture, as happened to Popper's philosophy of science at the hands of cladists. (shrink)

Advocates of cladistic parsimony methods have invoked the philosophy of Karl Popper in an attempt to argue for the superiority of those methods over phylogenetic methods based on Ronald Fisher's statistical principle of likelihood. We argue that the concept of likelihood in general, and its application to problems of phylogenetic inference in particular, are highly compatible with Popper's philosophy. Examination of Popper's writings reveals that his concept of corroboration is, in fact, based on likelihood. Moreover, because probabilistic assumptions (...) are necessary for calculating the probabilities that define Popper's corroboration, likelihood methods of phylogenetic inference—with their explicit probabilistic basis—are easily reconciled with his concept. In contrast, cladistic parsimony methods, at least as described by certain advocates of those methods, are less easily reconciled with Popper's concept of corroboration. If those methods are interpreted as lacking probabilistic assumptions, then they are incompatible with corroboration. Conversely, if parsimony methods are to be considered compatible with corroboration, then they must be interpreted as carrying implicit probabilistic assumptions. Thus, the non-probabilistic interpretation of cladistic parsimony favored by some advocates of those methods is contradicted by an attempt by the same authors to justify parsimony methods in terms of Popper's concept of corroboration. In addition to being compatible with Popperian corroboration, the likelihood approach to phylogenetic inference permits researchers to test the assumptions of their analytical methods (models) in a way that is consistent with Popper's ideas about the provisional nature of background knowledge. (shrink)

This chapter explores the idea that phylogenetic diversity plays a unique role in underpinning conservation endeavour. The conservation of biodiversity is suffering from a rapid, unguided proliferation of metrics. Confusion is caused by the wide variety of contexts in which we make use of the idea of biodiversity. Characterisations of biodiversity range from all-variety-at-all-levels down to variety with respect to single variables relevant to very specific conservation contexts. Accepting biodiversity as the sum of a large number of individual measures (...) results in an empirically intractable framework. However, large-scale decisions cannot be based on biodiversity variables inferred from local conservation imperatives because the variables relevant to the many systems being compared would be incommensurate with one another. We therefore need some general conception of biodiversity that would make tractable such large-scale environmental decision-marking. We categorise the large array of strategies for the measurement of biodiversity into four broad groups for consideration as general measures of biodiversity. We compare common moral justifications for the conservation of biodiversity and conclude that some form of instrumental value is the most plausible justification for biodiversity conservation. Although this is often interpreted as a reliance on option value, we opt for a broadly consequentialist characterisation of biodiversity conservation. We conclude that the best justified general measure of biodiversity will be some form of phylogenetic diversity. (shrink)

Phylogenetic trees are meant to represent the genealogical history of life and apparently derive their justification from the existence of the tree of life and the fact that evolutionary processes are treelike. However, there are a number of problems for these assumptions. Here it is argued that once we understand the important role that phylogenetic trees play as models that contain idealizations, we can accept these criticisms and deny the reality of the tree while justifying the continued use (...) of trees in phylogenetic theory and preserving nearly all of what defenders of trees have called the “importance of tree thinking.”. (shrink)

Tool use research has suffered from a lack of consistent theoretical frameworks. There is a plethora of tool use definitions and the most widespread ones are so inclusive that the behaviors that fall under them arguably do not have much in common. The situation is aggravated by the prevalence of anecdotes, which have played an undue role in the literature. In order to provide a more rigorous foundation for research and to advance our understanding of the interrelation between tool use (...) and cognition, we suggest the adoption of Fragaszy and Mangalam's (2018) tooling framework, which is characterized by the creation of a body-plus-object system that manages a mechanical interface between tool and surface. Tooling is limited to a narrower suite of behaviors than tool use, which might facilitate its neurocognitive investigation. Indeed, evidence in the literature indicates that tooling has distinct neurocognitive underpinnings not shared by other activities typically classified as tool use, at least in primates. In order to understand the extent of tooling incidences in previous research, we systematically surveyed the comprehensive tool use catalog by Shumaker et al. (2011). We identified 201 tool use submodes, of which only 81 could be classified as tooling, and the majority of the tool use examples across species were poorly supported by evidence. Furthermore, tooling appears to be phylogenetically less widespread than tool use, with the greatest variability found in the primate order. However, in order to confirm these findings and to understand the evolution and neurocognitive mechanisms of tooling, more systematic research will be required in the future, particularly with currently underrepresented taxa. (shrink)

Edvard August Vainio was a world-renowned Finnish lichenologist. In Finland, however, he was a controversial person due to his strong pro-Finnish political views. Equally disputed was his opinion that systematics should be based on evolutionary theory and phylogenetic thinking. Vainio was familiar with the ideas of the early German phylogeneticists—especially those of Carl Wilhelm von Nägeli – and, applying them, aimed to create an exact method for building a natural classification of lichens already at the end of the (...) nineteenth century. In this respect, Vainio was a true pioneer, as no actual phylogenetic method had yet been developed. In the general spirit of the time, Vainio focused on finding the ancestors of species and other taxa by comparing primitive and derived features of homologous characters. However, Vainio already understood the concept of sister groups in 1880, the identification of which is the basis of all modern phylogenetic research. Nevertheless, the distinctive method developed by Vainio was not so much focused on the construction of a phylogenetic tree, but on revealing the origin of species through the differentiation and fixation of their polymorphic variation. Indeed, Vainio’s species concept is surprisingly similar to the phylogenetic species concepts presented a hundred years later. Although in many ways progressive, Vainio’s views did not influence the development of phylogenetics more widely, but his discussions are nevertheless a valuable source to understanding the early development of phylogenetic theory. (shrink)

Tree diagrams are the prevailing form of visualization in biological classification and phylogenetics. Already during the time of the so-called Systematist Wars from the mid-1960s until the 1980s most journal articles and textbooks published by systematists contained tree diagrams. Although this episode of systematics is well studied by historians and philosophers of biology, most analyses prioritize scientific theories over practices and tend to emphasize conflicting theoretical assumptions. In this article, I offer an alternative perspective by viewing the conflict through (...) the lens of representational practices with a case study on tree diagrams that were used by numerical taxonomists (phenograms) and cladists (cladograms). I argue that the current state of molecular phylogenetics should not be interpreted as the result of a competition of views within systematics. Instead, molecular phylogenetics arose independently of systematics and elements of cladistics and phenetics were integrated into the framework of molecular phylogenetics, facilitated by the compatibility of phenetic and cladistic practices with the quantitative approach of molecular phylogenetics. My study suggests that this episode of scientific change is more complex than common narratives of battles and winners or conflicts and compromises. Today, cladograms are still used and interpreted as specific types of molecular phylogenetic trees. While phenograms and cladograms represented different forms of knowledge during the time of the Systematist Wars, today they are both used to represent evolutionary relationships. This indicates that diagrams are versatile elements of scientific practice that can change their meaning, depending on the context of use within theoretical frameworks. (shrink)

80 titles published between 1965 and 1996 in multiple languages attest to an increase in scholarly interest in the history of systematic biology, both among scientific practitioners and also among historians and philosophers of science. Topics studied have included the early history of the field (Ray, Linnaeus, Buffon), the influence of essentialism on systematics, the history of systematic diagrams, the development of cladistic analysis, the nature of species, and the growth of phylogenetic thinking.

Some plausibly necessary a posteriori theoretical claims include ‘water is H 2 O’, ‘gold is the element with atomic number 79’, and ‘cats are animals’. In this paper I challenge the necessity of the third claim. I argue that there are possible worlds in which cats exist, but are not animals. Under any of the species concepts currently accepted in biology, organisms do not belong essentially to their species. This is equally true of their ancestors. In phylogenetic systematics, (...) monophyletic clades such as the animal kingdom are composed of an ancestral stem species and all of its descendants. If the stem species had not existed, neither would the clade. Thus it could have been the case that all the organisms which actually belong to the animal kingdom might have existed yet not have been animals. (shrink)

Taxa and homologues can in our view be construed both as kinds and as individuals. However, the conceptualization of taxa as natural kinds in the sense of homeostatic property cluster kinds has been criticized by some systematists, as it seems that even such kinds cannot evolve due to their being homeostatic. We reply by arguing that the treatment of transformational and taxic homologies, respectively, as dynamic and static aspects of the same homeostatic property cluster kind represents a good perspective for (...) supporting the conceptualization of taxa as kinds. The focus on a phenomenon of homology based on causal processes (e.g., connectivity, activity-function, genetics, inheritance, and modularity) and implying relationship with modification yields a notion of natural kinds conforming to the phylogenetic-evolutionary framework. Nevertheless, homeostatic property cluster kinds in taxonomic and evolutionary practice must be rooted in the primacy of epistemological classification (homology as observational properties) over metaphysical generalization (series of transformation and common ancestry as unobservational processes). The perspective of individuating characters exclusively by historical-transformational independence instead of their developmental, structural, and functional independence fails to yield a sufficient practical interplay between theory and observation. Purely ontological and ostensional perspectives in evolution and phylogeny (e.g., an ideographic character concept and PhyloCode’s ‘individualism’ of clades) may be pragmatically contested in the case of urgent issues in biodiversity research, conservation, and systematics. (shrink)

The genetically and experimentally accessible organs of Drosophila, such as the heart or blood‐forming tissues, have become a fertile ground for systematic projects of gene discovery and for functional studies of gene networks and signaling pathways. One argument justifying this approach is the often‐tacit assumption that clear‐cut homologies can be established between the Drosophila organs and their vertebrate counterparts. Here we investigate this assumption by surveying pertinent aspects of vascular structure and development in different invertebrate phyla, in the hope that (...) this information will help to reveal the ancestral condition of the vascular system. Evolutionary scenarios that derive the structure of the cardiovascular system of extant animal taxa from the ancestral condition will be used to qualify hypotheses regarding homologies that are based on molecular similarities. BioEssays 28: 1203–1210, 2006. © 2006 Wiley Periodicals, Inc. (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)

Phylogenetic systematics is one of the most important analytical frameworks of modern Biology. It seems to be common knowledge that within phylogenetics, ‘groups’ must be defined based solely on the synapomorphies or on the “derived” characters that unite two or more taxa in a clade or monophyletic group. Thus, the idea of synapomorphy seems to be of fundamental influence and importance. Here I will show that the most common and straightforward understanding of synapomorphy as a shared derived character (...) is not sufficient and eventually must be rejected in favor of Nelson’s relational interpretation of such term. Arguing for this point and using three examples from previously published Apes’ genomic matrices, I explicitly demonstrate that the relationship )) with Hylobatidae as a sister taxon, may be successfully recovered by three-taxon statement analysis and three-taxon statement average consensus analysis even if all of the evident standard shared derived molecular characters of the relationship )) with Hylobatidae as a sister taxon, have been excluded from the molecular alignments. Neither conventional Maximum Parsimony nor Maximum Likelihood or Bayesian Inference can do this in such situation. Thus, our results show that the relationship )) with Hylobatidae as a sister taxon has appeared, in some way, behind standard shared derived characters: the last ones could be excluded, but the relationship remains the same. (shrink)

The taxa that appear in biological classifications are commonly seen as representing information about the traits of their member organisms. This paper examines in what way taxa feature in the storage and retrieval of such information. I will argue that taxa do not actually store much information about the traits of their member organisms. Rather, I want to suggest, taxa should be understood as functioning to localize organisms in the genealogical network of life on Earth. Taxa store information about where (...) organisms are localized in the network, which is important background information when it comes to establishing knowledge about organismal traits, but it is not itself information about these traits. The view of species and higher taxa that is proposed here follows from examining three problems that occur in contemporary biological systematics and are discussed here: the problem of generalization over taxa, the problem of phylogenetic inference, and the problematic nature of the Tree of Life. (shrink)

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

The role of scientific theories in classifying plants and animals is traced from Hennig's phylogenetics and the evolutionary taxonomy of Simpson and Mayr, through numerical phenetics, to present-day cladistics. Hennig limited biological classification to sister groups so that this one relation can be expressed unambiguously in classifications. Simpson and Mayr were willing to sacrifice precision in representation in order to include additional features of evolution in the construction of classifications. In order to make classifications more objective, precise and quantitative, numerical (...) pheneticists limited themselves to representing degrees of phenetic similarity. Finally, present-day cladists can be separated into phylogenetic cladists, who retain much of Hennig's theory of classification, and pattern cladists, who have stripped Hennig's system down to its bare essentials. (shrink)

In biological systematics, as well as in the philosophy of biology, species and higher taxa are individuated through their unique evolutionary origin. This is taken by some authors to mean that monophyly is a (relational) property not only of higher taxa, but also of species. A species is said to originate through speciation, and to go extinct when it splits into two daughter species (or through terminal extinction). Its unique evolutionary origin is said to bestow identity on a species (...) through time and change, and to render species names rigid designators. Species names are thus believed to function just like names of supraspecific taxa. However, large parts of the Web of Life are composed of species that do not have a unique evolutionary origin from a single population, lineage or stem-species. Further, monophyly is an ambiguous concept if it is defined simply in terms of ‘unique evolutionary origin’. Disambiguating the concept by defining a monophyletic taxon as ‘a taxon that includes the ancestor and all, and only, its descendant’ renders monophyly inapplicable to species. At the heart of the problem lies a fundamental distinction between species and monophyletic taxa, where species form mutually exclusive reticulated systems, while higher taxa form inclusive hierarchical systems. Examples are given both at the species level and below to illustrate the problems that result from the application of the monophyly criterion to species. The conclusion is that the concepts of exclusivity and monophyly should be treated as non-overlapping: exclusivity marks out a species synchronistically, i.e. in the present time. Monophyly marks out clades (groups of species) diachronistically, i.e. within an historical dimension. (shrink)

Short interspersed repetitive elements, or SINEs, are tRNA-derived retroposons that are dispersed throughout eukaryotic genomes and can be present in well over 104 total copies. The enormous volume of SINE amplifications per organism makes them important evolutionary agents for shaping the diversity of genomes, and the irreversible, independent nature of their insertion allows them to be used for diagnosing common ancestry among host taxa with extreme confidence. As such, they represent a powerful new tool for systematic biology that can be (...) strategically integrated with other conventional phylogenetic characters, most notably morphology and DNA sequences. This review covers the basic aspects of SINE evolution that are especially relevant to their use as systematic characters and describes the practical methods of characterizing SINEs for cladogram construction. It also discusses the limits of their systematic utility, clarifies some recently published misunderstandings, and illustrates the effective application of SINEs for vertebrate phylogenetics with results from selected case studies. BioEssays 22:148–160, 2000. ©2000 John Wiley & Sons, Inc. (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.

Although molecular systematists may use the terminology of cladism, claiming that the reconstruction of phylogenetic relationships is based on shared derived states , the latter is not the case. Rather, molecular systematics is based on the assumption, first clearly articulated by Zuckerkandl and Pauling , that degree of overall similarity reflects degree of relatedness. This assumption derives from interpreting molecular similarity between taxa in the context of a Darwinian model of continual and gradual change. Review of the history (...) of molecular systematics and its claims in the context of molecular biology reveals that there is no basis for the “molecular assumption.”. (shrink)

The role of scientific theories in classifying plants and animals is traced from Hennig's phylogenetics and the evolutionary taxonomy of Simpson and Mayr, through numerical phenetics, to present-day cladistics. Hennig limited biological classification to sister groups so that this one relation can be expressed unambiguously in classifications. Simpson and Mayr were willing to sacrifice precision in representation in order to include additional features of evolution in the construction of classifications. In order to make classifications more objective, precise and quantitative, numerical (...) pheneticists limited themselves to representing degrees of phenetic similarity. Finally, present-day cladists can be separated into phylogenetic cladists, who retain much of Hennig's theory of classification, and pattern cladists, who have stripped Hennig's system down to its bare essentials. (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)

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)

There is long-standing conflict between genealogical and developmental accounts of homology. This paper provides a general framework that shows that these accounts are compatible and clarifies precisely how they are related. According to this framework, understanding homology requires both an abstract genealogical account that unifies the application of the term to all types of characters used in phylogenetic systematics and locally enriched accounts that apply only to specific types of characters. The genealogical account serves this unifying role by (...) relying on abstract notions of ‘descent’ and ‘character’. As a result, it takes for granted the existence of such characters. This requires theoretical justification that is provided by enriched accounts, which incorporate the details by which characters are inherited. These enriched accounts apply to limited domains, providing the needed theoretical justification for recognizing characters within that domain. Though connected to the genealogical account of homology in this way, enriched accounts include phenomena that fall outside the scope of the genealogical account. They therefore overlap, but are not nested within, the genealogical account. Developmental accounts of homology are to be understood as enriched accounts of body part homology. Once they are seen in this light, the conflict with the genealogical account vanishes. It is only by understanding the fine conceptual structure undergirding the many uses of the term ‘homology’ that we can understand how these uses hang together. (shrink)