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

Why has it been so difficult to integrate paleontology and mainstream evolutionary biology? Two common answers are: (1) the two fields have fundamentally different aims, and (2) the tensions arise out of disciplinary squabbles for funding and prestige. This paper examines the role of fossil data in phylogeny reconstruction in order to assess these two explanations. I argue that while cladistics has provided a framework within which to integrate fossil character data, the stratigraphic (temporal) component of fossil data (...) has been harder to integrate. A close examination of how fossil data have been used in phylogeny reconstruction suggests that neither explanation is adequate. While some of the tensions between the fields may be intellectual turf wars, the second explanation downplays the genuine difficulty of combining the distinctive data of the two fields. Furthermore, it is simply not the case that the two fields pursue completely distinct aims. Systematists do disagree about precisely how to represent phylogeny (e.g., minimalist cladograms or trees with varying levels of detail) but given that every tree presupposes a pattern of branching (a cladogram), these aims are not completely distinct. The central problem has been developing methods that allow scientists to incorporate the distinctive bodies of data generated by these two fields. Further case studies will be required to determine if this explanation holds for other areas of interaction between paleontology and neontology. (shrink)

The concept that renders morphology a tool for phylogeny reconstruction is homology. The concept of homology is rooted in pre-evolutionary idealistic morphology. The claim that the goal of idealistic morphology was the seriability of form may sound paradoxical given that this discipline proceeded within a framework of strictly delimited types. But the types only demarcate where seriability starts and where it comes to an end. Carl Gegenbaur’s was recognized as a milestone in idealistic morphology. A comparison with the (...) second edition of 1870 illustrates Gegenbaur’s turn to evolutionary morphology. The methodology remained the same–seriability of form–but the series was no longer merely descriptive or conceptual but now a historical, evolutionary one. Gegenbaur emphasized that seriability of form was possible not only between species of the same type, but also between parts of organisms of the same type. Pursuing this project, he found that different parts of organisms evolve at different rates, resulting in an incongruence between the series of parts relative to the series of species under comparison. This incongrence was called chevauchement des spécialisations by Louis Dollo, Spezialisationskreuzungen by Othenio Abel, and heterobathmy of characters by Armen Takhtajan. Willi Hennig, the founder of modern methods in phylogenetic systematics, discovered that the heterobathmy of characters was a precondition for the establishment of the phylogenetic relationships based on shared derived characters. The result was a replacement of the search for ancestors by a search for relative degrees of phylogenetic relationships. (shrink)

Phylogenies are increasingly being used to investigate human history, diversification and cultural evolution. While using phylogenies in this way is not new, new modes of analysis are being applied to inferring history, reconstructing past states, and examining processes of change. Phylogenies have the advantage of providing a way of creating a continuous history of all current populations, and they make a large number of analyses and hypothesis tests possible even when other forms of historical information are patchy or nonexistent. In (...) common with approaches taken in other historical sciences, phylogenetics is a way of reconstructing past and processes using the traces left in the present day. Trees, based on DNA, language, cultural traits, or other evidence, are now sprouting all over the academic landscape. The increasing use of phylogenetic analysis to understand human cultural evolution has been embraced by some, and scorned by others. The purpose of this article is not to review methods and applications of phylogenetic analyses, nor to consider the growing field of cultural phylogenetics, but, more broadly, to explore how we interpret phylogenies as narratives about human diversification. The first half of the article deals with meaning: phylogenies are often interpreted as histories, but a bifurcating tree is at best an abstract representation of history, and its connections to past events and processes is dependent on the data used, the assumptions made in the analysis, and the degree to which nodes in the tree (where one lineage splits into two) can be connected to change and movement in real populations. The second half of the article explores the purpose of phylogenies: a tree does not have to be a literal history of human lineages in order to be useful for investigating processes of human diversification. Phylogenies should not be read as accurate records of history, but as a way of exploring plausible explanations for current patterns of diversity. Phylogenies provide important information that can be used to test ideas about human diversity, and can help to guard against errors of inference arising from statistical artifacts. (shrink)

Much is being written these days about integration, its desirability and even its necessity when complex research problems are to be addressed. Seldom, however, do we hear much about the failure of such efforts. Because integration is an ongoing activity rather than a final achievement, and because today’s literature about integration consists mostly of manifesto statements rather than precise descriptions, an examination of unsuccessful integration could be illuminating to understand better how it works. This paper will examine the case of (...) prokaryote phylogeny and its apparent failure to achieve integration within broader tree-of-life accounts of evolutionary history. Despite the fact that integrated databases exist of molecules pertinent to the phylogenetic reconstruction of all lineages of life, and even though the same methods can be used to construct phylogenies wherever the organisms fall on the tree of life, prokaryote phylogeny remains at best only partly integrated within tree-of-life efforts. I will examine why integration does not occur, compare it with integrative practices in animal and other eukaryote phylogeny, and reflect on whether there might be different expectations of what integration should achieve. Finally, I will draw some general conclusions about integration and its function as a ‘meta-heuristic’ in the normative commitments guiding scientific practice. (shrink)

Biology and the social sciences share an interest in phylogeny. Biologists know that living species are descended from past species, and use the pattern of similarities among living species to reconstruct the history of phylogenetic branching. Social scientists know that the beliefs, values, practices, and artifacts that characterize contemporary societies are descended from past societies, and some social science disciplines, linguistics and cross cultural anthropology for example, have made use of observed similarities to reconstruct cultural histories. Darwin appreciated that (...) his theory of descent with modification had many similarities of pattern and process to the already well developed field of historical linguistics. In many other areas of social science, however, phylogenetic reconstruction has not played a central role. (shrink)

The reconstruction of bacterial evolutionary relationships has proven to be a daunting task because variable mutation rates and horizontal gene transfer (HGT) among species can cause grave incongruities between phylogenetic trees based on single genes. Recently, a highly robust phylogenetic tree was constructed for 13 γ‐proteobacteria using the combined alignments of 205 conserved orthologous proteins.1 Only two proteins had incongruent tree topologies, which were attributed to HGT between Pseudomonas species and Vibrio cholerae or enterics. While the evolutionary relationships among (...) these species appears to be resolved, further analysis suggests that HGT events with other bacterial partners likely occurred; this alters the implicit assumption of γ‐proteobacteria monophyly. Thus, any thorough reconstruction of bacterial evolution must not only choose a suitable set of molecular markers but also strive to reduce potential bias in the selection of species. BioEssays 26:463–468, 2004. © 2004 Wiley Periodicals, Inc. (shrink)

Despite the promises made by molecular evolutionists since the early 1960s that phylogenies would be readily reconstructed using molecular data, the construction of molecular phylogenies has both retained many methodological problems of the past and brought up new ones of considerable epistemic relevance. The field is driven not only by changes in knowledge about the processes of molecular evolution, but also by an ever-present methodological anxiety manifested in the constant search for an increased objectivity—or in its converse, the avoidance of (...) subjectivity.This paper offers an exhaustive account of the methodological and conceptual difficulties embedded in each of the steps required to elaborate molecular phytogenies. The authors adopt a historical perspective on the field in order to follow the development of practices that seek to increase the objectivity of their methods and representations. These include the adoption and development of explicit criteria for evaluation of evidence, and of procedures associated with methods of statistical inference, quantification and automation. All these are linked to an increasing use of computers in research since the mid 1960s. We will show that the practices of objectivity described are highly dependent on the problems and tools of molecular phylogenetics. (shrink)

The theoretical biology of Jakob von Uexküll has had significant conceptual and practical afterlives, in Continental philosophy, biosemiotics and elsewhere. This paper will examine the utilisation of Uexküll in twentieth-century zoo biology and its significance for relating to wildlife in hybrid environments. There is an important though rarely analysed line of inheritance from von Uexküll to Heini Hediger, the Swiss zoo director and animal psychologist. Hediger’s fundamental theoretical position began from the construction of the world from the animal’s point of (...) view, as determined by factors including species specific phylogeny, individual and group biography, and anthropogenic circumstance. He operationalised Uexküll’s approach to animal worlds in order to optimise the design of zoo enclosures, considered as both physical and psychological habitats, in which captive wildlife could flourish. This subjectivist and phenomenological perspective has often been sidelined in zoo biology by more objectivist and mechanising approaches. Nonetheless, Hediger’s work and thought, through its inheritance from Uexküll, has important implications for twenty-first century relations with wildlife. (shrink)

Comparative biological morphology, incorporating the study of active reaction, is contrasted with genetics as the study of passive mutation. Geneticists investigate anatomical characters, never anatomical constructions, which are capable of reorganization when the biological-morphological equilibrium of the organism has been disturbed. The anatomy of Opisthocomus cristatus and Stringops habroptilus demonstrate that three successive disturbances in the bio-morphological equilibrium are reacted to purposively by anatomical reconstruction. These reactions are no accidental mutations, but are anatomical reactions, related to, and affecting, the (...) organism as a whole. In sharp contrast to such anatomical reaction, resulting, during phylogeny, in reorganization, are the “technics” [i.e., mechanistic bases] of individual development. The hereditary process is, like every physiological or embryological process, a fixed mechanism, which remains constant until an active reaction leads to reconstruction and at the same time an appropriate change of the mechanisms. The remolding of species is therefore no passive, “technical” process, but a creative act of the organisms themselves. [Original English abstract; not translated.]. (shrink)

"Habermas introduces the concept of “reconstructive science” with a double purpose: to place the “general theory of society” between philosophy and social science and reestablish the rift between the “great theorization” and the “empirical research”. The model of “rational reconstructions” represents the main thread of the surveys about the “structures” of the life-world (“culture”, “society” and “personality”) and their respective “functions” (cultural reproductions, social integrations and socialization). For this propose, the dialectics between “symbolic representation” of “the structures subordinated to all (...) worlds of life” (“internal relationships”) and the “material reproduction” of the social systems in their complex (“external relationships” between social systems and environment) has to be considered. This model finds an application, above all, in the “theory of the social evolution”, starting from the reconstruction of the necessary conditions for a phylogeny of the socio-cultural-life forms (the “hominization”) until an analysis of the development of “social formations”, which Habermas subdivides into primitive, traditional, modern and contemporary formations. This paper is an attempt, primarily, to formalize the model of “reconstruction of the logic of development” of “social formations” summed up by Habermas through the differentiation between vital world and social systems (and, within them, through the “rationalization of the life-world” and the “growth in complexity of the social systems”). Secondly, it tries to offer some methodological clarifications about the “explanation of the dynamics” of “historical processes” and, in particular, about the “theoretical meaning” of the evolutional theory’s propositions. Even if the German sociologist considers that the “ex-post rational reconstructions” and “the models system/environment” cannot have a complete “historiographical application”, these certainly act as a general premise in the argumentative structure of the “historical explanation”.". (shrink)

In a recent paper,(1) palaeontologist Mike Benton claimed that our ability to reconstruct accurately the tree of Life may not have improved significantly over the last 100 years. This implies that the cladistic and molecular revolutions may have promulgated as much bad “black box” science as rigorous investigation. Benton's assessment was based on the extent to which cladograms (typically constructed with reference only to distributions of character states) convey the same narrative as the geochronological ages of fossil taxa (an independent (...) data set). Fossil record quality varies greatly between major clades, and the palaeontological dating “yardstick” may be more appropriate for some groups than others. BioEssays 24:203–207, 2002. © 2002 Wiley Periodicals, Inc.; DOI 10.1002/bies.10065. (shrink)

Deep‐level arthropod phylogeny has been in a state of upheaval ever since the emergence of molecular tree reconstruction approaches. While a consensus has settled in that hexapods are more closely related to crustaceans than to myriapods, the phylogenetic position of the latter has remained a matter of debate. Mitochondrial, nuclear, and genome‐scale studies have proposed rejecting the long‐standing superclade Mandibulata, which unites myriapods with insects and crustaceans, in favor of a clade that unites myriapods with chelicerates and has (...) become known as Paradoxapoda or Myriochelata. Here we discuss the progress, problems, and prospects of arriving at the final arthropod tree. (shrink)

The German synthesis of evolutionary theory that grew out of opposition to idealistic morphology has been anchored in the systematic work at the species level and below pursued by the Berlin School around Erwin Stresemann (involving Bernhard Rensch and Ernst Mayr), in the 1939 German translation of Dobzhansky’s Genetics and the Origin of Species, and in a 1943 anthology on evolution edited by Gerhard Heberer. The latter volume opened with a philosophical essay written by Hugo Dingler that was intended to (...) provide the theoretical foundation for the theory of descent. Dingler and Heberer not only shared a commitment to the Darwinian evolutionary mechanisms of random mutation and natural selection, but also drew from Darwinism the justification for racial hygiene and eugenics. Dingler’s “unequivocal-methodological system” is a meta-scientific construct that offers no ontological grounding of evolutionary theory. Dingler’s voluntarism is subjectivist and stipulative and for those reasons ineffective in a critique of idealistic morphology. Dingler claimed descent with modification as a historical fact, but dealt only marginally with evolutionary mechanisms, and did not touch on issues of phylogeny reconstruction. (shrink)

One of the major criticisms of optimal foraging theory (OFT) is that it is not testable. In discussions of this criticism opposing parties have confused methodological concepts and used meaningless biological concepts. In this paper we discuss such misunderstandings and show that OFr has an empirically testable, and even well-confirmed, general core theory. One of our main conclusions is that specific model testing should not be aimed at proving optimality, but rather at identifying the context in which certain types of (...) behaviour are optimal. To do this, it is necessary to be aware of the assumptions made in testing a model. The assumptions that are explicitly stated in the literature up to now do not completely cover the actual assumptions made in testing OFT models in practice. We present a more comprehensive set of assumptions. Although all the assumptions play a role in testing models, they are not of equal status. Crucial assumptions concern constraints and the relation between fitness and currency. Therefore, it is essential to make such assumptions testable in practice. We show that a more explicit relationship between OFT modelling and evolutionary theory can help with this. Specifically, phylogeny reconstruction and population dynamic modelling can and should be used to formulate assumptions concerning constraints and currencies. (shrink)

A long-standing debate has dominated systematic biology and the ontological commitments made by its theories. The debate has contrasted individuals and the part – whole relationship with classes and the membership relation. This essay proposes to conceptualize the hierarchy of higher taxa is terms of a hierarchy of homeostatic property cluster natural kinds (biological species remain largely excluded from the present discussion). The reference of natural kind terms that apply to supraspecific taxa is initially fixed descriptively; the extension of those (...) natural kind terms is subsequently established by empirical investigation. In that sense, classification precedes generalization, and description provides guidance to empirical investigation. The reconstruction of a hierarchy of (homeostatic property cluster) natural kinds is discussed in the light of cladistic methods of phylogeny reconstruction. (shrink)

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

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)

One of the major criticisms of optimal foraging theory is that it is not testable. In discussions of this criticism opposing parties have confused methodological concepts and used meaningless biological concepts. In this paper we discuss such misunderstandings and show that OFr has an empirically testable, and even well-confirmed, general core theory. One of our main conclusions is that specific model testing should not be aimed at ‘proving’ optimality, but rather at identifying the context in which certain types of behaviour (...) are optimal. To do this, it is necessary to be aware of the assumptions made in testing a model. The assumptions that are explicitly stated in the literature up to now do not completely cover the actual assumptions made in testing OFT models in practice. We present a more comprehensive set of assumptions. Although all the assumptions play a role in testing models, they are not of equal status. Crucial assumptions concern constraints and the relation between fitness and currency. Therefore, it is essential to make such assumptions testable in practice. We show that a more explicit relationship between OFT modelling and evolutionary theory can help with this. Specifically, phylogeny reconstruction and population dynamic modelling can and should be used to formulate assumptions concerning constraints and currencies. (shrink)

Ferritins (FTs) are iron storage proteins that are involved in managing iron‐oxygen balance. In our work, we present a hypothesis on the putative effect of geological changes that have affected the evolution and radiation of ferritin proteins. Based on sequence analysis and phylogeny reconstruction, we hypothesize that two significant factors have been involved in the evolution of ferritin proteins: fluctuations of atmospheric oxygen concentrations, altering redox potential, and changing availability of water rich in bioavailable ferric ions.Fish, ancient amphibians, (...) reptiles, and placental mammals developed the broadest repertoire of singular FTs, attributable to embryonic growth in aquatic environments containing low oxygen levels and abundant forms of soluble iron. In contrast, oviparous land vertebrates, like reptiles and birds, that have developed in high oxygen levels and limited levels of environmental Fe2+ exhibit a lower diversity of singular FTs, but display a broad repertoire of subfamilies, particularly notable in early reptiles. (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.

We show that, contrary to long-standing assumptions, syntactic traits, modeled here within the generative biolinguistic framework, provide insights into deep-time language history. To support this claim, we have encoded the diversity of nominal structures using 94 universally definable binary parameters, set in 69 languages spanning across up to 13 traditionally irreducible Eurasian families. We found a phylogenetic signal that distinguishes all such families and matches the family-internal tree topologies that are safely established through classical etymological methods and datasets. We have (...) retrieved “near-perfect” phylogenies, which are essentially immune to homoplastic disruption and only moderately influenced by horizontal convergence, two factors that instead severely affect more externalized linguistic features, like sound inventories. This result allows us to draw some preliminary inferences about plausible/implausible cross-family classifications; it also provides a new source of evidence for testing the representation of diversity in syntactic theories. (shrink)

Planctomycetes, Verrucomicrobia and Chlamydia are prokaryotic phyla, sometimes grouped together as the PVC superphylum of eubacteria. Some PVC species possess interesting attributes, in particular, internal membranes that superficially resemble eukaryotic endomembranes. Some biologists now claim that PVC bacteria are nucleus‐bearing prokaryotes and are considered evolutionary intermediates in the transition from prokaryote to eukaryote. PVC prokaryotes do not possess a nucleus and are not intermediates in the prokaryote‐to‐eukaryote transition. Here we summarise the evidence that shows why all of the PVC traits (...) that are currently cited as evidence for aspiring eukaryoticity are either analogous (the result of convergent evolution), not homologous, to eukaryotic traits; or else they are the result of horizontal gene transfers. (shrink)

The ‘Tree of Life’ is intended to represent the pattern of evolutionary processes that result in bifurcating species lineages. Often justified in reference to Darwin’s discussions of trees, the Tree of Life has run up against numerous challenges especially in regard to prokaryote evolution. This special issue examines scientific, historical and philosophical aspects of debates about the Tree of Life, with the aim of turning these criticisms towards a reconstruction of prokaryote phylogeny and even some aspects of the (...) standard evolutionary understanding of eukaryotes. These discussions have arisen out of a multidisciplinary collaboration of people with an interest in the Tree of Life, and we suggest that this sort of focused engagement enables a practical understanding of the relationships between biology, philosophy and history. (shrink)

The origin of the bilaterian metazoans from radial ancestors is one of the biggest puzzles in animal evolution. A way to solve it is to identify the nature and main features of the last common ancestor of the bilaterians (LCB). Recent progress in molecular phylogeny has shown that many platyhelminth flatworms, regarded for a long time as basal bilaterians, now belong to the lophotrochozoan protostomates. In contrast, the LCB is now considered a complex organism bearing several features of modern (...) bilaterians. Here we discuss an alternative view, in which acoelomorph (Acoela + Nemertodermatida) flatworms, which do not belong to the Platyhelminthes, represent the earliest extant bilaterian clade. Sequences from ribosomal and other nuclear genes, Hox cluster genes, and reinterpretation of some morphological features strongly support the basal position of acoelomorphs arguing against a complex LCB. This reconstruction backs the old planuloid–acoeloid hypothesis and may help our understanding of the evolution of body axes, Hox genes and the Cambrian explosion. BioEssays 26:1046–1057, 2004. © 2004 Wiley Periodicals, Inc. (shrink)

The eukaryotic genome is a mosaic of eubacterial and archaeal genes in addition to those unique to itself. The mosaic may have arisen as the result of two prokaryotes merging their genomes, or from genes acquired from an endosymbiont of eubacterial origin. A third possibility is that the eukaryotic genome arose from successive events of lateral gene transfer over long periods of time. This theory does not exclude the endosymbiont, but questions whether it is necessary to explain the peculiar set (...) of eukaryotic genes. We use phylogenetic studies and reconstructions of ancestral first appearances of genes on the prokaryotic phylogeny to assess evidence for the lateral gene transfer scenario. We find that phylogenies advanced to support fusion can also arise from a succession of lateral gene transfer events. Our reconstructions of ancestral first appearances of genes reveal that the various genes that make up the eukaryotic mosaic arose at different times and in diverse lineages on the prokaryotic tree, and were not available in a single lineage. Successive events of lateral gene transfer can explain the unusual mosaic structure of the eukaryotic genome, with its content linked to the immediate adaptive value of the genes its acquired. Progress in understanding eukaryotes may come from identifying ancestral features such as the eukaryotic splicesome that could explain why this lineage invaded, or created, the eukaryotic niche. BioEssays 28:57–64, 2006. © 2005 Wiley Periodicals, Inc. (shrink)

Preservation permitting patterns of developmental evolution can be reconstructed within long extinct clades, and the rich fossil record of trilobite ontogeny and phylogeny provides an unparalleled opportunity for doing so. Furthermore, knowledge of Hox gene expression patterns among living arthropods permit inferences about possible Hox gene deployment in trilobites. The trilobite anteroposterior body plan is consistent with recent suggestions that basal euarthropods had a relatively low degree of tagmosis among cephalic limbs, possibly related to overlapping expression domains of cephalic (...) Hox genes. Trilobite trunk segments appeared sequentially at a subterminal generative zone, and were exchanged between regions of fused and freely articulating segments during growth. Homonomous trunk segment shape and gradual size transition were apparently phylogenetically basal conditions and suggest a single trunk tagma. Several derived clades independently evolved functionally distinct tagmata within the trunk, apparently exchanging flexible segment numbers for greater regionally autonomy. The trilobite trunk chronicles how different aspects of arthropod segmentation coevolved as the degree of tagmosis increased. BioEssays 25:386–395, 2003. © 2003 Wiley Periodicals, Inc. (shrink)

The origin of eukaryotes is one of the major challenges of evolutionary cell biology. Other than the endosymbiotic origin of mitochondria and chloroplasts, the steps leading to eukaryotic endomembranes and endoskeleton are poorly understood. Ras‐family small GTPases are key regulators of cytoskeleton dynamics, vesicular trafficking and nuclear function. They are specific for eukaryotes and their expansion probably traces the evolution of core eukaryote features. The phylogeny of small GTPases suggests that the first endomembranes to evolve during eukaryote evolution had (...) secretory, and not phagocytic, function. Based on the reconstruction of putative roles for ancestral small GTPases, a hypothetical scenario on the origins of the first endomembranes, the nucleus, and phagocytosis is presented. BioEssays 25:1129–1138, 2003. © 2003 Wiley Periodicals, Inc. (shrink)

In their 1987 Nature publication, “Mitochondrial DNA and Human Evolution,” Rebecca Cann, Mark Stoneking, and Allan C. Wilson gave a new reconstruction of human evolution on the basis of differences in mitochondrial DNA among contemporary human populations. This phylogeny included an African common ancestor for all human mitochondrial DNA (mtDNA) lineages, and Cann et al.’s reconstruction became known as the “Out of Africa” hypothesis. Since mtDNA is inherited exclusively through the maternal line, the common ancestor who was (...) first branded African Eve later became known as Mitochondrial Eve (mtEve, for short). -/- In this paper, I show that mtEve was not a single, successful, or purely scientific discovery. Instead, she was produced many times and in many ways, each of which informed the next. Importantly, though Wilson and colleagues heralded mitochondrial DNA as a source of certainty, objectivity, and consensus for evolutionary inference, their productions of Mitochondrial Eve depended as much on popular assumptions about the certainty of maternal inheritance as they did on new molecular and computational tools. This recognition lets us reevaluate the complex consequences of these productions, which, like mtEve herself, could not be confined to a purely social, material, or scientific dimension. (shrink)

Erwin Schrödinger’s 1944 publication What is Life? is a classic of twentieth century science writing. In his book, Schrödinger discussed the chromosome fibre as the seat of heredity and variation thanks to a hypothetical aperiodic structure – a suggestion that famously spurred on a generation of scientists in their pursuit of the gene as a physico-chemical entity. While historical attention has been given to physicists who were inspired by the book, little has been written about its biologist readers. This paper (...) examines the case of the English evolutionary botanist and cytologist Irène Manton, who took an interest in What is Life? for its relevance to her own research in chromosome structure as a clue to plant phylogeny. Drawing on recently discovered correspondence between Manton and Schrödinger, the paper reconstructs Manton ‘s path to the book and her response to it by way of throwing new light on a pivotal moment in the history of the debate on chromosome structure. (shrink)

The history of biological systematics documents a continuing tension between classifications in terms of nested hierarchies congruent with branching diagrams (the ‘Tree of Life’) versus reticulated relations. The recognition of conflicting character distribution led to the dissolution of the scala naturae into reticulated systems, which were then transformed into phylogenetic trees by the addition of a vertical axis. The cladistic revolution in systematics resulted in a representation of phylogeny as a strictly bifurcating pattern (cladogram). Due to the ubiquity of (...) character conflict—at the genetic or morphological level, or at any level in between—some characters will necessarily have to be discarded ( qua noise) in favor of others in support of a strictly bifurcating phylogenetic tree. Pattern analysts will seek maximal congruence in the distribution of characters (ultimately of any kind) relative to a branching tree-topology; process explainers will call such tree-topologies into question by reference to incompatible evolutionary processes. Pattern analysts will argue that process explanations must not be brought to bear on pattern reconstruction; process explainers will insist that the reconstructed pattern requires a process explanation to become scientifically relevant, i.e., relevant to evolutionary theory. The core question driving the current debate about the adequacy of the ‘Tree of Life’ metaphor seems to be whether the systematic dichotomization of the living world is an adequate representation of the complex evolutionary history of global biodiversity. In ‘Questioning the Tree of Life’, it seems beneficial to draw at least four conceptual distinctions: pattern reconstruction versus process explanation as different epistemological approaches to the study of phylogeny; open versus closed systems as expressions of different kinds of population (species) structures; phylogenetic trees versus cladograms as representations of evolutionary processes versus patterns of relationships; and genes versus species as expressions of different levels of causal integration and evolutionary transformation. (shrink)

Biological lineages move through time, space, and each other. As they do, they diversify, diverge, and grade away from and into one another. One result of this is genealogical discordance; i.e., the lineages of a biological entity may have different histories. We see this on numerous levels, from microbial networks, to holobionts, to population-level lineages. This paper considers how genealogical discordance impacts our study of species. More specifically, I consider this in the context of three framing questions: (1) How, if (...) at all, does genealogical discordance challenge, modify, or revise how we conceive of species? (2) How has growing appreciation of genealogical discordance impacted scientific practice? Of systematics in particular? (3) How do lineages at different levels diverge and diversify? All told, genealogical discordance enriches and complicates our taxonomic ontology, as well as the practice of reconstructing phylogenies. This presents both challenges and opportunities for the study of divergence and diversification. (shrink)

We discuss the impact of horizontal gene transfer (HGT) on phylogenetic reconstruction and taxonomy. We review the power of HGT as a creative force in assembling new metabolic pathways, and we discuss the impact that HGT has on phylogenetic reconstruction. On one hand, shared derived characters are created through transferred genes that persist in the recipient lineage, either because they were adaptive in the recipient lineage or because they resulted in a functional replacement. On the other hand, taxonomic (...) patterns in microbial phylogenies might also be created through biased gene transfer. The agreement between different molecular phylogenies has encouraged interpretation of the consensus signal as reflecting organismal history or as the tree of cell divisions; however, to date the extent to which the consensus reflects shared organismal ancestry and to which it reflects highways of gene sharing and biased gene transfer remains an open question. Preferential patterns of gene exchange act as a homogenizing force in creating and maintaining microbial groups, generating taxonomic patterns that are indistinguishable to those created by shared ancestry. To understand the evolution of higher bacterial taxonomic units, concepts usually applied in population genetics need to be applied. (shrink)

Since Darwin's theory of evolution by natural selection, the idea of descent with modification came to dominate systematics, and so the study of morphology became subgugated to the reconstruction of phylogenies. Reinstating the organism in the theory of evolution (Ho & Saunders, 1979; Webster & Goodwin, 1982) leads to a project inrational taxonomy (Ho, 1986, 1988a), which attempts to classify biological forms on the basis of transformations on a given dynamical structure.Does rational taxonomy correspond to thenatural system that Linnaeus (...) and his contemporaries as well as all pre-Darwinian morphologists had in mind? Here, we examine how rational taxonomy and the natural system can coincide in the dynamics of processes generating forms during development, which conferexclusivity, genericity androbustness to the forms that do exist. We use the example of segmentation, especially inDrosophila, as an illustration to explore the implications of rational taxonomy for evolution and systematics, and the relationship between ontogeny and phylogeny. (shrink)

Adaptive immunity is unique to the vertebrates, and the molecules involved (including immunoglobulins, T cell receptors and the major histocompatibility complex molecules) seem to have diversified very rapidly early in vertebrate history. Reconstruction of gene phylogenies has yielded insights into the evolutionary origin of a number of molecular systems, including the complement system and the major histocompatibility complex (MHC). These analyses have indicated that the C5 component of complement arose by gene duplication prior to the divergence of C3 and (...) C4, which suggests that the alternative complement pathway was the first to evolve. In the case of the MHC, phylogenetic analysis supports the hypothesis that MHC class II molecules evolved before class I molecules. The fact that the MHC‐linked proteasome components that specifically produce peptides for presentation by class I MHC appear to have originated before the separation of jawed and jawless vertebrates suggests that the MHC itself may have been present at this time. Immmune system gene families have evolved by gene duplication, interlocus recombination and (in some cases) positive Darwinian selection favoring diversity at the amino acid level. (shrink)

Dawkins in his The Selfish Gene(1) quite aptly applies the term “selfish” to parasitic repetitive DNA sequences endemic to eukaryotic genomes, especially vertebrates. Doolittle and Sapienza(2) as well as Orgel and Crick(3) enlivened this notion of selfish DNA with the identification of such repetitive sequences as remnants of mobile elements such as transposons. In addition, Orgel and Crick(3) associated parasitic DNA with a potential to outgrow their host genomes by propagating both vertically via conventional genome replication as well as infectiously (...) by horizontal gene transfer (HGT) to other genomes. Still later, Doolittle(4) speculated that unchecked HGT between unrelated genomes so complicates phylogeny that the conventional representation of a tree of life would have to be replaced by a thicket or a web of life.(4) In contrast, considerable data now show that reconstructions based on whole genome sequences are consistent with the conventional “tree of life”.(5–10) Here, we identify natural barriers that protect modern genome populations from the inroads of rampant HGT. BioEssays 27:741–747, 2005. © 2005 Wiley Periodicals, Inc. (shrink)

In the last thirty years systematics has transformed itself from a discipline concerned with classification into a discipline concerned with reconstructing the evolutionary history of life. This transformation has been driven by cladistic analysis, a set of techniques for reconstructing evolutionary trees. Long interested in the large-scale structure of evolutionary history, cladistically oriented systematists have recently begun to apply "tree thinking" to problems near the species level. ¶ In any local ("non-dimensional") situation species are usually well-defined, but across space and (...) time the grouping of populations into species is often problematic. Three views of species are in common use today, the biological species concept, the evolutionary species concept, and the phylogenetic species concept. Each of these has strengths and weaknesses, but no matter which is applied, exact counts of the number of species in any extended area will always be ambiguous no matter how much factual information is available. This ambiguity arises because evolution is an historical process, and the grouping of organisms into species always depends to some extent upon expectations of the future behavior of those organisms and their descendants, expectations that cannot be evaluated in the present. The existence and special character of the species problem is itself one of the central pieces of evidence for evolution. (shrink)

Phylogenetics is the study and reconstruction of evolutionary history and is filled with numerous foundational issues of interest to philosophers. This paper briefly introduces some central concepts in the field, describes some of the main methods for inferring phylogenies, and provides some arguments for the superiority of model-based methods such as Likelihood and Bayesian methods over nonparametric methods such as parsimony. It also raises some underdeveloped issues in the field of interest to philosophers.

From the point of view of a dynamic morphology, form is not only the result of process(es) — it is process. This process may be analyzed in terms of two pairs of fundamental processes: growth and decay, differentiation and dedifferentiation. Each of these processes can be analyzed in terms of various modalities (parameters) and submodalities. This paper deals with those of growth (see Table 1). For the purpose of systematits and phylogenetic reconstruction the modalities and submodalities can be considered (...) dynamic characters that have states. Each state of such a dynamic character is a more detailed process, hence not static. For example, determinate growth represents a state of the dynamic character (or modality) of growth duration.The processes of Table 1 can be applied to the whole plant kingdom (although in certain cases only some processes of the whole set may be applicable). Thus, the diversity of plant form is seen as a diversity of process combinations. From this point of view, change in form implies change in the process combination(s). Questions that arise are, for example, the following: Which process combinations actually occur? Which of these are the most frequent? How and why have process combinations changed during ontogeny and phylogeny? (shrink)

The present work analyzes the controversy within biological systematics regarding the status of cladistics. The use of the parsimony method for phylogenetic reconstruction has been defended by appealing to a methodological principle of simplicity, as well as to empirical principles that external to systematics. I propose new kind of approach, which consists in considering it an empirical theory, thus justifying its application by its empirical success. To defend this point, a formal structuralist reconstruction of cladistics will be provided, (...) which will help clarify what its central empirical statements are. At the same time, I show how these principles have been used to test that theory using experimental phylogenies. (shrink)

RubisCO is Earth's main enzyme responsible for CO2 fixation via carboxylation of ribulose-1,5-bisphosphate into organic matter. Besides the carboxylation reaction, RubisCO also catalyzes the oxygenation of RuBP by O2, which is probably as old as its carboxylation properties. Based on molecular phylogeny, the occurrence of the reactive oxygen species -removing system and kinetic properties of different RubisCO forms, we postulated that RubisCO oxygenase activity appeared in local microoxic areas, yet before the appearance of oxygenic photosynthesis. Here, in reviewing the (...) literature, we present a novel hypothesis: the RubisCO early oxygenase activity hypothesis. This hypothesis may be compared with the exaptation hypothesis, according to which latent RubisCO oxygenase properties emerged later during the oxygenation of the Earth's atmosphere. The reconstruction of ancestral RubisCO forms using ancestral sequence reconstruction techniques, as a promising way for testing of RubisCO early oxygenase activity hypothesis, is presented. RubisCO catalyzes carboxylation and oxygenation of ribulose-1,5-bisphosphate. Carboxylating properties of RubisCO are an ancient way for CO2 fixation into organic matter. However, an evolutionary importance of oxygenase activity of RubisCO remains enigmatic. Here, we formulated RubisCO early oxygenase activity hypothesis. According to this hypothesis, RubisCO oxygenase activity appeared yet before the appearance of oxygenic photosynthesis. (shrink)

The canonical view of a 3‐domain (3D) tree of life was recently challenged by the discovery of Asgardarchaeota encoding eukaryote signature proteins (ESPs), which were treated as missing links of a 2‐domain (2D) tree. Here we revisit the debate. We discuss methodological limitations of building trees with alignment‐dependent approaches, which often fail to satisfactorily address the problem of ‘‘gaps.’’ In addition, most phylogenies are reconstructed unrooted, neglecting the power of direct rooting methods. Alignment‐free methodologies lift most difficulties but require employing (...) realistic evolutionary models. We argue that the discoveries of Asgards and ESPs, by themselves, do not rule out the 3D tree, which is strongly supported by comparative and evolutionary genomic analyses and vast genomic and biochemical superkingdom distinctions. Given uncertainties of retrodiction and interpretation difficulties, we conclude that the 3D view has not been falsified but instead has been strengthened by genomic analyses. In turn, the objections to the 2D model have not been lifted. The debate remains open. Also see the video abstract here:https://youtu.be/-6TBN0bubI8. (shrink)

In this paper, I study the application of phylogenetic analysis in evolutionary archaeology. I show how transfer of this apparently general analytic tool is affected by salient differences in disciplinary context. One is that archaeologists, unlike many biologists, do not regard cladistics as a tool for classification, but are primarily interested in explanation. The other is that explanation is traditionally sought in terms of individual-level rather than population-level mechanisms. The latter disciplinary difference creates an ambiguity in the application and interpretation (...) of phylogenetic analyses. Moreover, I argue that, while archaeologists have claimed that “cladistics is useful for reconstructing artefact phylogenies”, these reconstructions only contribute minimally to the explanatory research agenda of evolutionary archaeology. (shrink)

William Whewell (1794–1866), polymathic Victorian scientist, philosopher, historian, and educator, was one of the great neologists of the nineteenth century. Although Whewell's name is little remembered today except by professional historians and philosophers of science, researchers in many scientific fields work each day in a world that Whewell named. "Miocene" and "Pliocene," "uniformitarian" and "catastrophist," "anode" and "cathode," even the word "scientist" itself—all of these were Whewell coinages. Whewell is particularly important to students of the historical sciences for another word (...) he coined, one that was unfortunately not as successful as many of his others because it is difficult to pronounce. This word, "palaetiology," was the name Whewell gave to the class of sciences that are concerned with historical causation: the class we might today refer to as historical sciences. Although the disciplines Whewell included under the heading of palaetiology might seem to cut across conventional academic boundaries of his day and ours—his exemplary palaetiological sciences were geology and comparative philology—all these fields may be examined together, Whewell argued, because of their common interest in reconstructing the past. ¶ This paper is an essay in the palaetiological sciences, dedicated to Whewell on the bicentennial of his birth, an essay that examines some of the principles, maxims, and rules of procedure that these sciences have all in common. Its first purpose is to demonstrate the continuing validity of Whewell's classification of these sciences through a study of historical representation in three different palaetiological fields: systematics, historical linguistics, and textual transmission. Its second purpose is to continue the development of an extended analogy between historical representation and cartographic representation that I began in an earlier paper (O'Hara, 1993, Systematic Biology), an analogy that makes especially clear the common representational practices that are found throughout palaetiology. (shrink)

Certain lines of reasoning common in evolutionary taxonomy have been termed viciously circular. They are quite obviously not logically circular. They do give the superficial appearance of epistemological circularity. This appearance arises from the method of successive approximation used by evolutionary taxonomists. It is argued that this method is not epistemologically circular, even when the only evidence that the taxonomist has to go on is the phenetic similarity of contemporary forms. The important criticism of evolutionary taxonomy is rather that in (...) the absence of fossil evidence phyletic reconstructions are not warranted. It is argued that this charge stems initially from a misunderstanding of the kind of certainty possible in empirical science. When this criticism is couched in appropriate terms, it may be seen to have some force. Many phyletic inferences are not as warranted as one might wish. However, there is a great deal of difference between arguing that a line of reasoning is unwarranted and arguing that it is viciously circular. (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)

Phylogenetic trees are a crucial backbone for a wide breadth of biological research spanning systematics, organismal biology, ecology, and medicine. In 2015, the Open Tree of Life project published a first draft of a comprehensive tree of life, summarizing digitally available taxonomic and phylogenetic knowledge. This paper reviews, investigates, and addresses the following questions as a follow-up to that paper, from the perspective of researchers involved in building this summary of the tree of life: Is there a tree of life (...) and should we reconstruct it? Is available data sufficient to reconstruct the tree of life? Do we have access to phylogenetic inferences in usable form? Can we combine different phylogenetic estimates across the tree of life? And finally, what is the future of understanding the tree of life? A schematic of the process of building a unified tree of life. Inputs are published phylogenetic estimates and a unified taxonomy. These inputs are combined into a summary supertree containing ≈2 million taxa. The relationships among any subset of taxa may easily be accessed and used in downstream analyses. (shrink)