The repugnant and the mature in phylogenetic inference: atemporal similarity and historical identity

The significance of “being similar” in the inference of species relationships is refuted once again (see also Hennig, 1966, Phylogenetic Systematics, Univ. of Illinois Press, Urbana, IL). Without merit is Rieppel and Kearney's (Biol. J. Linn. Soc., 2002, 75, 59–82) claim that submitting the relational property of topological similarity, their preferred definition of character, to falsifying tests of similarity benefits that kind of inference. Such a priori uses of similarity, in character analysis, are consistent with observational theory, where a character is defined intensionally in terms of immutable properties. However, the induced hypotheses that follow from this theory, not the deductive test that Rieppel and Kearney wanted, remain controversial, because their predictability is a consequence of circular reasoning, and their projectabality fails empirically from incongruent observation reports. Further, a category mistake is made when the abstract, similarity‐defined, group of organisms is reified, as a part of history. In addition, Rieppel and Kearney failed to provide a special theory for similarity, which renders similarity scientifically repugnant (Quine, 1969, Ontological Relativity and Other Essays, Columbia Univ. Press, New York). A return to Hennig's (1966) evolutionary concept of evidence, as transformation series, is urged, and from which a testable character hypothesis can be formulated. There is no one operation for determining character states in this system—it can be anything that leads to the testable hypothesis of synapomorphy, as an historical identity relation. Character compatibility and conjunction, but not similarity, provide a priori tests in phylogenetic character analysis. In turn, the phylogenetic system of inference leads to explanations of homology, as historical identities, which exemplifies the goal of achieving a mature state of historical knowledge (not of Quine, 1969). Such maturity obtains from attempts to falsify hypotheses of species relationships with severely tested evidence, not from induction of “the” observation statement that Rieppel and Kearney sought to justify their true belief in a hypothesis of relationships.     

The Evolution of Integration of Biological Systems: An Evolutionary Perspective through Studies on Cells, Tissues, and Organs

Biologists are integrating studies of morphology, development,physiology,and other disciplines in order to understand howspecies and lineages diversify and cope with their environments.An evolutionary perspective in such studies, including thoseof cells, tissues, and organs, is potentially useful for thestructure and analysis of such problems. Evolutionary biologyis the study of the history of evolution and the elucidationof its mechanisms. Comparative biology is the comparison ofa trait or traits in selected taxa, and may be, but need notbe, evolutionary in approach. A phylogenetic hypothesis is necessaryfor reconstruction of pattern in morphology, ecology, behavior,and other areas. Acquaintance with evolutionary and phylogeneticperspectives can guide selection of taxa for study and opennew approaches to analysis of data. Such an approach is notalways appropriate to problems in biology, but it could be utilizedbeneficially more frequently than is currently practiced. Studiesof cells, tissues, and organs may contribute to the constructionof new phylogenetic hypotheses and to analysis of patterns andmechanisms of change when pursued from an evolutionary perspective     

Phylogenetic frameworks: towards a firmer foundation for the comparative approach

The growing interest in using phylogenies to test evolutionary hypotheses has focused attention on the need for robust estimates of phylogenetic history. Whether specific branching structures are correct summaries of evolutionary history can be estimated only through the examination of congruence of many sets of characters. After consideration of practical and philosophical aspects of congruence, I conclude that taxonomic congruence (analysis of congruence of topologies produced from independent datasets) is preferable to character congruence (analysis of congruence between individual characters) for estimating accuracy of phylogenetic hypotheses. Existing methods for examining taxonomic congruence are discussed and the combinable components approach, when preceded by application of rigorous statistical manipulations (e.g. jackknifing or bootstrapping), found most appropriate. Implementation of the method of combinable components is described, and is demonstrated using published data for Menidia and Rana. The robust branching structure resulting from this analysis (a phylogenetic framework) contains those nodes (phylogenetic hypotheses) that are strongly supported by at least one dataset and are consistent with all others. This approach is the most appropriate/conservative for testing hypotheses about evolutionary history.     

Species concepts and phylogenetics

Concepts of species proposed within the phylogenetic paradigm arecritically reviewed. Most so called phylogenetic species concepts relyheavily on factors immaterial to phylogenetic hypotheses. Thus, theyhave limited empirical content and offer weak bases on which to makedecisions about real problems related to species. Any workable notion ofspecies relies on an explicit character analysis, rather than onabstract properties of lineages, narrative predications and speculationson tokogenetic relationships. Species only exist conjecturally, as thesmallest meaningful units for phylogenetic analysis, as based oncharacter evidence. Such an idea considers species to be conjecturesbased on similarity, that are subsequently subject to testing by theresults of analysis. Species, thus, are units of phylogenetic analysisin the same way as hypotheses of homology are units of comparablesimilarities, i.e. conjectures to be tested by congruence. Althoughmonophyly need not be demonstrated for species-level taxa, hypotheses ofrelationships are the only basis to refute species limits and guidenecessary rearrangements. The factor that leads to recognition ofspecies is similarity in observed traits. The concept of life cycle isintroduced as an important element in the discussion of species, as anefficient way to convey subsidiary notions of sexual dimorphism,polymorphism, polytypy and clusters of diagnosable semaphoronts. Thenotion of exemplars is used to expand the concept ofspecies-as-individual-organisms into a more generally usable concept.Species are therefore proposed for a diagnosable sample of(observed or inferred) life cycles represented by exemplars all of whichare hypothesized to attach to the same node in a cladogram, and whichare not structured into other similarly diagnosable clusters. Thisdefinition is character-based, potentially testable by reference to abranching diagram, and dispenses with reference to ancestor-descendantrelationships or regression into population concepts. It provides aworkable basis on which to proceed with phylogenetic analysis and abasis for that analysis to refute or refine species limits. A protocolis offered for testing hypotheses of species boundaries in cladograms.     

Miocene radiation of deep-sea hydrothermal vent shrimp (Caridea: Bresiliidae): evidence from mitochondrial cytochrome oxidase subunit I

The evolutionary history of deep-sea shrimp (Caridea: Bresiliidae) inhabiting deep-sea hydrothermal vent and hydrocarbon seep environments was assessed using the mitochondrial Cytochrome c Oxidase subunit I (COI) gene (600 bp). Phylogenetic analyses (parsimony, likelihood, and neighbor-joining) recovered three distinct clades (A, Rimicaris/Chorocaris/Opaepele; B, Alvinocaris; and C, Mirocaris) consistent with higher level taxonomy based on morphology. However, robust phylogenetic results suggested that Chorocaris is paraphyletic and that Mirocaris fortunata and M. keldyshi may not be genetically distinct. A Kishino-Hasegawa likelihood approach was used to test alternative phylogenetic hypotheses based on biogeography and morphology. Evolutionary relationships of vent-endemic shrimp species did not appear to be correlated either with their extant biogeographic distribution or with the history of sea floor spreading. Additionally, COI data suggested that these vent-endemic organisms are not remnants of a Mesozoic vent assemblage; instead, they radiated in the Miocene.     

Hypothesis testing in biogeography

Often, biogeography is applied only as a narrative addition to phylogenetic studies and lacks scientific rigour. However, if research questions are framed as hypotheses, biogeographical scenarios become testable. In this review, we explain some problems with narrative biogeography and show how the use of explicit hypotheses is changing understanding of how organisms came to be distributed as they are. Developing synergies between biogeography, ecology, molecular dating and palaeontology are providing novel data and hypothesis-testing opportunities. New approaches are challenging the classic 'Gondwana' paradigm and a more complicated history of the Southern Hemisphere is emerging, involving not only general drivers such as continental drift and niche conservatism, but also drowning and re-emergence of landmasses, biotic turnover and long-distance colonization.     

A maximum likelihood approach to generate hypotheses on the evolution and historical biogeography in the Lower Volga Valley regions (southwest Russia)

The evolution of the diverse flora in the Lower Volga Valley (LVV) (southwest Russia) is complex due to the composite geomorphology and tectonic history of the Caspian Sea and adjacent areas. In the absence of phylogenetic studies and temporal information, we implemented a maximum likelihood (ML) approach and stochastic character mapping reconstruction aiming at recovering historical signals from species occurrence data. A taxon-area matrix of 13 floristic areas and 1018 extant species was constructed and analyzed with RAxML and Mesquite. Additionally, we simulated scenarios with numbers of hypothetical extinct taxa from an unknown palaeoflora that occupied the areas before the dramatic transgression and regression events that have occurred from the Pleistocene to the present day. The flora occurring strictly along the river valley and delta appear to be younger than that of adjacent steppes and desert-like regions, regardless of the chronology of transgression and regression events that led to the geomorphological formation of the LVV. This result is also supported when hypothetical extinct taxa are included in the analyses. The history of each species was inferred by using a stochastic character mapping reconstruction method as implemented in Mesquite. Individual histories appear to be independent from one another and have been shaped by repeated dispersal and extinction events. These reconstructions provide testable hypotheses for more in-depth investigations of their population structure and dynamics.     

Phylogenetic analysis, structural evolution and functional divergence of the 12-oxo-phytodienoate acid reductase gene family in plants

Background

Whenever different data sets arrive at conflicting phylogenetic hypotheses, only testable causal explanations of sources of errors in at least one of the data sets allow us to critically choose among the conflicting hypotheses of relationships. The large (28S) and small (18S) subunit rRNAs are among the most popular markers for studies of deep phylogenies. However, some nodes supported by this data are suspected of being artifacts caused by peculiarities of the evolution of these molecules. Arthropod phylogeny is an especially controversial subject dotted with conflicting hypotheses which are dependent on data set and method of reconstruction. We assume that phylogenetic analyses based on these genes can be improved further i) by enlarging the taxon sample and ii) employing more realistic models of sequence evolution incorporating non-stationary substitution processes and iii) considering covariation and pairing of sites in rRNA-genes.

Results

We analyzed a large set of arthropod sequences, applied new tools for quality control of data prior to tree reconstruction, and increased the biological realism of substitution models. Although the split-decomposition network indicated a high noise content in the data set, our measures were able to both improve the analyses and give causal explanations for some incongruities mentioned from analyses of rRNA sequences. However, misleading effects did not completely disappear.

Conclusion

Analyses of data sets that result in ambiguous phylogenetic hypotheses demand for methods, which do not only filter stochastic noise, but likewise allow to differentiate phylogenetic signal from systematic biases. Such methods can only rely on our findings regarding the evolution of the analyzed data. Analyses on independent data sets then are crucial to test the plausibility of the results. Our approach can easily be extended to genomic data, as well, whereby layers of quality assessment are set up applicable to phylogenetic reconstructions in general.     

The phylogenetic placement of Chondrichthyes: inferences from analysis of multiple genes and implications for comparative studies

Elasmobranch fishes (sharks and rays) have proven valuable for inferring general and specific properties of molecular evolution through comparative studies with crown group vertebrates because they are the most ancient group of gnathostomes. Recent studies have questioned the conventional phylogenetic placement of sharks in the vertebrate tree, however. In this paper I review the importance of the basal position of Chondrichthyes for comparative biology and compile evidence from multiple, independent genes to evaluate the phylogenetic placement of sharks. The results suggests that alternative phylogenetic hypotheses of the relationships among the Chondrichthyes, Actinopterygii and Sarcopterygii can not be refuted with available data, implying that the assumption of the basal placement of sharks in the vertebrate tree is suspect. Resolving the phylogeny of basal vertebrates is important for testing hypotheses about the evolution of vertebrates, and the current lack of a robust phylogeny limits evolutionary inferences that can be gained from comparative studies that include sharks and rays.     

The evolution of plant development

The last decade has witnessed a resurgence in the study of the evolution of plant development, combining investigations in systematics, developmental morphology, molecular developmental genetics, and molecular evolution. The integration of phylogenetic studies, structural analyses of fossil and extant taxa, and molecular developmental genetic information allows the formulation of explicit and testable hypotheses for the evolution of morphological characters. These comprehensive approaches provide opportunities to dissect the evolution of major developmental transitions among land plants, including those associated with apical meristems, the origins of the root/shoot dichotomy, diversification of leaves, and origin and subsequent modification of flower structure. The evolution of these major developmental innovations is discussed within both phylogenetic and molecular genetic contexts. We conclude that it is the combination of these approaches that will lead to the greatest understanding of the evolution of plant development.     

Modèle de mélange et analyse comparative des traits démographiques des poissons: Mixture models and comparative analysis of life history traits of Teleost fishes

The comparative approach in biology is used to investigate the attributes of actual species. Differences between them are used to formulate adaptive hypotheses of life history, morphology, demography, etc. Difficulties arise from the phylogenetic heritage: species are not independent and neglecting this fact can lead to inaccurate results. Fishing can have important effects but usually they are mixed with other variations such as environmental ones. Single species studies cannot succeed completely in analysing the effect of fishing. To disentangle them from orher fluctuations, we are led to the use of a comparative approach. Thus, we need to account for phylogenetic links between populations. In this paper we propose a new family of models, which allows us to split life history traits into parts.     

Phylogenetic studies of insect-plant interactions: Insights into the genesis of diversity

Phylogenetic analyses (and related historical evidence) can be used to test hypotheses about the oppurtinity for coevolution among plants and insect herbivores, the role of plant chemistry in mediating host shifts, the reality of coevolutionary 'arms races', and the role of novel defensive or counterdefensive characteristics in enhancing rates of diversification. Recent analyses provide evidence on each of these hypotheses. The strong phylogenetic component in many insect-plant associations suggests that host shifts by insects are often strongly constrained, that insect diets may not be 'optimal', and that the trophic structure and species diversity of local communities strongly reflect evolutionary history.     

Homoplasy, character function, and nemertean systematics

We question recent claims that cladistic analysis is inapplicable in nemerteans (phylum Nemertea) due to a supposedly high degree of convergence. We further argue that terms like convergence and parallelism are historical sayings and only make sense in a phylogenetic context. Therefore, an approach aiming to produce phylogenetic hypotheses cannot be rejected on the grounds of a high degree of convergence before the actual hypothesis. Convergence is not an empirical observation, but a conclusion made after an analysis. We also discuss the view that knowledge of a character's function is a prerequisite for phylogenetic analysis and conclude that this is an invalid approach. Function, like any other way of sharpening our observations, helps in formulating non-phylogenetic hypotheses of homology, but the crucial test is congruence with other characters on a phylogeny.     

Perspectives on the ecomorphology of bony fishes

Synopsis The field of ecomorphology has a long history with early roots in Europe. In this half of the century the application of ecomorphology to the biology of fishes has developed in the former Soviet Union, Poland and Czechoslovakia, The Netherlands, and in North America. While the specific approaches vary among countries, many North American studies begin by comparing morphological variation with variation in ecological characteristics at the intra or interspecific levels. These initial correlative studies form the ground work for hypotheses that explore the mechanistic underpinnings of the observed ecomorphological associations. Supporting these mechanistic hypotheses are insights from functional studies which demonstrate the limits to potential resource use resulting from a particular morphology; however, the actual resource use is likely to be more limited due to additional constraints provided by internal (e.g., behavior, physiology) and external (e.g., resource abundance, predator distribution) factors. The results from performance studies in the laboratory or field can be used to test specific ecomorphological hypotheses developed from the initial correlational and functional studies. Such studies may, but rarely do, incorporate an ontogenetic analysis of the ecomorphological association to determine their effect on performance. Finally, input from phylogenetic analyses allow an investigator to examine the evolution of specific features and to assess the rates and directionality of character evolution. The structural and ecological diversity of fishes provides a fertile ground to investigate these interactions. The contributions in this volume highlight some of the specific directions for ecomorphological research covering a variety of biological processes in fishes. These include foraging, locomotion, reproduction, respiration, and sensory systems. Running through these papers are new insights into universal ecomorphological issues, i.e., the relationships between form and ecological role and the factors that modify these relationships.     

Phylogenetic meta-analysis

Meta-analysis is a powerful statistical technique that combines the results of independent studies to identify general trends. When the species under examination are not independent however, it is also necessary to incorporate phylogenetic information into the analysis. Unfortunately, current meta-analytic approaches cannot account for lack of independence resulting from shared evolutionary history, so a general solution to this problem is lacking. In this article, I derive a model for phylogenetic meta-analysis, so that data across studies may be summarized with evolutionary history explicitly incorporated. The approach takes advantage of common aspects of linear statistical models used by both meta-analysis and the phylogenetic comparative method, thereby allowing them to be analytically combined. In this manner, the correlation structure generated by phylogenetic history can be incorporated directly into the meta-analytic procedure. I illustrate the approach by examining the prevalence of body size clines in mammals. The approach is general, and can also be used to incorporate correlation structure among studies generated by other factors, such as spatial or temporal proximity, or environmental similarity. Therefore, this procedure provides a general statistical template for meta-analytic techniques that can account for attributes that generate nonindependence among studies. Implications of the phylogenetic meta-analysis are discussed.     

The Use of Functional and Adaptive Criteria in Phylogenetic Systematics

SYNOPSIS. The controversy over whether functional data can contributeto phylogenetic inference has grown in recent years. Steps canbe taken toward its resolution if the relevance of functionaldata is judged for each component of phylogenetic analysis.These components are (1)recognizing of basic taxa (species orsupraspecific taxa), (2) formulating hypotheses of homologyfollowed by character analysis, (3) evaluating character phylogeny,(4) formulating phylogenetic hypotheses, and (5) evaluatingalternative phylogenetic hypotheses. It can be shown that functionaldata do not play a necessary or unique role in any of thesecomponents of phylogenetic analysis. Arguments to the contraryhave failed to provide a rigorous, repeatable methodto incorporatefunctional data; proponents of a functional approach to phylogeneticreconstruction rely too often on subjective, authoritarian argumentation. Students of functional evolutionary morphology frequently havefailed to understand the kinds of information necessary to studyor apply the causal process of adaptation via natural selection.This information, required by the very nature of the theoryitself, includes knowing the pattern of heredity of the phenotypiccharacters being studied, relating intrapopulational phenotypicvariability to variation in fitness, and knowing a sufficientamount about population structure to specify the componentsof natural selection. Studies within functional evolutionarymorphology are not designed to satisfy these requirements. Functionalevolutionary morphology uses the concepts of adaptation andnatural selection axiomatically, and thus such studies contributenothing to our understanding of the evolutionary process becausehypotheses about that process are not being evaluated. Thisalso suggests that, if functional evolutionary morphology wishesto engage in analyses of the evolutionary dynamics of the phenotype,a reorientation of its research strategy and goals will be necessary.     

Developmental constraints in a comparative framework: a test case using variations in phalanx number during amniote evolution

Constraints are factors that limit evolutionary change. A subset of constraints is developmental, and acts during embryonic development. There is some uncertainty about how to define developmental constraints, and how to formulate them as testable hypotheses. Furthermore, concepts such as constraint-breaking, universal constraints, and forbidden morphologies present some conceptual difficulties. One of our aims is to clarify these issues. After briefly reviewing current classifications of constraint, we define developmental constraints as those affecting morphogenetic processes in ontogeny. They may be generative or selective, although a clear distinction cannot always be drawn. We support the idea that statements about constraints are in fact statements about the relative frequency of particular transformations (where 'transformation' indicates a change from the ancestral condition). An important consequence of this is that the same transformation may be constrained in one developmental or phylogenetic context, but evolutionarily plastic in another. In this paper, we analyse developmental constraints within a phylogenetic framework, building on similar work by previous authors. Our approach is based on the following assumptions from the literature: (1) constraints are identified when there is a discrepancy between the observed frequency of a transformation, and its expected frequency; (2) the 'expected' distribution is derived by examining the phylogenetic distribution of the transformation and its associated selection pressures. Thus, by looking for congruence between these various phylogenetic distribution patterns, we can test hypotheses about constraint. We critically examine this approach using a test case: variation in phalanx-number in the amniote limb.     

Intercontinental community convergence of ecology and morphology in desert lizards

Evolutionary ecologists have long debated the extent to which communities in similar environments but different geographic regions exhibit convergence. On the one hand, if species' adaptations and community structure are determined by environmental features, convergence would be expected. However, if historical contingencies have long-lasting effects convergence would be unlikely. Most studies to date have emphasized the differences between communities in similar environments and little quantitative evidence for convergence exists. The application of comparative phylogenetic methods to ecological studies provides an opportunity to further investigate hypotheses of convergence. We compared the evolutionary patterns of structural ecology and morphology of 42 species of iguanian lizards from deserts of Australia and North America. Using a comparative approach, we found that evolutionary convergence of ecology and morphology occurs both in overall, community-wide patterns and in terms of pairs of highly similar intercontinental pairs of species. This result indicates that in these desert lizards, deterministic adaptive evolution shapes community patterns and overrides the historical contingencies unique to particular lineages.     

Rejecting "the given" in systematics

How morphology and systematics come together through morphological analysis, homology hypotheses and phylogenetic analysis is a topic of continuing debate. Some contemporary approaches reject biological evaluation of morphological characters and fall back on an atheoretical and putatively objective (but, in fact, phenetic) approach that defers to the test of congruence for homology assessment. We note persistent trends toward an uncritical empiricism (where evidence is believed to be immediately “given” in putatively theory‐free observation) and instrumentalism (where hypotheses of primary homology become mere instruments with little or no empirical foundation for choosing among competing phylogenetic hypotheses). We suggest that this situation is partly a consequence of the fact that the test of congruence and the related concept of total evidence have been inappropriately tied to a Popperian philosophy in modern systematics. Total evidence is a classical principle of inductive inference and does not imply a deductive test of homology. The test of congruence by itself is based philosophically on a coherence theory of truth (coherentism in epistemology), which is unconcerned with empirical foundation. We therefore argue that coherence of character statements (congruence of characters) is a necessary, but not a sufficient, condition to support or refute hypotheses of homology or phylogenetic relationship. There should be at least some causal grounding for homology hypotheses beyond mere congruence. Such causal grounding may be achieved, for example, through empirical investigations of comparative anatomy, developmental biology, functional morphology and secondary structure. © The Willi Hennig Society 2006.     

Historical relationships among Neotropical lowland forest areas of endemism as determined by mitochondrial DNA sequence variation within the Wedge-billed Woodcreeper (Aves: Dendrocolaptidae: Glyphorynchus spirurus)

Studies of the distribution of South American taxa have identified several areas of endemism that may have contributed to the historical diversification of the region. We constructed a phylogeny of Glyphorynchus spirurus (Aves: Dendrocolaptidae) populations using mtDNA sequence data from portions of cytochrome b, NADH dehydrogenase subunit II (ND2), and complete NADH dehydrogenase subunit III (ND3). Using this phylogeny we evaluate five previous hypotheses of area-relationships, two based on phylogenetic studies of morphological characters in birds and three based on parsimony analysis of endemism in birds and primates. Maximum likelihood and maximum parsimony analyses recovered two phylogenetic hypotheses that differed in the placement of one of the areas. Within each of the areas of endemism, the two analyses support the same clades. Neither of the phylogenetic hypotheses for Glyphorynchus exactly matches any of the five previous hypotheses of area-relationships, although ambiguous support exists for one of them. Five areas-Central America, Inambari, Napo, Pará, and Rond?nia-are supported as composites with component taxa having phylogenetic affinities with more than one area. Data reported here also indicate high levels of sequence divergence within Glyphorynchus. Genetic breaks within Glyphorynchus are only partially congruent with subspecific taxonomy. The regional sampling design used makes this study the largest scale genetic assay of a widespread Neotropical avian taxon published to date.