Classification or Phylogenetic Estimates?

[m]3ta is a method that seeks to implement a taxic view of homology. The method is consistent with Patterson's tests for discriminating homology from nonhomology. Contrary to the claims of Kluge and Farris, (1999, Cladistics 15, 205–212), m3ta is not a phenetic method—nor does it necessarily place the basal split in a tree between the phenetically most divergent taxa. [m]3ta does not seek to accurately recover phylogeny but rather it seeks to maximize the information content of taxic homology propositions. [m]3ta is a method of classification in which the unit of analysis is the relation of homology. [m]3ta differs from all phylogenetic methods because the units of analyses in phylogenetic methods, including sca, are transformation series.     

A/The Brief History of Three-Taxon Analysis

Recent claims by its advocates notwithstanding, three-taxon analysis (3ta) provides no method for recognizing reversals or for applying them as apomorphies. Accordingly, 3ta could be used as a phylogenetic method only under an assumption of irreversibility. Being a method for calculating trees from character data, 3ta is not connected to any particular rule (“interpretation”) for selecting resolutions of consensus trees considered as abstract diagrams. 3ta cannot be justified simply by invoking a general minimization principle such as Occam's Razor, since that would cover almost any method. Some more specific basis is needed, and consideration of proposed bases for 3ta shows that none is even remotely adequate.     

Taxic and Transformational Homology: Different Ways of Seeing

Hypotheses of taxic homology are hypotheses of taxa (groups). Hypotheses of transformational homology are hypotheses of transformations between character states within the context of an explicit model of character evolution. Taxic and transformational homology are discussed with respect to secondary loss and reversal in the context of three-taxon statement analysis and standard cladistic analysis. We argue that it is important to distinguish complement relation homologies from those that we term paired homologues. This distinction means that the implementation of three-taxon statement analysis needs modification if all data are to be considered potentially informative. Modified three-taxon statement analysis and standard cladistic analysis yield different results for the example of character reversal provided by Kluge (1994) for both complement relation data and paired homologues. We argue that these different results reflect the different approaches of standard cladistic analysis and modified t.t.s. analysis. In the standard cladistic approach, absence, as secondary loss, can provide evidence for a group. This is because the standard cladistic approach implements a transformational view of homology. In the t.t.s approach discussed in this paper, absence can only be interpreted as secondary loss by congruence with other data; absence alone can never provide evidence for a group. In this respect, the modified t.t.s. approach is compatible with a taxic view of homology.     

Cycles

Intended to support three-taxon analysis (3ta), the proposal that all character states be regarded as terminal would instead undercut that method. The same is true of the idea that cladistic methods should not account for plesiomorphies. Parsimony does not correspond to interpretation 1 for incompletely resolved cladograms. The main argument common to Nelson's (1996) and Nelson and Platnick's (1991) advocacy of three-taxon analysis rests on presupposing its conclusion. While suggesting that parsimony rests on an inferior evolutionary model, Nelson (1996) neither offered nor provided evidence for any alternative. 3ta sometimes favors reversal over parallelism, but in other cases may disregard reversed characters, so that the method seems to lack any coherent theoretical basis.     

It is normal for classification approaches to be diverse

It is asserted that the postmodern concept of science, unlike the classical ideal, presumes necessary existence of various classification approaches (schools) in taxonomy, each corresponding to a particular aspect of consideration of the "taxic reality". They are set up by diversity of initial epistemological and ontological backgrounds which fix in a certain way a) fragments of that reality allowable for investigation, and b) allowable methods of exploration of the fragments being fixed. It makes it possible to define a taxonomic school as a unity of the above backgrounds together with consideration aspect delimited by them. Two extreme positions of these backgrounds could be recognized in recent taxonomic thought. One of them follows the scholastic tradition of elaboration of a formal and, hence, universal classificatory method ("new typology", numerical phenetics, pattern cladistics). Another one asserts dependence of classificatory approach on the judgment of the nature of taxic reality (natural philosophy, evolutionary schools of taxonomy). Some arguments are put forward in favor of significant impact of evolutionary thinking onto the theory of modern taxonomy. This impact is manifested by the correspondence principle which makes classificatory algorithms (and hence resulting classifications) depending onto initial assumptions about causes of taxic diversity. It is asserted that criteria of "quality" of both classifications proper and classificatory methods can be correctly formulated within the framework of a particular consideration aspect only. For any group of organisms, several particular classifications are rightful to exist, each corresponding to a particular consideration aspect. These classifications could not be arranged along the "better-worse" scale, as they reflect different fragments of the taxic reality. Their mutual interpretation depends on degree of compatibility of background assumptions and of the tasks being resolved. Extensionally, classifications are compatible as much as they coincide by context and hierarchical structure of included taxa. Intentionally, typological classifications are compatible if included taxa are comparable by their diagnoses, while phylogenetic classifications are compatible if the included taxa are ascribed monophyletic status. A brief consideration is given to the "new phylogenetics" (= "genophyletics") as to a classificatory approach aimed at elaboration of parsimonious phylogenetic hypotheses based on molecular biology data and employing numerical methods of cladistic analysis. This approach is shown to borrows some phenetic ideas and revives scholastic principle of unified classificatory basis. It is supposed that, in a time, biological classification would get escaping from plethora of positivistic ideas (including those being developed by nowaday cladistics) and would assimilate (revive) more actively holistic worldview.     

Homology and logical fallacy

The meaning of homology, is analyzed in a phylogenetic context. The notion of homology is unequivocal only if tied to recency of common ancestry, i.e. to the concept of monophyly. Processual analysis of the biological causes of similarity cannot provide guidance in the search for relations of homology. Instead, it is the reconstruction of the taxic hierarchy based on homology and congruence which provides guidance in the search for underlying causes of similarity. Pattern reconstruction is shown to have logical precedence over process explanations.     

Recycled

Both three-taxon analysis (3ta) and conventional parsimony analysis (CPA) fall within the cladistic framework. Attempts to exclude 3ta from the general cladistic framework so far seem to amount to declaring CPA as the only permissible analytic technique within cladistics. Critics of 3ta have failed to fully implement it in examples; as a result this criticism is faulty and does not support the claims made. Ultimately, the relative merit of 3ta will be resolved empirically, by comparison of classifications produced from it with classifications using other methods.     

Taxic Revisions

Parsimony analysis provides a straightforward way of assessing homology on a tree: a state shared by two terminals comprises homologous similarity if optimization attributes that state to all the stem species lying between those terminals. Three-taxon statements (3ts), although seemingly "exact" in that each either fits a tree or does not, do not provide a satisfactory assessment of homology, because that assessment can be internally contradictory and because 3ts systematically exclude homologous resemblance in reversed states. Modified 3ts analysis (m3ta), a method in which both plesiomorphic and apomorphic states of "paired homologue" (PH) characters (those other than presence/absence data) are regarded as "informative" (able to distinguish groups), can (obviously) group by symplesiomorphy and so form paraphyletic groups unless data are clocklike enough. Patterson's pattern analysis (ppa) has the same shortcoming, to which it adds the drawback that only characters fitting the tree perfectly are used, a restriction that can easily lead to discarding most of the structure in the data. Revised m3ta (rm3ta), a method in which plesiomorphic states are not taken as informative, can also form paraphyletic groups, because it cannot apply reversals as apomorphies. The idea that knowledge of phylogeny has been derived from classifications does not imply that nonevolutionary methods should be employed for classification, but instead means that systematic methods must be logically capable of phylogenetic interpretation. Neither m3ta nor rm3ta satisfies that requirement because of their contradictory assessments of homology.     

Phylogeny of the superfamily Gelechioidea (Lepidoptera: Obtectomera), with an exploratory application on geometric morphometrics

The superfamily Gelechioidea (Lepidoptera: Obtectomera) has a high species diversity. It consists of more than 18,400 described species and has a global distribution. Among it, large numbers of species were reported to be economically important to people's production and life. However, relationships among families or subfamilies in Gelechioidea have been exceptionally difficult to resolve using morphology or single gene genealogies. Multiple gene genealogies had been used in the molecular phylogenetic studies on Gelechioidea during the past years, but their phylogenetic relationships remain to be controversial mainly due to their limited taxa sampling relative to such high species diversity. In this paper, 89 ingroup species representing 55 genera are sequenced and added to the data downloaded from GenBank, and six species representing four closely related superfamilies are chosen as outgroup. The molecular phylogeny of Gelechioidea is reconstructed based on the concatenated data set composed of one mitochondrial marker (COI) and seven nuclear markers (CAD, EF-1ɑ, GAPDH, IDH, MDH, RpS5, wingless). The phylogenetic results, taking into consideration of the comparative morphological study, show that the clade of Gelechioidea is strongly supported and separated from other superfamilies, which further proves its monophyly. Five families are newly defined: Autostichidae sensu nov., Depressariidae sensu nov., Peleopodidae sensu nov., Ashinagidae sensu nov. and Epimarptidae sensu nov. Meanwhile, a monophyletic “SSABM” clade considered to be closely related is proposed for the first time, consisting of Stathmopodidae, Scythrididae, Ashinagidae, Blastobasidae and Momphidae. Moreover, geometric morphometric analyses using merged landmark data set from fore and hind wings of 118 representative species are conducted. The phenetic tree shows that the monophyly and phylogenetic relationships correspond with the results of molecular phylogeny largely, which well proves its importance and potential application in both phylogenetic reconstruction and species identification.     

PTP is Meaningless,T-PTP is Contradictory: A Reply to Trueman

Abstract — The T-PTP test for monophyly can attribute significance to entirely unsupported groups and even to both of two contradictory alternatives. The method of evaluating “support” after replacing selected groups of terminals with reconstructed ancestors has similar drawbacks. The proposed placement of Onychophora among Arthropoda is unsupported by 12S data, and strongly refuted by other evidence. Attempts to justify T-PTP on Popperian grounds rest entirely on misunderstanding Popper's ideas. The PTP test assesses neither Popperian corroboration nor statistical confidence of phylogenetic conclusions. It can attribute high significance to data that support no resolved grouping. Efforts to salvage PTP by proposing new interpretations share the weakness that none of the proposed interpretations generally holds. None of these methods seems useful in phylogenetic     

Evolutionary theory and systematics: relationships between process and patterns

The relevance of the Modern Evolutionary Synthesis to the foundations of taxonomy (the construction of groups, both taxa and phyla) is reexamined. The nondimensional biological species concept, and not the multidimensional, taxonomic, species notion which is based on it, represents a culmination of an evolutionary understanding. It demonstrates how established evolutionary mechanisms acting on populations of sexually reproducing organisms provide the testable ontological basis of the species category. We question the ontology and epistemology of the phylogenetic or evolutionary species concept, and find it to be a fundamentally untenable one. We argue that at best, the phylogenetic species is a taxonomic species notion which is not a theoretical concept, and therefore should not serve as foundation for taxonomic theory in general, phylogenetics, and macroevolutionary reconstruction in particular. Although both evolutionary systematists and cladists are phylogeneticists, the reconstruction of the history of life is fundamentally different in these two approaches. We maintain that all method, including taxonomic ones, must fall out of well corroborated theory. In the case of taxonomic methodology the theoretical base must be evolutionary. The axiomatic assumptions that all phena, living and fossil, must be holophyletic taxa (species, and above), resulting from splitting events, and subsequently that evaluation of evolutionary change must be based on a taxic perspective codified by the Hennig ian taxonomic species notion, are not testable premises. We discuss the relationship between some biologically, and therefore taxonomically, significant patterns in nature, and the process dependence of these patterns. Process-free establishment of deductively tested “genealogies” is a contradiction in terms; it is impossible to “recover” phylogenetic patterns without the investment of causal and processual explanations of characters to establish well tested taxonomic properties of these (such as homologies, apomorphies, synapomorphies, or transformation series). Phylogenies of either characters or of taxa are historical-narrative explanations (H-N Es), based on both inductively formulated hypotheses and tested against objective, empirical evidence. We further discuss why construction of a “genealogy”, the alleged framework for “evolutionary reconstruction”, based on a taxic, cladistic outgroup comparison and a posteriori weighting of characters is circular. We define how the procedure called null-group comparison leads to the noncircular testing of the taxonomic properties of characters against which the group phylogenies must be tested. This is the only valid rooting procedure for either character or taxon evolution. While the Hennig -principle is obviously a sound deduction from the theory of descent, cladistic reconstruction of evolutionary history itself lacks a valid methodology for testing transformation hypotheses of both characters and species. We discuss why the paleontological method is part of comparative biology with a critical time dimension ana why we believe that an “ontogenetic method” is not valid. In our view, a merger of exclusive (causal and interactive, but best described as levels of organization) and inclusive (classificatory) hierarchies has not been accomplished by a taxic scheme of evolution advocated by some. Transformational change by its very nature is not classifiable in an inclusive hierarchy, and therefore no classification can fully reflect the causal and interactive chains of events constituting phylogeny, without ignoring and contradicting large areas of corroborated evolutionary theory. Attempts to equate progressive evolutionary change with taxic schemes by Haeckel were fundamentally flawed. His ideas found 19th century expression in a taxic perception of the evolutionary process (“phylogenesis”), a merger of typology, hierarchic and taxic notions of progress, all rooted in an ontogenetic view of phylogeny. The modern schemes of genealogical hierarchies, based on punctuation and a notion of “species” individuality, have yet to demonstrate that they hold promise beyond the Haeckel ian view of progressive evolution.     

系统发育多样性测度及其在生物多样性保护中的应用

生物多样性保护面临两个基本问题：如何确定生物多样性测度以及如何保护生物多样性。传统的生物多样性测度是以物种概念为基础的,用生态学和地理学方法确定各种生物多样性指数。其测度依赖于样方面积的大小,并且所有的物种在分类上同等对待。系统发育多样性测度基于系统发育和遗传学的理论和方法,能确定某一物种对类群多样性的贡献大小。该方法比较复杂,只有在类群的系统发育或遗传资料比较齐全时方能应用。本文认为,物种生存力途径和系统发育多样性测度相结合有助于确定物种和生态系统保护的优先秩序。     

Myriapod monophyly and relationships among myriapod classes based on nearly complete 28S and 18S rDNA sequences

Myriapods play a pivotal position in the arthropod phylogenetic tree. The monophyly of Myriapoda and its internal relationships have been difficult to resolve. This study combined nearly complete 28S and 18S ribosomal RNA gene sequences (3,826 nt in total) to estimate the phylogenetic position of Myriapoda and phylogenetic relationships among four myriapod classes. Our data set consists of six new myriapod sequences and homologous sequences for 18 additional species available in GenBank. Among the six new myriapod sequences, those of the one pauropod and two symphylans are very important additions because they were such difficult taxa to classify in past molecular-phylogenetic studies. Phylogenetic trees were constructed with maximum parsimony, maximum likelihood, and Bayesian analyses. All methods yielded moderate to strong support for the monophyly of Myriapoda. Symphyla grouped strongly with Pauropoda under all analytical conditions. The KH test rejected the traditional view of Dignatha and Progoneata, and the topology obtained here, though not significantly supported, was Diplopoda versus ((Symphyla + Pauropoda) + Chilopoda).     

Parsimony analysis and ecological communities, phytosociology, nested subsets analysis, forest fragmentation: a reply to Giannini and Keller

We suggested using parsimony analysis to study community evolution in terms of species composition and to apply these results in the context of forest fragmentation as a replacement for the so‐called “nested subsets analysis” or other phenetic synecological or phytosociological methods ( Pellens et al., 2005 ). Giannini and Keller (2007 ) took issue with this new application on the basis of three misunderstandings. We re‐emphasize that communities themselves are analyzed, not landscape parts such as forest fragments. Therefore, it must be clear that communities are analogous to taxa and landscape parts such as fragments are analogous to distributions of taxa. Community evolution is the change in community composition by immigration, emigration and local extinction. Thus, gains and losses of species should not be confused with horizontal transfer. Parsimony analysis does not necessarily group communities based on shared absences of rare species. Rare species are not necessarily absent in the same communities and these absences are not necessarily inferred to be synapomorphies after rooting. This is the main advance expected when cladistics is used instead of the previously cited phenetic methods working with overall similarity. © The Willi Hennig Society 2007.     

Phylogenetic reconstruction and phenetic taxonomy

Cladistic analysis should not be equated with phylogenetic reconstruc Instead it is a means of describing character-state distributions among organisms and in this it resembles phenetic analysis. However, the claim that cladistic methods meet phenetic ('Gilmour-natural') criteria for classification as well as or better than traditional phenetic ones is shown to be based on an inadequate interpretation of these criteria. Instead, a new measure of naturalness is proposed in which the most natural classification is that which describes the distribution of all character states by the smallest number of statements. The possibility of extending this measure to provide a criterion for an optimally simple classification is noted. It is concluded that phylogenetic reconstruction must not only reflect the branching patterns suggested by cladistic analysis but also take account of the tionary history that is reflected in an optimal phenogram.     

Phylogeny of the sabertoothed felids (Carnivora: Felidae: Machairodontinae)

In recent years, advances in our understanding of feline relationships have cast light on their evolutionary history. In contrast, there have been no phylogenetic analyses on machairodont felids, making it difficult to develop an evolutionary hypothesis based on the recent surge of studies on their craniomandibular morphology and functional anatomy. In this paper, I provide the first phylogenetic hypothesis of machairodont relationships based on 50 craniomandibular and dental characters from a wide range of sabercats spanning more 11 Myr. Exact searches produced 19 most‐parsimonious trees, and a strict consensus was well resolved. The Machairodontinae comprise a number of basal taxa (Promegantereon, Machairodus, Nimravides, Dinofelis, Metailurus) and a well‐supported clade of primarily Plio‐Pleistocene taxa (Megantereon, Smilodon, Amphimachairodus, Homotherium, Xenosmilus) for which the name Eumachairodontia taxon novum is proposed. Previous phenetic grouping of machairodont taxa into three distinct groups, the Smilodontini, Homotherini and Metailurini, was not supported by cladistic parsimony analysis, and forcing monophyly of these groups was significantly incompatible with character distribution. Machairodonts as a clade are not characterized by saberteeth, i.e. hypertrophied, blade‐like upper canines, but by small lower canines, as well as small M¹; and large P³ parastyle. True saberteeth arose later and are a synapomorphy of the Eumachairodontia.     

Convergent and parallel evolution: a model illustrating selection, phylogeny and phenetic similarity

A model is proposed which considers the structural relationships of body characteristics and their role in a concept wherein the phylogenetic relationship of the organisms under study is interpreted as constituting degrees of convergent or parallel evolution. The model also accounts for the relationships between selection pressures, phylogeny, and phenetic expression. The phenomena of convergent and parallel evolution are based upon the observations of similar characteristics, the geometric concept, magnitude and similarity of selection pressures, and the phylogenetic relationship of the groups in question.     

Searching for natural lineages within the Cerylonid Series (Coleoptera: Cucujoidea)

Phylogenetic relationships within the diverse beetle superfamily Cucujoidea are poorly known. The Cerylonid Series (C.S.) is the largest of all proposed superfamilial cucujoid groups, comprising eight families and representing most of the known cucujoid species diversity. The monophyly of the C.S., however, has never been formally tested and the higher-level relationships among and within the constituent families remain equivocal. Here we present a phylogenetic study based on 18S and 28S rDNA for 16 outgroup taxa and 61 C.S. ingroup taxa, representing seven of the eight C.S. families and 20 of 39 subfamilies. We test the monophyly of the C.S., investigate the relationships among the C.S. families, and test the monophyly of the constituent families and subfamilies. Phylogenetic reconstruction of the combined data was achieved via standard static alignment parsimony analyses, Direct Optimization using parsimony, and partitioned Bayesian analysis. All three analyses support the paraphyly of Cucujoidea with respect to Tenebrionoidea and confirm the monophyly of the C.S. The C.S. families Bothrideridae, Cerylonidae, Discolomatidae, Coccinellidae and Corylophidae are supported as monophyletic in all analyses. Only the Bayesian analysis recovers a monophyletic Latridiidae. Endomychidae is recovered as polyphyletic in all analyses. Of the 14 subfamilies with multiple terminals in this study, 11 were supported as monophyletic. The corylophid subfamily Corylophinae and the coccinellid subfamilies Chilocorinae and Scymninae are recovered as paraphyletic. A sister grouping of Anamorphinae+Corylophidae is supported in all analyses. Other taxonomic implications are discussed in light of our results.     

Using genetic networks and homology to understand the evolution of phenotypic traits

Homology can have different meanings for different kinds of biologists. A phylogenetic view holds that homology, defined by common ancestry, is rigorously identified through phylogenetic analysis. Such homologies are taxic homologies (=synapomorphies). A second interpretation, "biological homology" emphasizes common ancestry through the continuity of genetic information underlying phenotypic traits, and is favored by some developmental geneticists. A third kind of homology, deep homology, was recently defined as "the sharing of the genetic regulatory apparatus used to build morphologically and phylogenetically disparate features." Here we explain the commonality among these three versions of homology. We argue that biological homology, as evidenced by a conserved gene regulatory network giving a trait its "essential identity" (a Character Identity Network or "ChIN") must also be a taxic homology. In cases where a phenotypic trait has been modified over the course of evolution such that homology (taxic) is obscured (e.g. jaws are modified gill arches), a shared underlying ChIN provides evidence of this transformation. Deep homologies, where molecular and cellular components of a phenotypic trait precede the trait itself (are phylogenetically deep relative to the trait), are also taxic homologies, undisguised. Deep homologies inspire particular interest for understanding the evolutionary assembly of phenotypic traits. Mapping these deeply homologous building blocks on a phylogeny reveals the sequential steps leading to the origin of phenotypic novelties. Finally, we discuss how new genomic technologies will revolutionize the comparative genomic study of non-model organisms in a phylogenetic context, necessary to understand the evolution of phenotypic traits.