Aphyly: A Systematic Designation for a Taxonomic Problem

A taxon is aphyletic when it is deemed to be non-monophyletic or unresolved, therefore aphyletic taxa are a taxonomic problem rather than an evolutionary anomaly. A problem arises in systematics when taxonomic names assigned to aphyletic taxa are treated as if they were natural groups. In the absence of a taxonomic and systematic revision, anomalous taxa should be labelled as aphyletic without recourse to phylogenetic inference (i.e., interpretation). Doing so avoids the validation of aphyletic names and the creation of dubious results in fields that rely on systematic and taxonomic data.     

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.     

Genotypic and phenotypic evaluation of Rutilus spp. from Skadar, Ohrid and Prespa Lakes supports revision of endemic as well as taxonomic status of several taxa

One mtDNA gene (cytochrome b), one nuclear DNA fragment, five microsatellites and a suite of morphological characters were evaluated in samples of Rutilus spp. from Skadar, Ohrid and Prespa Lakes. Both genetic and phenotypic data supported two sympatric taxa in Lake Skadar, whereby Prespa and Ohrid Lakes revealed only a single taxon each. One of the taxa from Lake Skadar was similar to samples from Lake Prespa, whereas the second taxon was the most divergent in the data set. The estimated time to the most recent common ancestor of these two sympatric taxa in Lake Skadar was between 125 000 and 500 000 years. The data did not support existing taxonomic schemes for Rutilus in these lakes. This study poses the following working hypothesis: (1) Rutilus prespensis lives both in Lake Prespa and Lake Skadar and therefore is not endemic to Lake Prespa, (2) Rutilus ohridanus lives in Lake Ohrid only and therefore could be considered an endemic if its species status is retained and (3) a third recently described taxon (Rutilus albus) sympatric to R. prespensis lives in Lake Skadar.     

Multiple data sets, high homoplasy, and the phylogeny of softshell turtles (Testudines: Trionychidae)

We present a phylogenetic hypothesis and novel, rank-free classification for all extant species of softshell turtles (Testudines:Trionychidae). Our data set included DNA sequence data from two mitochondrial protein-coding genes and a approximately 1-kb nuclear intron for 23 of 26 recognized species, and 59 previously published morphological characters for a complimentary set of 24 species. The combined data set provided complete taxonomic coverage for this globally distributed clade of turtles, with incomplete data for a few taxa. Although our taxonomic sampling is complete, most of the modern taxa are representatives of old and very divergent lineages. Thus, due to biological realities, our sampling consists of one or a few representatives of several ancient lineages across a relatively deep phylogenetic tree. Our analyses of the combined data set converge on a set of well-supported relationships, which is in accord with many aspects of traditional softshell systematics including the monophyly of the Cyclanorbinae and Trionychinae. However, our results conflict with other aspects of current taxonomy and indicate that most of the currently recognized tribes are not monophyletic. We use this strong estimate of the phylogeny of softshell turtles for two purposes: (1) as the basis for a novel rank-free classification, and (2) to retrospectively examine strategies for analyzing highly homoplasious mtDNA data in deep phylogenetic problems where increased taxon sampling is not an option. Weeded and weighted parsimony, and model-based techniques, generally improved the phylogenetic performance of highly homoplasious mtDNA sequences, but no single strategy completely mitigated the problems of associated with these highly homoplasious data. Many deep nodes in the softshell turtle phylogeny were confidently recovered only after the addition of largely nonhomoplasious data from the nuclear intron.     

Integration of morphological data sets for phylogenetic analysis of Amniota: the importance of integumentary characters and increased taxonomic sampling

Several mutually exclusive hypotheses have been advanced to explain the phylogenetic position of turtles among amniotes. Traditional morphology-based analyses place turtles among extinct anapsids (reptiles with a solid skull roof), whereas more recent studies of both morphological and molecular data support an origin of turtles from within Diapsida (reptiles with a doubly fenestrated skull roof). Evaluation of these conflicting hypotheses has been hampered by nonoverlapping taxonomic samples and the exclusion of significant taxa from published analyses. Furthermore, although data from soft tissues and anatomical systems such as the integument may be particularly relevant to this problem, they are often excluded from large-scale analyses of morphological systematics. Here, conflicting hypotheses of turtle relationships are tested by (1) combining published data into a supermatrix of morphological characters to address issues of character conflict and missing data; (2) increasing taxonomic sampling by more than doubling the number of operational taxonomic units to test internal relationships within suprageneric ingroup taxa; and (3) increasing character sampling by approximately 25% by adding new data on the osteology and histology of the integument, an anatomical system that has been historically underrepresented in morphological systematics. The morphological data set assembled here represents the largest yet compiled for Amniota. Reevaluation of character data from prior studies of amniote phylogeny favors the hypothesis that turtles indeed have diapsid affinities. Addition of new ingroup taxa alone leads to a decrease in overall phylogenetic resolution, indicating that existing characters used for amniote phylogeny are insufficient to explain the evolution of more highly nested taxa. Incorporation of new data from the soft and osseous components of the integument, however, helps resolve relationships among both basal and highly nested amniote taxa. Analysis of a data set compiled from published sources and data original to this study supports monophyly of Amniota, Synapsida, Reptilia, Parareptilia, Eureptilia, Eosuchia, Diapsida, Neodiapsida, Sauria, Lepidosauria, and Archosauriformes, as well as several more highly nested divisions within the latter two clades. Turtles are here resolved as the sister taxon to a monophyletic Lepidosauria (squamates + Sphenodon), a novel phylogenetic position that nevertheless is consistent with recent molecular and morphological studies that have hypothesized diapsid affinities for this clade.     

THE QUANTIFICATION OF SOME PLANT-TAXA CIRCUMSCRIPTIONS

Intrataxon variability is estimated by the technique of numerical taxonomy. Taxa which had been determined by orthodox taxonomic methods are subjected to analyses by numerical phenetics. Comparisons are made of the amounts of cohesion among the members of different taxa in the Bromeliaceae at several taxonomic levels by computing the degrees of similarities between members of the same taxon and of different taxa. Results of these comparisons with several taxa reveal that the variability of a taxon in terms of phenetic distance computed with a large number of characters does not necessarily agree with a subjective appraisal of variability. The method facilitated difficult taxonomic determinations and provided an estimate of the relative evolutionary ages of different taxonomic groups.     

A problem of truth in biological systematics

A possibility to put a question of truth of knowledge in biological systematics is studied. It is shown that the problem of truth in reference to systematics is wider than a question of classified information reliability. Prerequisites needed for logically accurate formulation of a definition and criteria of truth are considered. It is shown that such prerequisites are present in taxonomic practice, namely in a process of diagnosis compiling. Philosophical analysis of this work has been carried out. Interpretation of an essence of systematics as classification is connected with use of classical concept of truth (which defines truth as correspondence between knowledge and object) in its undeveloped form. Carried analysis allows supposing that a theory of systematics based on diagnostics rather than on classification would be more prospective. Use of imperfect concept of truth can be seen also in notions that system of taxa must reflect its evolutionary history. Development and modernization of Aristotle's orientation to discovery of the object form can become an alternative to such opinions. An aspiration to achieve the truth is the main motive of systematic work. An influence of this aspiration on a selection of purposes of taxonomic work and theoretical comprehension of its bases is shown. Such features of modern biological systematics as its accessibility for new results, criticism in respect of external morphological characters, and interest in intraspecific variability are connected with this aspiration. This motive comes into contradiction with a tendency to withdraw the problem of truth as such, which takes place in some brunches of theoretic systematics.     

Coordinating the traditional and modern approaches in the systematics of insects

The problem of coordinating the traditional and modern approaches to systematics is ever-lasting due to the continuous development and enrichment of our knowledge of biodiversity, means of analysis, and concepts. Comparative morphology was and still is the cornerstone of studies of insect taxonomy. It gives the most extensive and diverse information on the organisms studied, particularly when it is supported by the data on embryology and functional morphology as well as by analysis of adaptive significance of morphological characters. The limitations of this approach are often related to the presence of homoplasies, reversions, etc. Comparative paleontology is the only approach providing direct evidence of the historical succession of taxa and their characters. However, this approach is fully applicable only to some groups due to the specific features of their morphology and taphonomy. All the modern approaches (molecular, cytogenetic, etc.) are very informative but also have their own limitations; they should not be contrasted with the traditional approaches and certainly should not replace them. The traditional approaches do not become obsolete; it is only their comparative importance in the set of taxonomic tools that may be reevaluated. No single approach can be considered universal for an unambiguous reconstruction of phylogeny and substantiation of the natural system of taxa. Each approach has its own advantages and limitations, and only combined use of different approaches allows a broader range of the problems to be solved. Different approaches may prevail in the studies of different groups of insects and at different levels of taxonomic hierarchy. The intuition of the taxonomist, which is so often criticized by the followers of “objective” systematics, is based on taxonomic experience and scope of knowledge of a particular taxon. It does not imply a subjective bias, but allows the taxonomist to choose the instruments adequate to a particular case.     

Charles Girard: Relationships and Representation in Nineteenth Century Systematics

Early nineteenth century systematists sought to describe what they called the Natural System or the Natural Classification. In the nineteenth century, there was no agreement about the basis of observed patterns of similarity between organisms. What did these systematists think they were doing, when they named taxa, proposed relationships between taxa, and arranged taxa into representational schemes? In this paper I explicate Charles Frederic Girard’s (1822–1895) theory and method of systematics. A student of Louis Agassiz, and subsequently (1850–1858) a collaborator with Spencer Baird, Girard claimed that natural classificatory methods do not presuppose either a special creationist or an evolutionary theory of the natural world. The natural system, in Girard’s view, comprises three distinct ways in which organisms can be related to each other. Girard analyzed these relationships, and justified his classificatory methodology, by appeal to his embryological and physiological work. Girard offers an explicit theoretical answer to the question, what characters are evidence for natural classificatory hypotheses? I show that the challenge of simultaneously depicting the three distinct types of relationship led Girard to add a third dimension to his classificatory diagrams.     

歧隐翅虫亚族(鞘翅目,隐翅虫科)分类特征分析及单系性质疑

一个分类阶元是否单系,在当代分类学与系统学的意义上直接决定这个分类阶元的有效性,即可否广泛接受.形态学分析和动物地理分布模式可以提供非常有价值的启示.鞘翅目隐翅虫科隐翅虫亚科隐翅虫族的歧隐翅虫亚族Anisolinina Hayashi,1993(Coleoptera,Staphylinidae,Staphylininae),尽管是一个建立不久的分类单元,但亚族的分类定义、包括的种属、以及各属种的相互关系等,最近发生了较大的变化.通过分析一些重要的、用来定义这个亚族的关键形态性状,如前胸背板缘折上缘线与下缘线、上颚须、下唇须等,比较亚族内各属级单元的分类历史与动物地理分布,揭示歧隐翅虫亚族Anisolinina不是一个单系类群.     

Molecular phylogeny and evolutionary patterns of the european satyrids (Lepidoptera: Satyridae) as revealed by mitochondrial gene sequences

We examine the phylogenetic relationships of more than 40 species of European satyrids representing six tribes (Coenonymphini, Erebiini, Maniolini, Satyrini, Melanargiini, and Lethini). The analyses are based on comparisons of morphological data and mitochondrial genes encoding the large ribosomal subunit (16S rDNA) and NADH dehydrogenase subunit 1 (ND1). The cladistic reassessment of systematics based on morphological characters differs from the view retained by Miller by a lack in resolution due to the low number of characters used. Furthermore, some level of incongruence about the monophyly of the tribes is found between topologies from morphological and molecular analyses. In the case of Aphantopus hyperantus, molecular data and reexamination of morphology of this taxon indicate that this species has to be included within Maniolini. Contrary to the other clades, Erebia displays a radiate systematic pattern which cannot be explained by a lack of variable or informative sites. The combined spatial and temporal specialization found in the Erebia species may explain the rapid diversification of this genus relative to other satyrids. Finally, the subfamily level as defined by Miller for the taxa presented in the data set (Satyrinae and Elymninae) is not consistent with the molecular data. Given the reassessment of satyrids as a subfamily within Nymphalidae (Satyrinae), it seems more appropriate to retain the tribes as valid taxonomic ranks only in Satyrinae.     

A phylogenetic investigation of the Neotropical genus Alphamenes van der Vecht, 1977 (Hymenoptera,Vespidae, Eumeninae)

Advances in potter wasp systematics have been achieved recently, with classificatory changes resulting from analyses based upon large scale molecular datasets. For the Neotropics, recent hypotheses point to the occurrence of an exclusive clade recognized within the tribe Eumenini. In this group, several contributions regarding taxonomy and systematics have been proposed in the last five years, including the genus Alphamenes. This taxon contains seven described species whose distribution is exclusively Neotropical. Females are morphologically homogeneous, and characters related to copulatory organs are useful in male diagnosis. This contribution forms the first phylogenetic approach to include all species of Alphamenes, hence the first to strongly test for group monophyly. Our cladistic results recovered Alphamenes as a monophyletic group supported by male genital features. Relationships among included species also rely upon genitalic characters, highlighting the importance of these attributes for eumenine systematics. Recent phylogenetic investigations applied to the Neotropical fauna of potter wasps represent desirable advancements towards a natural classification for the group.     

Age rank/clade rank metrics--sampling,taxonomy, and the meaning of "stratigraphic consistency"

Paleontologists frequently contrast clade rank (i.e., nodal or patristic distance from the base of a cladogram) with age rank (i.e., relative first known appearances of the analyzed taxa) to measure the degree of congruence between the estimated phylogeny and the fossil record. Although some potential biases of these methods have been examined (e.g., the effect of tree imbalance), other properties of age rank/clade rank (ARCR) comparisons have not been studied in detail. A basic premise of ARCR metrics is that outgroup taxa diverged earlier than ingroups and thus should first appear in older strata. For example, given phylogeny (A,(B,C)), then taxon A should be sampled before either taxon B or taxon C. We examine this premise in the context of (1) phylogenetic theory, (2) taxonomic practice, (3) sampling intensity (R), and (4) factors other than sampling intensity (including cladogram accuracy). Simulations combining clade evolution and sampling over time indicate a poor relationship between ARCR metrics and R when all taxa are apomorphy-based monophyletic groups. However, a good relationship exists when taxa are either stem-based monophyletic groups or if workers include taxa without a priori decisions about monophyly or paraphyly. These results are not surprising because cladograms predict the order in which lineages diverged (which applies to stem-based monophyletic taxa) and the order in which morphologic grades appeared (which applies to paraphyletic taxa relative to derived monophyletic groups). Other factors that increase ARCR metrics when the average R stays the same include high temporal variation in R, budding instead of bifurcating speciation patterns, low extinction rates, cladogram inaccuracy, and (to a much lesser extent) large clade size. These results suggest several plausible explanations for patterned differences in ARCR metrics among clades, thereby compromising their validity as measures of the quality of the fossil record.     

The phylogenetic trunk: maximal inclusion of taxa with missing data in an analysis of the lepospondyli (Vertebrata,Tetrapoda)

The importance of fossils to phylogenetic reconstruction is well established. However, analyses of fossil data sets are confounded by problems related to the less complete nature of the specimens. Taxa that are incompletely known are problematic because of the uncertainty of their placement within a tree, leading to a proliferation of most-parsimonious solutions and "wild card" behavior. Problematic taxa are commonly deleted based on a priori criteria of completeness. Paradoxically, a taxon's problematic behavior is tree dependent, and levels of completeness are not directly associated with problematic behavior. Exclusion of taxa on the basis of completeness eliminates real character conflict and, by not allowing incomplete taxa to determine tree topology, diminishes the phylogenetic hypothesis. Here, the phylogenetic trunk approach is proposed to allow optimization of taxonomic inclusion and tree stability. The use of this method in an analysis of the Paleozoic Lepospondyli finds a single most-parsimonious tree, or trunk, after the removal of one taxon identified as being problematic. Moreover, the 38 trees found at one additional step from this primary trunk were reduced to 2 by removal of one additional taxon. These trunks are compared with the trees that were found by excluding taxa with various degrees of completeness, and the effects of incomplete taxa are explored with regard to use of the trunk. Correlated characters associated with limblessness are discussed regarding the assumption of character independence; however, inclusion of intermediate taxa is found to be the single best method for breaking down long branches.     

No speed dating please! Patterns of social preference in male and female house mice

Glass sponges (Class Hexactinellida) are important components of deep-sea ecosystems and are of interest from geological and materials science perspectives. The reconstruction of their phylogeny with molecular data has only recently begun and shows a better agreement with morphology-based systematics than is typical for other sponge groups, likely because of a greater number of informative morphological characters. However, inconsistencies remain that have far-reaching implications for hypotheses about the evolution of their major skeletal construction types (body plans). Furthermore, less than half of all described extant genera have been sampled for molecular systematics, and several taxa important for understanding skeletal evolution are still missing. Increased taxon sampling for molecular phylogenetics of this group is therefore urgently needed. However, due to their remote habitat and often poorly preserved museum material, sequencing all 126 currently recognized extant genera will be difficult to achieve. Utilizing morphological data to incorporate unsequenced taxa into an integrative systematics framework therefore holds great promise, but it is unclear which methodological approach best suits this task. Here, we increase the taxon sampling of four previously established molecular markers (18S, 28S, and 16S ribosomal DNA, as well as cytochrome oxidase subunit I) by 12 genera, for the first time including representatives of the order Aulocalycoida and the type genus of Dactylocalycidae, taxa that are key to understanding hexactinellid body plan evolution. Phylogenetic analyses suggest that Aulocalycoida is diphyletic and provide further support for the paraphyly of order Hexactinosida; hence these orders are abolished from the Linnean classification. We further assembled morphological character matrices to integrate so far unsequenced genera into phylogenetic analyses in maximum parsimony (MP), maximum likelihood (ML), Bayesian, and morphology-based binning frameworks. We find that of these four approaches, total-evidence analysis using MP gave the most plausible results concerning congruence with existing phylogenetic and taxonomic hypotheses, whereas the other methods, especially ML and binning, performed more poorly. We use our total-evidence phylogeny of all extant glass sponge genera for ancestral state reconstruction of morphological characters in MP and ML frameworks, gaining new insights into the evolution of major hexactinellid body plans and other characters such as different spicule types. Our study demonstrates how a comprehensive, albeit in some parts provisional, phylogeny of a larger taxon can be achieved with an integrative approach utilizing molecular and morphological data, and how this can be used as a basis for understanding phenotypic evolution. The datasets and associated trees presented here are intended as a resource and starting point for future work on glass sponge evolution.     

系统发育和被子植物“多系-多期-多域”系统——兼答傅德志的评论

历史地回顾了谱系、系统发育、单系、多系概念的由来和发展。详细讨论了单系(monophyly )和多系(polyphyly)在叙述分类群起源和描述分类群内部系统发育线多样性的不同含义。以此为基础,说明在“多系_多2_多域”的八纲系统中“多系”和“单系”是指自然分类群内系统发育线的多样性,不等同于“多源”和“单源”。通过与近年来所发表的被子植物分类系统的比较,认为基于化石、形态、分子和地理分布证据提出的八纲系统所显示的预言性,必将在今后的科学实践中经受检验。     

Infraspecific variation in Cerastium alpinum s. lat. (Caryophyllaceae) in Central Norway

Within Cerastium alpinum s. lat., several taxa have been circumscribed. Three infraspecific taxa are currently recognized in Nordic literature, but the opinions have varied about their taxonomic levels and relations to each other. Populations of these three taxa — alpinum, glabratum and lanatum — were investigated in Central Norway to elucidate taxonomic relationships on a regional scale. Morphometric analyses (PCA ordination) based on two data sets; one with 61 characters, and one without the 35 indumentum characters, gave different results. The former partly support the use of a higher rank for glabratum , but this analysis may put too much weight on indumentum characters. The reduced data set gave a more continuous transition between the three taxa, with no special status for glabratum. Crossing experiments between the taxa indicated full cross compatibility as far as relative seed set was concerned. Finally, vegetation analysis (DCA ordination) indicated that the three taxa have somewhat different ecological demands. This suggests that they, in spite of hybridization abilities, normally are maintained as separate entities due to habitat differences. A delimitation of three infraspecific taxa at the same level within C. alpinum s. lat. is thus supported by this study. Whether these taxa should be treated as subspecies or varieties strongly depends upon the definitions of these categories. Viewed in a wide geographical context, however, the arguments for subspecific rank are strengthened.     

Benzyltetrahydroisoquinoline-derived alkaloids as systematic markers

Basic unsubstituted skeleta within biogenetic groups can be codified according to their position along biosynthetic pathways aligned in order of probability of occurrence. Correlation of the codified structural type with the taxa in which they are found is effected through frequency indices associated with the relative number of derivatives of the skeleton in the taxon and the relative number of subunits of the taxon which contain derivatives thereof. The benzyltetrahydroisoquinolines, a biogenetically related group of compounds, serves as example of the applicability of the method, which is part of an integrated system of expression of biochemical data in biological systematics.     

Multivariate analysis of morphological variation among closely related species Bromus japonicus, B. squarrosus and B. arvensis (Poaceae) in comparison with isozyme evidences

Morphological variation and taxonomic continuum of 110 specimens of Bromus arvensis, B. japonicus and B. squarrosus were analysed to assess the morphological variability of these close taxa. The importance of every morphological character was estimated by the GLM procedure. Stepwise discriminant analysis was used in order to find a set of morphological characters that maximises differences among the species. According to this, the most important characters appeared to be the length of anthers, width of lemma and lemma margin angle shape. The most important characters for the determination of Bromus arvensis was the length of anthers and for B. squarrosus width of lemma. According to the UPGMA cluster analysis, canonical discriminant, classificatory discriminant and principal component analyses, all specimens were separated into three moderately distinct groups which corresponded to the three traditional species. The study showed that qualitative characters were the best for the delimitation of the tax a by statistical analyses and demonstrated the incongruity between the pattern of morphological and genetic (allozyme) variation among the three Bromus species.