Definitions in phylogenetic taxonomy: critique and rationale

A general rationale for the formulation and placement of taxonomic definitions in phylogenetic taxonomy is proposed, and commonly used terms such as "crown taxon" or "node-based definition" are more precisely defined. In the formulation of phylogenetic definitions, nested reference taxa stabilize taxonomic content. A definitional configuration termed a node-stem triplet also stabilizes the relationship between the trio of taxa at a branchpoint, in the face of local change in phylogenetic relationships or addition/deletion of taxa. Crown-total taxonomies use survivorship as a criterion for placement of node-stem triplets within a taxonomic hierarchy. Diversity, morphology, and tradition also constitute heuristic criteria for placement of node-stem triplets.     

Phylogenetic definitions and taxonomic philosophy

An examination of the post-Darwinian history of biological taxonomy reveals an implicit assumption that the definitions of taxon names consist of lists of organismal traits. That assumption represents a failure to grant the concept of evolution a central role in taxonomy, and it causes conflicts between traditional methods of defining taxon names and evolutionary concepts of taxa. Phylogenetic definitions of taxon names (de Queiroz and Gauthier 1990) grant the concept of common ancestry a central role in the definitions of taxon names and thus constitute an important step in the development of phylogenetic taxonomy. By treating phylogenetic relationships rather than organismal traits as necessary and sufficient properties, phylogenetic definitions remove conflicts between the definitions of taxon names and evolutionary concepts of taxa. The general method of definition represented by phylogenetic definitions of clade names can be applied to the names of other kinds of composite wholes, including populations and biological species. That the names of individuals (composite wholes) can be defined in terms of necessary and sufficient properties provides the foundation for a synthesis of seemingly incompatible positions held by contemporary individualists and essentialists concerning the nature of taxa and the definitions of taxon names.     

菌物分类学研究中常见的物种概念

物种是生物多样性与分类学研究的基本单元, 物种识别是生物学研究的基本问题之一。物种的划分一直以来都没有一个明确统一的标准, 这使得分类学多少带有主观的色彩, 并经常被看作艺术而不完全是科学的研究。本文简要概述了菌物分类学研究中常见的3个物种概念, 即形态学种、生物学种和系统发育学种的背景和应用现状, 并通过实例讨论了这3个物种概念的特点及应用中存在的问题, 特别是各个物种概念之间的交错, 以期为菌物分类学研究和物种概念探讨提供参考。     

Taxonomic names and phylogenetic trees

This paper addresses the issue of philosophy of names within the context of biological taxonomy, more specifically how names refer. By contrasting two philosophies of names, one that is based on the idea that names can be defined and one that they cannot be defined, I point out some advantages of the latter within phylogenetic systematics. Due to the changing nature of phylogenetic hypotheses, the former approach tends to rob taxonomy from its unique communicative value since a name that is defined refers to whatever fits the definition. This is particularly troublesome should the hypothesis of phylogenetic relationship change. I argue that, should we decide to accept a new phylogenetic hypothesis, it is also likely that our view of what to name may change. A system where names only refer acknowledge this, and accordingly leaves it open whether to keep a name (and accept the way it refers in the new hypothesis) or discard a name and introduce new names for the parts of the tree that we find scientifically interesting. One of the main differences between a phylogenetic system of definition (PSD) and a phylogenetic system of reference (PSR) is that the former is governed by laws of language while the latter by communicative needs of taxonomists. Thus, a PSR tends to give primacy to phylogenetic trees rather than phylogenetic definitions of names should our views of which phylogenetic hypothesis to accept change. © 1998 The Norwegian Academy of Sciences and Letters     

Phylogeny of Early Cretaceous spatangoids (Echinodermata: Echinoidea) and taxonomic implications

A phylogenetic analysis of 36 species provides a test for the taxonomy and the history of Early Cretaceous spatangoids. Most taxonomic units from genera to suborders are consistent with the proposed phylogenetic framework. We retain Hemiasterina, Micrasterina, Hemiasteridae, Schizasteridae, Hemiaster , Heteraster , Mecaster , and Periaster as original monophyletic groups. However, all of these clades originate without the classical apomorphies normally ascribed to them. We suggest a revision of their diagnoses and of the generic attributions of basal species. Some ill-defined, 'primitive', and paraphyletic taxa are recognised: Toxaster , Epiaster , Palhemiaster , and Toxasteridae. Even if they do not have phylogenetic meaning, they are retained here, pending a more complete revision.     

Mitochondrial phylogeny of Locustella and related genera

We used maximum likelihood analysis of complete mitochondrial ND2 sequences (1041 bp) to clarify the taxonomy and relationships of various species and genera of grass and bush warblers. The tree revealed two clades of grass and bush warblers. One clade was comprised of all four western Palearctic Locustella and two species of Asian Bradypterus . The other clade included five eastern Palearctic Locustella (including the distinctive Sakhalin warbler Locustella amnicola ) and the marsh grassbird Megalurus pryeri . African Bradypterus and Australian little grassbird Megalurus gramineus were distantly related to their Asian congeners. Therefore, current taxonomy of these genera does not reflect their evolutionary history and needs revision. It is proposed that a phylogenetic analysis of morphology and ecological preferences would show that the current taxonomy of grass and bush warblers reflects species' habitat preferences and morphology related to locomotion and foraging in their habitats, rather than their shared ancestry. Distinct clades were found in grasshopper warbler Locustella naevia and Pallas's grasshopper warbler L. certhiola . Detailed phylogeographic studies are needed to elucidate the species status of the clades within these two species.     

Conodonts Meet Cladistics: Recovering Relationships and Assessing the Completeness of the Conodont Fossil Record

A numerical cladistic analysis of the conodont family Palmatolepidae has been undertaken to determine the applicability of the technique to group-wide systematic revision. Results suggest a new hypothesis of relationships that is considerably more parsimonious than trees compatible with existing hypotheses of relationships, or trees that are even loosely constrained stratigraphically. This may occur either because the fossil record is incomplete, because taxon sampling for the cladistic analysis is low, or because the most parsimonious trees approximate the true tree less well than do stratigraphically-constrained trees (or because of a combination of these factors). Although more taxa and more characters would be preferable in choosing between these possibilities, the tree derived solely from morphological data is adopted. Thus, stratigraphic data can be used to test hypotheses of relationships and construct phylogenies; hypotheses of relationships can be used to test the completeness of the conodont fossil record. Existing schemes of classification within the Palmatolepidae are rejected because most groups within them are either polyphyletic or paraphyletic. A new scheme is presented. Character changes suggest correlated, progressive and mosaic evolution within the Palmatolepidae. Parsimony analysis of partitioned datasets indicates that more phylogenetic information can be recovered from S rather than P or M element positions, although data from all three positional groups are preferable to data from just one. Thus, multielement taxonomy is essential to the resolution of conodont interrelationships.     

Linking of digital images to phylogenetic data matrices using a morphological ontology

Images are paramount in documentation of morphological data. Production and reproduction costs have traditionally limited how many illustrations taxonomy could afford to publish, and much comparative knowledge continues to be lost as generations turn over. Now digital images are cheaply produced and easily disseminated electronically but pose problems in maintenance, curation, sharing, and use, particularly in long-term data sets involving multiple collaborators and institutions. We propose an efficient linkage of images to phylogenetic data sets via an ontology of morphological terms; an underlying, fine-grained database of specimens, images, and associated metadata; fixation of the meaning of morphological terms (homolog names) by ostensive references to particular taxa; and formalization of images as standard views. The ontology provides the intellectual structure and fundamental design of the relationships and enables intelligent queries to populate phylogenetic data sets with images. The database itself documents primary morphological observations, their vouchers, and associated metadata, rather than the conventional data set cell, and thereby facilitates data maintenance despite character redefinition or specimen reidentification. It minimizes reexamination of specimens, loss of information or data quality, and echoes the data models of web-based repositories for images, specimens, and taxonomic names. Confusion and ambiguity in the meanings of technical morphological terms are reduced by ostensive definitions pointing to features in particular taxa, which may serve as reference for globally unique identifiers of characters. Finally, the concept of standard views (an image illustrating one or more homologs in a specific sex and life stage, in a specific orientation, using a specific device and preparation technique) enables efficient, dynamic linkage of images to the data set and automatic population of matrix cells with images independently of scoring decisions.     

A species concept for bacteria based on adaptive divergence

Bacterial strains are currently grouped into species based on overall genomic similarity and sharing of phenotypes deemed ecologically important. Many believe this polyphasic taxonomy is in need of revision because it lacks grounding in evolutionary theory, and boundaries between species are arbitrary. Recent taxonomy efforts using multilocus sequence typing (MLST) data are based on the identification of distinct phylogenetic clusters. However, these approaches face the problem of deciding the phylogenetic level at which clusters are representative of evolutionary or taxonomically distinct units. In this review, I propose classifying two phylogenetic clusters as separate species only when they have statistically significantly diverged as a result of adaptive evolution. More than a method for classification, the concept of adaptive divergence can be used in a 'reverse ecology' approach to identify lineages that are in the process of speciation or genes involved in initial adaptive divergence.     

Molecular systematics and species limits in the Philippine fantails (Aves: Rhipidura)

Islands have long-attracted scientists because of their relatively simple biotas and stark geographic boundaries. However, for many islands and archipelagos, this simplicity may be overstated because of methodological and conceptual limitations when these biotas were described. One archipelago that has received relatively little recent attention is the Philippine islands. Although much of its biota was documented long ago, taxonomic revision and evolutionary study has been surprisingly scarce, and only a few molecular phylogenetic studies are beginning to appear. We present a molecular phylogeny and taxonomic revision for the Philippine fantails (Aves: Rhipidura) using nuclear and mitochondrial DNA sequences. Our results suggest that current taxonomy underestimates diversity in the group. Some morphologically distinct subspecies warrant species status, whereas one was indistinguishable genetically and morphologically and should not be retained. A few taxa require additional sampling for thorough taxonomic assessment. Patterns of diversity within Philippine Rhipidura mostly corroborate predictions of the Pleistocene aggregate island complex (PAIC) hypothesis, in which diversity is expected to be partitioned by deep water channels separating Pleistocene aggregate islands rather than by current islands. Substantial structure within PAIC clades indicates that additional drivers of diversification should be considered.     

Systematic revision of Entomobryidae (Collembola) by integrating molecular and new morphological evidence

Entomobryidae, the largest collembolan family, is traditionally classified at suprageneric level using a limited set of morphological structures, such as scales, antennal segmentation. Most tribal and subfamilial delimitations appear, however, disputable in the light of recent works. Integrating molecular and morphological evidence, we propose here a revision of the systematics of the family. In addition to traditional taxonomic characters, tergal specialized chaetae (S‐chaetae) are newly introduced, and their patterns are shown to be diversified at all levels from species to subfamilies. S‐chaetotaxic pattern on phylogenetic tree shows that evolution of S‐chaetae is not parallel between the different terga and that their patterns coincide well with the known molecular phylogeny, providing a powerful tool for the systematics of Entomobryidae. Orchesellinae sensu Soto‐Adames et al. (Annals of the Entomological Society of America, 101, 2008, 501); is divided into three subfamilies: Orchesellinae s. s., Bessoniellinae and Heteromurinae, the latter two upgraded from the original tribal level. Entomobryinae sensu Szeptycki (Morpho‐Systematic Studies on Collembola. IV. Chaetotaxy of the Entomobryidae and its Phylogenetical Significance, 1979), is no longer divided into scaled and unscaled tribes, and Lepidosira‐group is transferred from Seirinae to Entomobryinae. A key to subfamilies and tribes and a comparison with previous classifications of the Entomobryidae are provided. This study greatly improves the understanding of primary and secondary characters and erects the fundamental framework for the taxonomy of Entomobryidae.     

Analysis of ribosomal DNA internal transcribed spacer sequences of the Leishmania donovani complex

To understand phylogenetic relationships of species and strains within the Leishmania donovani complex, we have analyzed the ribosomal DNA internal transcribed spacer (ITS) sequences of 27 Leishmania infantum, 2 Leishmania chagasi, 18 L. donovani and 5 Leishmania archibaldi strains of different zymodemes and geographical origin. Eight ITS sequence types were found. All detected sequence variation within ITS1 and ITS2 was based on 12 polymorphic microsatellites. The L. infantum strains from the Mediterranean region, China and L. chagasi from the New World formed a phylogenetic group well separated from the second main group including all strains from East Africa and India. Within the latter group three distinct phylogenetic subgroups could be differentiated: (1) L. donovani (Sudan/Ethiopia, China) + L. archibaldi (Sudan), (2) L. donovani (Sudan/Ethiopia) + L. infantum (Sudan) + L. archibaldi (Sudan/Ethiopia), and (3) L. donovani (Kenya, India). These groups are not consistent with previous species definitions based on isoenzyme analyses, e.g. L. infantum is polyphyletic and L. archibaldi is not supported as a distinct species. Two groups of Indian strains could be differentiated, one of which has an identical sequence type to the strains from Kenya. Three main lineages of strains can thus be differentiated in East Africa: two quite distantly related groups of strains from Sudan/Ethiopia, and a third group including all strains from Kenya, which is more closely related to part of the Indian strains than to any of the Sudanese/Ethiopian groups. The ITS sequence analysis presented here supports the need for revision of the taxonomy of the L. donovani complex.     

Should taxon names be explicitly defined?

Overviews are provided for traditional and phylogenetic nomenclature. In traditional nomenclature, a name is provided with a type and a rank. In the rankless phylogenetic nomenclature, a taxon name is provided with an explicit phylogenetic definition, which attaches the name to a clade. Linnaeus’s approach to nomenclature is also reviewed, and it is shown that, although the current system of nomenclature does use some Linnaean conventions (e.g., certain rank-denoting terms, binary nomenclature), it is actually quite different from Linnaean nomenclature. The primary differences between traditional and phylogenetic nomenclature are reviewed. In phylogenetic nomenclature, names are provided with explicit phylogenetic definitions, whereas in traditional nomenclature names are not explicitly defined. In phylogenetic nomenclature, a name remains attached to a clade regardless of how future changes in phylogeny alter the clade’s content; in traditional nomenclature a name is not “married” to any particular clade. In traditional nomenclature, names must be assigned ranks (an admittedly arbitrary process), whereas in phylogenetic nomenclature there are no formal ranks. Therefore, in phylogenetic nomenclature, the name itself conveys no hierarchical information, and the name conveys nothing regarding set exclusivity. It is concluded that the current system is better able to handle new and unexpected changes in ideas about taxonomic relationships. This greater flexibility, coupled with the greater information content that the names themselves (i.e., when used outside the context of a given taxonomy or phytogeny) provide, makes the current system better designed for use by all users of taxon names.     

Metabolic fingerprinting as an indicator of biodiversity: towards understanding inter-specific relationships among Homoscleromorpha sponges

Sponges are an important source of secondary metabolites showing a great diversity of structures and biological activities. Secondary metabolites can display specificity on different taxonomic levels, from species to phylum, which can make them good taxonomic biomarkers. However, the knowledge available on the metabolome of non-model organisms is often poor. In this study, we demonstrate that sponge chemical diversity may be useful for fundamental issues in systematics or evolutionary biology, by using metabolic fingerprints as indicators of metabolomic diversity in order to assess interspecific relationships. The sponge clade Homoscleromorpha is particularly challenging because its chemistry has been little studied and its phylogeny is still debated. Identification at species level is often troublesome, especially for the highly diversified Oscarella genus which lacks the fundamental characters of sponge taxonomy. An HPLC–DAD–ELSD–MS metabolic fingerprinting approach was developed and applied to 10 Mediterranean Homoscleromorpha species as a rapid assessment of their chemical diversity. A first validation of our approach was to measure intraspecific variability, which was found significantly lower than interspecific variability obtained between two Oscarella sister-species. Interspecific relationships among Homoscleromorpha species were then inferred from the alignment of their metabolic fingerprints. The resulting classification is congruent with phylogenetic trees obtained for a DNA marker (mitochondrial COI) and demonstrates the existence of two distinct groups within Homoscleromorpha. Metabolic fingerprinting proves a useful complementary tool in sponge systematics. Our case study calls for a revision of Homoscleromorpha with further phylogenetic studies and identification of additional chemical synapomorphic characters.     

A bare-bones scheme to choose between the species,subspecies, and ‘evolutionarily significant unit’ categories in taxonomy and conservation

Species are the fundamental unit of analysis in biodiversity. They are often divided into subspecies, or into ‘evolutionarily significant units’ (ESUs). The imprecise demarcation between these categories causes disagreements and practical difficulties in taxonomy and conservation. I argue that it is possible to unequivocally choose which category to use for referring to any plant or animal population on the basis of a simple classificatory scheme, here proposed, resulting from the combination of stages of morphological and genetic differentiation. This scheme is the first to consider the three categories together in a unified frame of reference. Numerous definitions have been proposed for specific and infraspecific groupings. Most of them are redundant, or unnecessarily conflate the properties of species, subspecies, and ESUs. Based on feedback from the scheme, I reword general definitions for these categories to make them complementary, both conceptually and operationally.     

Immunological responses from brachiopod skeletal macromolecules; a new technique for assessing taxonomic relationships using shells

The phylogenetic relationships of living, calcareous-shelled brachiopods have been assessed on the basis of immunological responses from intracrystalline macromolecules. Calculations of immunological distances between representatives of the order Terebratulida have revealed a primary threefold division which correlates precisely with a proposed subdivision of the order into three superfamilies but refutes attempts to establish a fourth superfamily. This conclusion was confirmed by carrying out immunological investigations of small shell fragments from other brachiopod genera which were so rare that no antisera could be prepared. The immunological results also indicate a fundamental subdivision of the long-looped brachiopods, with one group perhaps being derived from short-looped terebratuloids rather than long-looped terebratelloids. Sero-taxonomy of skeletal macromolecules provides an ideal method of acquiring molecular phylogenetic data in many groups because a large number of taxa can be surveyed in a short period of time, and microscopic pieces of shell contain sufficient antigenic determinants for many reactions. The technique can also be applied to specimens which have been stored without special treatment in museum collections, making the technique particularly applicable to rare taxa for which no other form of molecular data is available. □ Immunology, Brachiopoda, taxonomy, intracrystalline molecules.     

Revision des evolutionsdenkens?

My criticism of the »organism-centered concept of evolution« as exposed once again by Bonik et al. (this vol.) refers to its phylogenetic application but not to the underlying basic assumption. The misunderstanding is due to the fact that Bonik et al. do not distinguish between evolutionary theory (which is concerned with general problems) and phylogenetic research. Nobody would deny that organisms are energy converting systems. Variants with increased energetic efficiency will of course be favoured by natural selection. But this premise alone does not allow to reconstitute phylogenetic lineages. Evolutionary success of a given variant is determined by many other factors as, for instance, the presence or absence of certain predators. Therefore phylogenetic research depends largely on circumstantial evidence from compared anatomy. Finally it should be stressed that the elimination of »abnormal ontogenetic stages« does not contribute to phylogenetic transformation. Bonik et al. are wrong when they believe that this kind of internal selection constitutes a new, hitherto neglected element which necessitates a fundamental revision of evolutionary theory.     

Resolving current disagreements and ambiguities in the terminology of animal communication

Communication is central to most interactions between organisms. There is currently considerable controversy about the evolution, function and even about the most basic definition of communication. The controversy is linked to definitional ambiguities and disagreements. Here we discuss how some recent disagreements can be resolved and offer a clear set of definitions. Central to our approach is a definition of communication as being a trade between one organism (the informer) and another (the perceiver). The informer exerts influence on the perceiver through the communication process, and the perceiver experiences a change in its informational state (that is, gains information) as a consequence of detecting the communication. We define both influence and information explicitly and delineate between signalling, deceptive communication, and situations where perceivers respond to cues rather than signals. We demonstrate how our definitions allow resolution of conflicts arising in recent publications on the definitions on communication and related terms.