Family-level Classification Research Articles

PHYLOGENETIC RELATIONSHIPS OF PALAEACANTHOCEPHALA (ACANTHOCEPHALA) INFERRED FROM SSU AND LSU rDNA GENE SEQUENCES

The Palaeacanthocephala is traditionally represented by 2 orders, Echinorhynchida and Polymorphida, with 10 and 3 families, respectively. To test the monophyly of the class, these 2 orders, and certain families, phylogenies were inferred using nuclear small-subunit (SSU) and large-subunit (LSU) ribosomal DNA sequences obtained for 29 species representing 10 families, 2 other classes of acanthocephalans, and 3 rotifer outgroups. Phylogenetic relationships were inferred by analyzing combined SSU and LSU sequences using maximum parsimony (MP) and maximum likelihood (ML) methods. Parsimony and ML trees inferred from combined analysis of these rDNA data strongly supported monophyly of Palaeacanthocephala and provided good resolution among species. Neither Polymorphida nor Echinorhynchida was monophyletic. Gorgorhynchoides bullocki (Echinorhynchida) was nested within the 6 species representing Polymorphida, and this clade was nested within species representing Echinorhynchida. Three of 4 palaeacanthocephalan families that could be evaluated were not monophyletic, and this finding was strongly supported. These results indicate that the family level classification of palaeacanthocephalans, which is mainly based on combinations of shared characters (not shared derived characters), needs to be reevaluated with respect to comprehensively sampled phylogenetic hypotheses.

System of Families of Enoplan Nemerteans of the Order Eumonostilifera (Nemertea: Enopla)

The paper is a review of the family-level classifications of the order Eumonostilifera. The elaboration of a phylogenetic system of Eumonostilifera is not yet possible. Some new groups, presumably monophyletic, have been distinguished, which are considered to be of a family rank (Neesiidae fam. n., Sacconemertidae stat. n., Zygonemertidae fam. n., Prostomatidae, Acteonemertidae fam. n., and Potamonemertidae fam. n.). The proposed system is a typological one; however, it can be used as a basis for further taxonomical and phylogenetic studies.

Phylogeny of the superfamily Gelechioidea (Lepidoptera: Ditrysia): an exemplar approach.

Phylogenetic relationships within the megadiverse lepidopteran superfamily Gelechioidea have been poorly understood and consequently the family level classification has been problematic. An analysis of phylogeny using 193 characters, including 241 informative character states, derived from larval, pupal and adult morphology and larval ecology, was performed to resolve the phylogeny of the Gelechioidea. 143 species representing the diversity of the putative Gelechioidea were included, supplemented with 13 species representing 11 other Ditrysian families. The monophyly of the Gelechioidea was supported, although only with homoplastic characters. The putative position of the Gelechioidea as the sister group of the Apoditrysia was not supported, since the Gelechioidea was nested within this clade. The Gelechioidea was divided into two main lineages: (1) the gelechiid lineage constituting Deoclonidae, Syringopainae, a re-composed Coleophoridae (including Coelopoetinae and Batrachedrinae as paraphyletic with Stathmopodinae, and Coleophorinae nested within it), Momphidae, Pterolonchidae, Scythrididae, Cosmopterigidae, and Gelechiidae, and (2) the oecophorid lineage constituting the "autostichid" family assemblage (including taxa formerly assigned to Autostichinae, Holcopogoninae, Symmocinae, Glyphidoceridae and Lecithoceridae), Xyloryctidae s.l. (including a paraphyletic Xyloryctidae of authors, some oecophorids of authors, Deuterogoniinae and Blastobasinae), Oecophoridae s.s., Amphisbatidae s.s., Carcinidae, Stenomati[n/d]ae, Chimabachidae and Elachistidae (including Depressariinae s.s., Telechrysis, Ethmiinae, Hypertrophinae s.l., miscellaneous "amphisbatids"sensu authors, Aeolanthinae, Parametriotinae, Agonoxeninae and Elachistinae). Detritivory/fungivory may have evolved only twice within Gelechioidea, though the evolution of larval food substrate use frequently reverses. To avoid an unnecessary further proliferation of names, it is recommended that no further family group names are introduced within the Gelechioidea, unless based on a rigorous analysis of inter-relationships.

Phylogeny and classification of the extant basal lineages of the Hymenoptera (Insecta)

The phylogeny of the basal hymenopteran lineages, including representatives of all ‘symphytan’ families, is anal; In total, 236 morphological characters were scored for 44 exemplars, including six outgroup, two xyelic tenthredinoid, five pamphilioid, three cephoid, five ‘siricoid’, one orussid, and six apocritan taxa. The datas analysed with parsimony under equal weights and under implied weights. The monophyly of the Hymenopte strongly supported but the sistergroup of the Hymenoptera cannot be identified with confidence. The relations of the ‘symphytan’ lineages are Xyeloidea +(Tenthredinoidea+ (Pamphilioidea + (Cephoidea + (Ariaxyelic (Siricidae + (Xiphydriidae +(Orussoidea+Apocrita))))))). Many of the relationships between the superfamilies, especially in the basal branching pattern, are only weakly corroborated. The monophyly of most superfamilies is supported, and all may be monophyletic except the ‘Siricoidea’, which is clearly paraphyletic. It is difficult to di whether the Siricidae or the Anaxyelidae are the closest relatives of Xiphydriidae + (Orussoidea + Apocrita). support for the sistergroup relationship between the Orussoidea and the Apocrita is substantial, putative apomorphies being provided by most character systems. There is also good evidence in favour of the monophj the Apocrita. The internal phylogeny of the Tenthredinoidea differs considerably from the results of earlier anal The Blasticotomidae are the sistergroup of the Tenthredinoidea s.s. Relationships at the base of the Tenthredini s.s. are weakly supported. It is uncertain whether the Tenthredinidae are monophyletic or comprise a 1 paraphyletic grade within the Tenthredinoidea s.s. The Diprionidae may be the sistergroup to Cimbicidae +(Argidae+ Pergidae). Most relationships within the Cimbicidae + (Argidae + Pergidae) clade are corroborated, with the exception of the monophyly of the Argidae. It is proposed to elevate the Anaxyelidae the Xiphydriida both to superfamily status. The family-level classification of the Tenthredinoidea will probably have to be changed, but this must await further clarification of the phylogeny of this superfamily.

Phylogeny and classification of the extant basal lineages of the Hymenoptera (Insecta)

The phylogeny of the basal hymenopteran lineages, including representatives of all «symphytan» families, is analysed. In total, 236 morphological characters were scored for 44 exemplars, including six outgroup, two xyelid, 16 tenthredinoid, five pamphilioid, three cephoid, five «siricoid», one orussid, and six apocritan taxa. The dataset is analysed with parsimony under equal weights and under implied weights. The monophyly of the Hymenoptera is strongly supported but the sistergroup of the Hymenoptera cannot be identified with confidence. The relationships of the «symphytan» lineages are Xyeloidea+(Tenthredinoidea+(Pamphilioidea+(Cephoidea+(Anaxyelidae+ (Siricidae+(Xiphydriidae+(Orussoidea+Apocrita))))))). Many of the relationships between the superfamilies, especially in the basal branching pattern, are only weakly corroborated. The monophyly of most superfamilies is well supported, and all may be monophyletic except the «Siricoidea», which is clearly paraphyletic. It is difficult to decide whether the Siricidae or the Anaxyelidae are the closest relatives of Xiphydriidae+(Orussoidea+Apocrita). The support for the sistergroup relationship between the Orussoidea and the Apocrita is substantial, putative synapomorphies being provided by most character systems. There is also good evidence in favour of the monophyly of the Apocrita. The internal phylogeny of the Tenthredinoidea differs considerably from the results of earlier analyses. The Blasticotomidae are the sistergroup of the Tenthredinoidea s.s. Relationships at the base of the Tenthredinoidea s.s. are weakly supported. It is uncertain whether the Tenthredinidae are monophyletic or comprise a basal paraphyletic grade within the Tenthredinoidea s.s. The Diprionidae may be the sistergroup to the Cimbicidae+(Argidae+Pergidae). Most relationships within the Cimbicidae+(Argidae+Pergidae) clade are well corroborated, with the exception of the monophyly of the Argidae. It is proposed to elevate the Anaxyelidae and the Xiphydriida both to superfamily status. The family-level classification of the Tenthredinoidea will probably also have to be changed, but this must await further clarification of the phylogeny of this superfamily.

Evolutionary Relationships among Mucoralean Fungi (Zygomycota): Evidence for Family Polyphyly on a Large Scale

Mucorales (Zygomycota) are ubiquitous, morphologically simple terrestrial fungi that are united taxonomically by possession of a coenocytic mycelium upon which nonmotile mitotic spores are produced asexually in uni-to multispored sporangia, and zygospores, where known, are produced following fusion of sexually compatible hyphae. Here we report the first comprehensive phylogenetic analysis of essentially all genera of Mucorales (63 species, 54 genera and 13 families) based on partial nucleotide sequence data of nuclear small subunit (18S) ribosomal RNA and nuclear large subunit (28S) ribosomal RNA genes, translation elongation factor-1α gene exons, and a morphological data set consisting of 1826, 389, 1092 and 11 characters, respectively. Individual and combined data sets were analyzed by unequally weighted maximum parsimony (MP) to investigate evolutionary relationships among and within mucoralean families. A Micromucor—Umbelopsis clade, traditionally included in the Mortierellaceae, was identified as the basal sister-group to all other Mucorales. A major discovery of this study is that traditional family-level classification schemes for this order appear to be highly artificial as evidenced by polyphyly of four of the seven families containing two or more genera. As presently circumscribed, these four families include 83% of the Mucorales. In addition, the largest and best known genera, Mucor and Absidia, were resolved as polyphyletic. The results provide a robust phylogenetic framework for additional evolutionary studies of the Mucorales.

Evolutionary relationships among mucoralean fungi (Zygomycota): Evidence for family polyphyly on a large scale

Mucorales (Zygomycota) are ubiquitous, morphologically simple terrestrial fungi that are united taxonomically by possession of a coenocytic mycelium upon which nonmotile mitotic spores are produced asexually in uni-to multispored sporangia, and zygospores, where known, are produced following fusion of sexually compatible hyphae. Here we report the first comprehensive phylogenetic analysis of essentially all genera of Mucorales (63 species, 54 genera and 13 families) based on partial nucleotide sequence data of nuclear small subunit (18S) ribosomal RNA and nuclear large subunit (28S) ribosomal RNA genes, translation elongation factor-1α gene exons, and a morphological data set consisting of 1826, 389, 1092 and 11 characters, respectively. Individual and combined data sets were analyzed by unequally weighted maximum parsimony (MP) to investigate evolutionary relationships among and within mucoralean families. A Micromucor—Umbelopsis clade, traditionally included in the Mortierellaceae, was identified as the basal sister-group to all other Mucorales. A major discovery of this study is that traditional family-level classification schemes for this order appear to be highly artificial as evidenced by polyphyly of four of the seven families containing two or more genera. As presently circumscribed, these four families include 83% of the Mucorales. In addition, the largest and best known genera, Mucor and Absidia, were resolved as polyphyletic. The results provide a robust phylogenetic framework for additional evolutionary studies of the Mucorales.

IDENTIFYING BRITISH INSECTS and ARACHNIDS, AN ANNOTATED BIBLIOGRAPHY OF KEY WORKS. Edited by Peter C. Barnard. Cambridge University Press, Cambridge, U.K. 1999. Pp. 353. Hardcover. f50.00. ISBN 0521 63241 2.

IDENTIFYING BRITISH INSECTS and ARACHNIDS, AN ANNOTATED BIBLIOGRAPHY OF KEY WORKS. Edited by Peter C. Barnard. Cambridge University Press, Cambridge, U.K. 1999. Pp. 353. Hardcover. f50.00. ISBN 0521 63241 2.

Cladistics and Family Level Classification of the Gentianales

Abstract—The most recent classification of the angiosperm order Gentianales (Thorne, 1992) includes four principal families: Apocynaceae, Gentianaceae, Loganiaceae, and Rubiaceae. Ever since Bentham (1857) the status of Loganiaceae has been questioned, and several segregates of that family have been proposed both before and after his treatment. In this study we present cladistic results that show Loganiaceae, sensu lato, to be a paraphyletic group definable only by plesiomorphies, with members showing closest relationships to other families of the order. As the impact of different character-state representations of polymorphic terminals remains largely untested, our morphological and phytochemical data were analysed both with restricted polymorphism coding as well as with the monomorphic "subtaxon" recoding method of Nixon and Davis (1991). Both approaches yield highly compatible results, and we here discuss a new classification of the Gentianales based on (i) monophyletic groups identified by outgroup analysis, and (ii) the maximal portrayal of evidence provided by subtaxon polymorphism recoding. Most prominently, the Loganiaceae sensu lato are divided into four segregate families, two previously named (Loganiaceae sensu stricto and Strychnaceae), and two defined as a result of this study (Gelsemiaceae, L. Struwe & V. A. Albert, stat. nov. and Geniostomaceae, L. Struwe & V. A. Albert, fam. nov.). Apocynaceae (incl. Asclepiadaceae), Gentianaceae (incl. Loganiaceae—Potalieae), and Rubiaceae remain as monophyletic families. Outgroup analysis supports both the monophyly of the Gentianales as well as the exclusion from the order of Buddleja, Desfontainia, Plocosperma, Polypremum, and Retzia (all Loganiaceae sensu Leeuwenberg and Leenhouts).

A phylogenetic analysis of the isopod family Janiridae (Crustacea)

A phylogeny of the isopod family Janiridae and genera from presumptive outgroups, Acanthaspidiidae, Joeropsididae and Microparasellidae is estimated. Characters were gathered from the published literature, and assembled into a data matrix for cladistic analysis. The data, when evaluated with heuristic search algorithms, yielded eight most-parsimonious trees, none of which supported the monophyly of the Janiridae. To evaluate the impact of homoplasy, characters with a rescaled consistency less than 0.1 were deleted, resulting in four somewhat different trees that were non-monophyletic for the janirids. With the smaller data set, trees supporting janirid monophyly were 10 steps longer. A permutation tail probability test found substantially more hierarchical information in the janirid data set than in randomised data. Internal topologies of the shortest trees were evaluated as hypotheses for new family-level groups, although new family-level classifications cannot be proposed at this time owing to insufficient evidence. The Janiridae therefore cannot be considered monophyletic.

A revision of the genus Lomanella Pocock and its implications for family level classification in the Travunioidea (Arachnida: Opiliones: Triaenonychidae)

A revision of the genus Lomanella Pocock and its implications for family level classification in the Travunioidea (Arachnida: Opiliones: Triaenonychidae)

The influence of seasonal and taxonomic factors on the ordination and classification of running‐water sites in Great Britain and on the prediction of their macro‐invertebrate communities

SUMMARY. 1. Macro‐invertebrate samples were collected from 268 running‐water sites in Great Britain in each of three seasons (spring, summer and autumn). A combined seasons’treatment was generated by amalgamating the individual seasons’data. These four seasonal options were each subjected to four distinct taxonomic analyses differing in level of identification and whether the data were quantitative or qualitative. Thus sixteen data‐sets were available for analysis. Environmental data on physical and chemical variables, macrophyte cover and date of sampling were also recorded for each site.2. All sixteen data‐sets were ordinated by detrended correspondence analysis and classified by two‐way indicator species analysis. There were strong correlations between the sixteen ordinations and significant concordance between classifications.3. The relationships between ordination scores and single environmental variables were investigated. Muhiple discriminant analysis was used to fit environmental data to eight selected classifications covering the full range of seasonal and taxonomic treatments. The environmental variables most useful in distinguishing between rivers were substratum characteristics, alkalinity and total oxidized nitrogen. Within‐river differences were often highly correlated with discharge, distance from source, width and depth. Slope and altitude contributed strongly to both between‐river and within‐river distinctions.4. Between‐site variation (beta diversity), eigenvalues of ordination, the reliability of classifications, the proportion of sites correctly assigned to their biological group using environmental data and the standardized similarity between observed and predicted fauna were all higher when identifications were taken to species level, rather than one of three family treatments. Qualitative data on a reduced list of families gave comparable or better results than more detailed family treatments.5. Combined seasons’data enabled better categorization and prediction than single season's.6. The values of the Czekanowski Index of Similarity between the observed and predicted fauna of test sites were close to realistic maximum values.7. Recommendations are made concerning potential usages of the various classifications. The species level classification has uses in the field of conservation and in the prediction of biological response to environmental change. The family level classifications have value in developing local site inventories and in the interpretation of pollution surveillance programmes.