Cladistic analysis of the Sepsidae (Cyclorrhapha: Diptera) based on a comparative scanning electron microscopic study of larvae

The result of a phylogenetic analysis of the Sepsidae based on larval characters is presented. It is shown that cyclorrhaphan larvae can be as rich a source of characters as Nematocera immatures when investigated using an SEM. The cladistic analysis comprised fifty-two species in sixteen genera of the Sepsidae and five outgroup species and used fifty-seven morphological characters. It found seven parsimonious trees which only differed with respect to the arrangement of some species within the genus Themira. The basal dichotomies of the phylogenetic trees are particularly well supported, indicating the conservative nature of larval characters. Orygma is confirmed as the sister group of all the remaining sepsids, the Sepsinae. There is good larval evidence that Ortalischema is the sister group of all remaining Sepsinae and that the Toxopodinae constitute an early radiation within the Sepsidae. According to larval data, some genera are paraphyletic ( Themira, Palaeosepsis ), but adult characters appear to contradict these findings. Among the traditionally recognized higher taxa within the Sepsidae, Hennig's Themira species-group and Steysbal's Sepsini have to be rejected as polyphyletic. However, Hennig's Sepsis species-group is confirmed as monophyletic and will probably constitute one major element of a future phylogenetic system of the Sepsidae. States of the strongly modified fore-legs of some adult sepsid males are mapped onto the phylogenetic tree, largely confirming Šulc's ideas about the evolution of these features. The origin and evolution of male sternites with brushes and a gland on the tibiae of the males ('osmeterium') are discussed. Whereas adult characters point to a sister-group relationship between the Sepsidae and the Ropalomeridae, larval characters appear to indicate a sister-group relationship between the Coelopidae and the Sepsidae. The evidence for both hypotheses is critically evaluated.     

Phylogeny of Aphantochilinae and Strophiinae sensu Simon (Araneae; Thomisidae)

This study tests the monophyly of ant‐mimicking Thomisidae (Aphantochilinae and Strophiinae sensu Simon), redefines the composition of these taxa, proposes tribes and discusses aspects of their myrmecomorphy and biogeography. The analysis is based on a matrix composed of 113 morphological characters and 37 terminal taxa (11 Aphantochilinae, 16 Strophiinae and 10 belonging to the out‐group). The 12 most parsimonious trees with 232 steps, obtained with equally weighted characters, support the monophyly of Aphantochilinae sensu Simon. Strophiinae emerges as a paraphyletic group divided into two clades: a basal clade that groups Strophius and Strigoplus (Strophiini new status) and another clade that includes Ceraarachne, Simorcus and Ulocymus (Ceraarachnini new status) as the sister group of Aphantochilus + Bucranium (Aphantochilini new status). Diagnoses are presented for the tribes and genera in this analysis. The synonymy between Bucranium and Aphantochilus is rejected. Majellula and Acracanthostoma are considered junior synonyms of Bucranium, and Synstrophius of Ceraarachne. The monophyly of Synstrophius is not recovered, S. blanci is transferred to Ceraarachne and S. muricatus is transferred to Ulocymus. Ant‐preying behaviour appears to be basal and has been documented for Strophiini and Aphantochilini species. Myrmecomorphy, which was documented for Aphantochilus, is presumably derived. The biogeographical analysis of Aphantochilinae and Strophiinae suggests an ancient relation between Neotropical, Afrotropical and Oriental species, with probable origin after the breakup of Gondwana, that is, in early Paleogene.     

Morphology and systematics of anthracomartidae (Arachnida: Trigonotarbida)

Abstract: X‐ray microtomography (XMT) was applied to three species of the extinct arachnid order Trigonotarbida, hosted in siderite nodules from the late Carboniferous British Middle Coal Measures. All three, Cryptomartus hindi ( Pocock, 1911 ), Cleptomartus plautus ( Petrunkevitch, 1949 ) and Maiocercus celticus ( Pocock, 1902 ), belong to the family Anthracomartidae. As well as providing interactive three‐dimensional visualisations of their likely appearance in life, XMT study has resolved new morphological detail, yielding key data about the likely stance and habitus of these early arachnids. Similarities in the form of the carapace, eyes and coxosternal region between anthracomartids and the Devonian genus Palaeocharinus Hirst, 1923 are highlighted. Analogies in limb disposition are drawn between anthracomartids and modern crab spiders (Araneae: Thomisidae), which exhibit sit‐and‐wait style predatory behaviour. Anthracomartids are relatively common fossils at many Coal Measures localities. Although they appear to exhibit limited morphological variation, numerous genera and species have been proposed in the literature, often distinguished from each other on rather trivial characters. From our reconstruction of well‐preserved examples, we have resolved a number of common features likely to be present in a typical anthracomartid. Comparisons to this model suggest that the characters on which many anthracomartid genera are distinguished may be artefacts of preservation. We hence treat Promygale Fri?, 1901 , Brachylycosa Fri?, 1904 , Perneria Fri?, 1904 , Coryphomartus Petrunkevitch, 1945 , Cryptomartus Petrunkevitch, 1945 , Pleomartus Petrunkevitch, 1945 , Cleptomartus Petrunkevitch, 1949 and Oomartus Petrunkevitch, 1953 as junior synonyms of the type genus of the family, Anthracomartus Karsch, 1882 . Only the genera Brachypyge Woodward, 1878 and Maiocercus Pocock, 1911 are additionally retained. Within Anthracomartus, Cleptomartus planus Petrunkevitch, 1949 , Cryptomartus meyeri Guthörl, 1964 , Cleptomartus hangardi Guthörl, 1965 and Cryptomartus rebskei Brauckmann, 1984 are treated as junior synonyms of A. hindi Pocock, 1911 (all syn. nov.). Cleptomartus plautus Petrunkevitch, 1949 and Anthracomartus denuiti Pruvost, 1922 are treated as junior synonyms of A. priesti Pocock, 1911 (both syn. nov.).     

Anatomy and phylogenetic analysis of the neotropical callichthyid catfishes (Ostariophysi, Siluriformes)

Based mainly on morphological characters, the phylogenetic relationships among genera and some species groups of the neotropical family Callichthyidae were examined. A study of the osteology of a generalized callichthyid, Callichthys callichthys (Linnaeus), with detailed comparisons among representatives of the remaining genera in the family, is presented and used as a basis for the phylogenetic analysis. A single most parsimonious tree supported the monophyly of the family Callichthyidae based on 28 derived features and the division of the family in the subfamilies Corydoradinae and Callichthyinae. In the subfamily Corydoradinae, the genus Aspidoras is the sister-group of the clade formed by Corydoras plus Brochis. Five derived features support the monophyly of this clade and four support the monophyly of Brochis. No characters, however, were found to support the genus Corydoras. In the subfamily Callichthyinae, Dianema and Hopbstemum are sister-taxa. Megalechis represents the sister-group of Dianema plus Hoplosternum and Lepthoplosternum represents the sister-group to Megalechis plus Dianema plus Hopbstemum. Finally, Callichthys is considered the least derived member of the subfamily, and is hypothesized as the sister-group of the remaining species. A key to all callichthyid genera is provided.     

Blackflies (Diptera: Simuliidae) of the Genera Hellichiella Rivosecchi et Cardinali, 1975 and Boreosimulium Rubzov et Yankovsky, 1982

The genus name Boreosimulium Rubzov et Yankovsky, 1982 was considered for a long time as a junior synonym of the genus name Hellichiella Rivosecchi et Cardinali, 1975. The status of Boreosimulium as a separate generic (or subgeneric) name has been confirmed in the monograph by Adler, Currie and Wood (2004). In the present paper the indentification keys for the Boreosimulium and Hellichiella (subfam. Simuliinae) species are given. Also the keys for the genera Stegopterna Enderlein, 1930 and Greniera Doby et David, 1959 (subfam. Prosimuliinae) being similar in some characters to the above genera are provided. The lists of species of these genera, together with their distribution ranges (according to the data obtained to the end of 2004) are added.     

18S rDNA phylogeny of the Tubificidae (Clitellata) and its constituent taxa: dismissal of the Naididae

The phylogeny of the Tubificidae, and of most of its subfamilies and some of its genera, is revisited, on the basis of sequences of 18S ribosomal DNA in a selection of species. Forty-six new 18S sequences of Naididae (6), Tubificidae (37), Phreodrilidae (1), Lumbriculidae (1), and Enchytraeidae (1) are reported and aligned together with corresponding sequences of 21 previously studied taxa. The 18S gene of Insulodrilus bifidus provides the first molecular evidence that phreodrilids are closely related to tubificids, corroborating previous conclusions based on morphology. The data further support the monophyletic status of Tubificidae, provided that the "Naididae" is regarded a part of this family; "naidids" may not even constitute a monophyletic group. It is thus suggested that the family name Naididae is formally suppressed as a junior synonym of the Tubificidae. The 18S gene also resolves a number of relationships within the tubificids. Among the subfamilies, Tubificinae is supported, Rhyacodrilinae and Phallodrilinae are revealed as nonmonophyletic, and Limnodriloidinae remains unresolved. Most tubificid genera tested for monophyly are corroborated by the data, only one (Tubifex) is refuted, and two (Tubificoides and Limnodriloides) are unresolved from other taxa. It is concluded that it will be valuable to expand the taxonomic sampling for 18S rDNA in clitellates, and in annelids in general, as this is likely to improve the resolution at many levels. However, it will be equally important to combine the annelid 18S data with other gene sequences and nonmolecular characters, to estimate the phylogeny of these common and diverse worms with greater precision.     

SYSTEMATICS OF THE CAMBRIAN TRILOBITE FAMILY NEPEIDAE, WITH REVISION OF AUSTRALIAN SPECIES

Abstract:  Taxonomy of the Cambrian trilobite family Nepeidae is revised. Morphometric analyses of the genera Nepea and Penarosa demonstrate that use of proportions of linear dimensions of the cranidium to differentiate species is invalid, and that infrageneric variation is continuous. The taxonomy and biostratigraphy of all Australian species of Nepeidae is revised. Species considered valid herein include: Nepea narinosa (type species), N. tonsillata , N. nans , Penarosa retifera (type species), P. elaticeps , P. rhinodelphis , P. netenta , Loxonepea loxophrys (type species) and Ferenepea hispida (type species). Folliceps is considered to be a junior subjective synonym of Nepea , Trinepea is regarded as a junior subjective synonym of Penarosa , and Ascionepea is considered to be a junior subjective synonym of Ferenepea .     

Clade stability and the addition of data: A case study from erigonine spiders (Araneae: Linyphiidae, Erigoninae)

This study presents a new phylogeny of erigonine spiders with emphasis on genera from the Neotropics. Thirty‐nine exemplar taxa representing mostly Neotropical genera were added to a global sample of 31 erigonine and 12 outgroup exemplar taxa analyzed in a previous study. These 82 taxa were coded for 176 (172 informative) mostly morphological characters. Eighty‐one characters were identical to or modified from the 73 (67 informative) characters included in a previous study; the remaining 95 characters are new. The complete data set includes 70 erigonine exemplars representing 65 genera, seven nonerigonine linyphiid exemplars, and five exemplars representing four araneoid families in the outgroup. Cladistic analysis resulted in a single most parsimonious tree (L =904, CI = 0.23, RI = 0.58; uninformative characters excluded: L = 900, CI = 0.23). This paper explores the implications of the new topology for the evolution of several characters of interest in erigonine evolution. The phylogeny implies that the desmitracheate condition is a synapomorphy of erigonines, with a reversal to the haplotracheate condition in one large clade within Erigoninae. We infer that the loss of the paracymbium in Neotropical erigonines occurred twice and may have progressed by different evolutionary pathways. Our phylogeny differs markedly from the previous cladistic hypothesis of erigonine relationships. We investigate how the addition of characters and taxa (alone and together) have altered the earlier hypothesis of erigonine phylogeny. We conclude that topological changes from the previous study to the current one are largely the result of adding and modifying characters, not adding taxa. Continuous Jackknife Function (CJF) analysis predicts that the inclusion of additional character data will continue to imply changes in the relationships among taxa in our analysis.     

Phylogenetic consideration of two scuticociliate genera, Philasterides and Boveria (Protozoa, Ciliophora) based on 18 S rRNA gene sequences

Many scuticociliates are facultative parasites of aquatic organisms and are among the most problematic ciliate taxa regarding their systematic relationships. The main reason is that most species, especially taxa in the order Thigmotrichida have similar morphology and have not been studied yet using molecular methods. In the present work, two scuticociliate genera, represented by two rare parasitic species, Philasterides armatalis (order Philasterida) and Boveria subcylindrica (order Thigmotrichida), were studied, and phylogenetic trees concerning these two genera were constructed based on their 18 S rRNA gene sequences. The results indicate that: 1) Philasterides forms a sister group with Philaster, supporting the classification that these two genera belong to the family Philasteridae; 2) it is confirmed that the nominal species, Philasterides dicentrarchi Dragesco et al., 1995 should be a junior synonym of Miamiensis avidus as revealed by both previous investigations and the data revealed in the present work; and 3) the poorly known form B. subcylindrica, the only member in the order Thigmotrichida, of which molecular data are available so far, always clusters with Cyclidium glaucoma, a highly specialized scuticociliate, indicating a sister relationship between the orders Thigmotrichida and Pleuronematida.     

THE TAXONOMIC AND PHYLOGENETIC POSITION OF THE PLESIOSAUROIDEA FROM THE LOWER JURASSIC POSIDONIA SHALE OF SOUTH-WEST GERMANY

Abstract:  The two plesiosauroid species from the Posidonia shale of Holzmaden, ' Plesiosaurus ' guilelmiimperatoris and ' Plesiosaurus ' brachypterygius , do not belong to Plesiosaurus but form new monotypic genera. The new genus Hydrorion is erected for ' P .' brachypterygius , and the genus Seeleyosaurus is re-established for ' P .' guilelmiimperatoris . The recently described species Plesiopterys wildii is regarded as a junior synonym of S. guilelmiimperatoris . A short phylogenetic analysis shows that S. guilelmiimperatoris and Muraenosaurus are basal elasmosaurs. H. brachypterygius , Occitanosaurus tournemirensis from France and Microcleidus homalospondylus from England form a monophyletic clade, which is the sister taxon to the Cretaceous elasmosaurs. A palaeobiogeographical comparison of plesiosaur localities in the Lower Jurassic shows distinct palaeobiogeographical zones for the Toarcian, with different plesiosaur taxa in England, Germany and France.     

Phylogenetic relationship among Libellula, Ladona and Plathemis (Odonata: Libellulidae) based on DNA sequence of mitochondrial 16S rRNA gene

Abstract. The type genus for the dragonfly family Libellulidae is Libellula. At present, Libellula s.l. includes twenty-nine species, whose distribution is largely Nearctic. Whether two other libellulid taxa, Ladona and Plathemis, should be considered synonyms of Libellula, subgenera of Libellula, or separate genera, has been a subject of intermittent debate for over a century. Earlier proposals concerning Ladona and Plathemis were based on a limited number of morphological characters and lacked rigorous phylogenetic analyses. Therefore, we used the DNA sequence of a portion of the mitochondrial 16S rRNA gene and parsimony, maximum likelihood and neighbour-joining analyses to explore whether Ladona and Plathemis are monophyletic lineages distinct from Libellula. We obtained ≈ 415 bp of DNA sequence from twenty-three taxa including thirteen species of Libellula s.s., all three recognized species of Ladona, the two species of Plathemis and representatives of four other libellulid genera. Tetragoneuria williamsoni (Odonata: Corduliidae) was included as the outgroup. Parsimony analysis suggested that Ladona and Plathemis are monophyletic lineages distinct from Libellula s.s. with a sister group relationship between Libellula and Ladona. The monophyly of Ladona, Plathemis and Libellula was supported in > 90% of bootstrap replications and in trees five to ten steps longer than the most parsimonious trees. Relationships inferred from maximum likelihood and neighbour-joining analyses also supported the monophyly of Ladona and Plathemis. The four other libellulid genera included in the study formed a monophyletic clade distinct from Libellula, Ladona and Plathemis. Based on our analysis, we propose that Ladona and Plathemis be considered either genera or subgenera within Libellulidae.     

安徽东至寒武纪早期三叶虫动物群

描述了采自皖南东至庞汪剖面和潘冲剖面黄柏岭组和大陈岭组中的三叶虫,共计9种8属和1亚属,其中Chuchiaspis dongzhiensissp.nov.是1新种;认为TingyuaniaSunetChang,1937是DinesusEtheridge,1896的晚出异名。     

Phylogenetic relationships in the tribe Oxyptilini (Lepidoptera,Pterophoridae, Pterophorinae) based on morphological data of adults

The monophyly of the tribe Oxyptilini and phylogenetic relationships of the genera embraced in this tribe were examined using 171 (75 binary and 96 multistate) characters of adult morphology. The study material included 98 species of 30 genera, representing all previously recognized genera of Oxyptilini, together with the genera Sphenarches, Antarches, Diacrotricha, and Cosmoclostis, four species of Oidaematophorini, three species of Platyptiliini, as well as three and two other species belonging to Pterophorini and Exelastini respectively. Two Agdistis species were used as outgroups. The cladistic analysis resulted in six equally parsimonious trees. A majority of the recovered synapomorphic characters have previously been used in the taxonomy of the subfamily. However, 25 novel characters were found. The monophyly of Oxyptilini was supported, although only with homoplastic characters and low amounts of tree confidence; the genera Capperia, Procapperia, Paracapperia, Oxyptilus, Megalorhipida, and Trichoptilus were found to be nonmonophyletic; Sphenarches and Antarches were recovered as members of Oxyptilini; the two genera Cosmoclostis and Diacrotricha were placed out of Oxyptilini, inside the tribe Pterophorini; and close affinity of the genus Dejongia to Stangeia, Stenodacma, Megalorhipida, Trichoptilus, and Buckleria species was revealed. Four new combinations, Cosmoclostis lanceata (Arenberger) comb. nov. , Nippoptilia regulus (Meyrick) comb. nov. , Capperia tadzhica (Zagulajev) comb. nov. , and Buckleria negotiosus (Meyrick) comb. nov. are proposed; Capperia insomnis Townsend was considered as a senior synonym of Procapperia hackeri Arenberger syn. nov. , Buckleria negotiosus (Meyrick) as a senior synonym of Buckleria vanderwolfi Gielis syn. nov. , and Oxyptilus variegatus Meyrick syn. nov. as a junior synonym of Oxyptilus secutor Meyrick. © 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 163 , 484–547.     

Sericolophus (Pheronematidae) and Hernandeziana (Corythophoridae) are synonyms (Hexactinellida: Porifera)

Eight specimens of Sericolophus reflexus (Ijima) were compared with the holotype of Hernandeziana ijimae (Hernandez) to assess suspected synonymy of these taxa. An extensive morphometric analysis of the spicules available in these specimens of variable condition show that the H. ijimae holotype exhibits characters near the centre of the range of S. reflexus. It is concluded that H. ijimae is a junior synonym of S. reflexus , and that the family Corythophoridae de Laubenfels is invalid by preoccupation of its base generic name.     

Phylogeny of the Archiborborinae (Diptera: Sphaeroceridae) Based on Combined Morphological and Molecular Analysis

The Archiborborinae is a diverse Neotropical subfamily of Sphaeroceridae, with many undescribed species. The existing generic classification includes three genera consisting of brachypterous species, with all other species placed in the genus Archiborborus. We present the first phylogenetic hypothesis for the subfamily based on morphological, molecular, and combined datasets. Morphological data include 53 characters and cover all valid described taxa (33 species in 4 genera) in the subfamily, as well as 83 undescribed species. Molecular data for five genes (mitochondrial 12S rDNA, cytochrome c oxidase subunit I, and cytochrome B, and nuclear alanyl-tRNA synthetase and 28S rDNA) were obtained for 21 ingroup taxa. Data support the separation of the Archiborborinae from the Copromyzinae, with which they were formerly combined. Analyses support consistent groups within the subfamily, but relationships between groups are poorly resolved. The validity of the brachypterous genera Penola Richards and Frutillaria Richards is supported. The former genus Archiborborus Duda is paraphyletic, and will be divided into monophyletic genera on the basis of this work. Aptery and brachyptery have evolved multiple times in the subfamily. Antrops Enderlein, previously including a single brachypterous species, is a senior synonym of Archiborborus.     

Gelechioidea (Insecta: Lepidoptera) systematics: a reexamination using combined morphology and mitochondrial DNA data

Phylogenies of Gelechioidea (Insecta: Lepidoptera) historically have been in disagreement and definitions vary at the family and subfamily levels. Addition of new taxa or new characters drastically changes relationships indicating that current phylogenetic schemes require more investigation. This study is the first phylogenetic analysis of Gelechioidea to include molecular data. Here we present a combined analysis using Maximum Parsimony to investigate sister-group relationships within Gelechioidea. The addition of Cytochrome oxidase I and II to revised published morphological matrices gives 453 parsimony informative characters for the 42 taxa for which we have sequence data. The combined analysis resulted in two trees with mostly novel sister-group relationships. These results challenge current concepts of Gelechioidea, suggesting that traditional morphological characters that have united taxa may not be homologous structures and are in need of further investigation. A combination of morphological data with new molecular data will be the most robust method of study for Gelechioidea phylogenetics.     

Squamate phylogeny and the relationships of snakes and mosasauroids

Cladistic analysis of extant and fossil squamates (95 characters, 26 taxa) finds the fossil squamate, Coniasaurus Owen, 1850, to be the sister-group of the Mosasauroidea (mosasaurs and aigialosaurs). This clade is supported in all 18 shortest cladograms (464 steps; CI 0.677; HI 0.772) by nine characters of the dermatocranium, maxilla, and mandible. A Strict Consensus Tree of the 18 shortest trees collapses to a basal polytomy for most major squamate clades including the clade (Coniasaurus, Mosasauroidea). A Majority Rule Consensus Tree shows that, in 12 of 18 shortest cladograms, the clade Coniasaurus- Mosasauroidea is the sister-group to snakes (Scolecophidia (Alethinophidia, Dinilysia); this entire clade, referred to as the Pythonomorpha ([[Scolecophidia [Alethinophidia, Dinilysia]], [Coniasaurus, Mosasauroidea]]) is the sister-group to all other scleroglossans. Pythonomorpha is supported in these 12 cladograms by nine characters related to the lower jaw and cranial kinesis. In 6 of 18 shortest cladograms, snakes are the sister-group to the clade (Amphisbaenia (Dibamidae (Gekkonoidea, Eublepharidae))). None of the cladograms support the hypothesis that coniasaurs and mosasauroids are derived varanoid anguimorphs. Two additional analyses were conducted: (1) manipulation and movement of problematic squamate clades while constraining ‘accepted’ relationships; (2) additional cladistic analyses beginning with extant taxa, and sequentially adding fossil taxa. From Test I, at 467 steps, Pythonomorpha can be the sister-group to the Anguimorpha, Scincomorpha, ‘scinco-gekkonomorpha’ [scincomorphs, gekkotans, and amphibaenids-dibamids]. At 471 steps Pythonomorpha can be placed within Varanoidea. Treating only mosasauroids and coniasaurs as a monophyletic group: 469 steps, mosasauroids and coniasaurs as sister-group to Anguimorpha; 479 steps, mosasauroids and coniasaurs nested within Varanoidea. Test II finds snakes to nest within Anguimorpha in a data set of only Mosasauroidea + Extant Squamates; the sistergroup to snakes + anugimorphs is (Amphisbaenia (Dibarnidae (Gekkonoidea, Eublepharidae))). No one particular taxon is identified as a keystone taxon in this analysis, though it appears truc that fossil taxa significantly alter the structure of squamate phylogenetic trees.