Phylogeny,classification and biogeography of bombyliine bee flies (Diptera,Bombyliidae)

The Bombyliinae comprises over 1100 described species in 73 known genera distributed worldwide. It is one of the largest subfamilies of bee flies (Diptera: Bombyliidae). We present the first phylogenetic hypothesis for this subfamily, based on 157 adult morphological characters scored for 123 species representing 60 genera, including all the tribes of Bombyliinae, and the related subfamilies Lordotinae and Toxophorinae. Four most parsimonious trees were generated from our analysis under equal weighting schemes. The monophyly of Bombyliinae is supported, and Lordotinae is sister to the Bombyliinae. Within Bombyliinae, Conophorini is sister to the remaining tribes. Five previously recognized tribes are revised and four new tribes are erected. We placed almost all genera in our tribal classification, based on our phylogenetic results and available character evidence. The genus Parabombylius is proposed as a synonym of Bombylius. The Gondwanan origin for the major lineages of Bombyliinae is strongly indicated by our biogeographic analysis which reconstructs ancestral areas. This published work has been registered in ZooBank, http://zoobank.org/urn:lsid:zoobank.org:pub: 1EC5C827‐34D5‐4A95‐BA78‐4ACF457F6D40.     

A new milliped of the Genus Chonodesmus,with a proposed reclassification of the family Cryptodesmidae (Diplopoda,Polydesmida)

Chonodemus gervaisi, n. sp., is described from San Agustín, Colombia. The genus Chonodesmus in many ways provides a link between the three nominal families Cryptodesmidae, Pterodesmidae, and Peridontodesmidae, and it is proposed that the three be combined into a single taxon, Cryptodesmidae, with the other two names retained for subordinate groups. A provisional classification of the family is suggested, including generic synonymy, and the new tribes Lampodesmini, Ophrydes‐mini, Dyakryptini, and Trichopeltini are proposed. The family name Otodesmidae is reduced to subfamily rank, and the name Niponiellidae reduced to tribal status. A key is provided for four West African genera of the subfamily Pterodesminae.     

A molecular phylogeny of the Trichogrammatidae (Hymenoptera: Chalcidoidea), with an evaluation of the utility of their male genitalia for higher level classification

Abstract A molecular phylogeny of the Trichogrammatidae (Hymenoptera: Chalcidoidea) is presented. This group of minute egg parasitoids is known from a broad range of host insects. The phylogeny produced, the first of any kind for the family, utilizes 121 taxa in fifty‐two of the eighty‐four recognized genera. Results were inferred from the ribosomal RNA regions 18S, 28S‐D2 and 28S‐D3, all aligned according to secondary structure models. Parsimony analysis was performed on both a complete and reduced dataset, in which ambiguous regions as defined by secondary structure were eliminated. The reduced dataset produced a much less resolved phylogenetic hypothesis. Only the complete dataset was utilized for Bayesian and maximum likelihood analyses. A robust‐choice Bayesian hypothesis stemmed from the concatenation of five distinct character set parameters. The results are compared with the current classification based primarily on male genitalia. Although our conclusions are partially congruent with the accepted hypothesis of trichogrammatid relationships, none of the currently adopted assemblages was recovered as monophyletic. Nevertheless, the structure of the male genitalia does correspond with relative taxon position in the molecular hypotheses. In general, the greatest genitalic simplification and fusion characterizes taxa treated as the most derived in the molecular hypotheses. Several groups are consistently recovered, but relationships between these groups and other genera vary with the analytical method. A new classification of the Trichogrammatidae is proposed. It includes one tribe, Trichogrammatini, within the subfamily Trichogrammatinae, and three tribes, Paracentrobiini, Chaetostrichini and Oligositini, within the Oligositinae. All tribes are more narrowly defined than previously and many genera are treated as incertae sedis within each subfamily. The results are interpreted in the light of morphological evidence, and the placement of genera not represented in molecular analysis is inferred by morphology alone. Host associations throughout the family are varied, although the molecular results suggest that Trichogrammatidae primitively parasitized Coleoptera, with more recent radiations onto other insect orders, such as Lepidoptera and Hemiptera.     

The monophyly of the subfamily Perisphaeriinae (Dictyoptera: Blattaria: Blaberidae)

A new definition of the subfamily Perisphaeriinae is given on the basis of five synapomorphies described from male and female genitalia and head morphology. The subfamily comprises eighteen genera: Bantua , Compsagis , Cyrtotria , Derocalymma , Ellipsica , Elliptoblatta , Gymnonyx , Hostilia , Laxta , Neolaxta , Perisphaeria , Perisphaerus , Pilema , Platysilpha , Poeciloblatta , Pseudoglomeris , Trichoblatta , Zuluia . Two genera are newly assigned to the Perisphaeriinae: Laxta and Neolaxta . Four genera are removed from the Perisphaeriinae: Aptera , Blepharodera , Eustegasta , Isoniscus . One new generic synonymy is proposed: Pronaonota as a synonym of Pilema . The subfamily has a large geographical range including tropical Africa, the Arabian Peninsula, Asia and Australia.     

Phylogenetic relationships of subfamilies and circumscription of tribes in the family Hesperiidae (Lepidoptera: Hesperioidea)

A comprehensive tribal‐level classification for the world’s subfamilies of Hesperiidae, the skipper butterflies, is proposed for the first time. Phylogenetic relationships between tribes and subfamilies are inferred using DNA sequence data from three gene regions (cytochrome oxidase subunit I‐subunit II, elongation factor‐1α and wingless). Monophyly of the family is strongly supported, as are some of the traditionally recognized subfamilies, with the following relationships: (Coeliadinae + (“Pyrginae” + (Heteropterinae + (Trapezitinae + Hesperiinae)))). The subfamily Pyrginae of contemporary authors was recovered as a paraphyletic grade of taxa. The formerly recognized subfamily Pyrrhopyginae, although monophyletic, is downgraded to a tribe of the “Pyrginae”. The former subfamily Megathyminae is an infra‐tribal group of the Hesperiinae. The Australian endemic Euschemon rafflesia is a hesperiid, possibly related to “Pyrginae” (Eudamini). Most of the traditionally recognized groups and subgroups of genera currently employed to partition the subfamilies of the Hesperiidae are not monophyletic. We recognize eight pyrgine and six hesperiine tribes, including the new tribe Moncini. © The Willi Hennig Society 2008.     

Carboniferous bryozoans of the order Rhabdomesida of the East European Platform

This paper deals with the revision of Carboniferous bryozoans of the order Rhabdomesida from the East European Platform. Based on evolutionary and morphological analysis and on astogenetic method, the criteria of classification are refined and the taxonomic system of Carboniferous rhabdomesids of this region is studied in more detail. The history of studying rhabdomesids, their taphonomy and paleoecology, morphology and classification, and the dynamics of the diversity and phylogenetic relationships of rhabdomesid genera and species are discussed. A phenomenon of polymerization in the development of the subfamily Ascoporinae Gorjunova subfam. nov. has been discovered for the first time. Sixty-eight rhabdomesid species (31 new), 24 genera (8 new), belonging to six subfamilies (one new), eight familie, and three suborders are described.     

Phylogenetic analysis of the family Ariidae (Ostariophysi: Siluriformes), with a hypothesis on the monophyly and relationships of the genera

Ariid monophyly and intrafamilial relationships are investigated based on cladistic analysis of 230 morphological characters. Terminal taxa examined include whenever possible type‐species, or the most morphologically similar species to the type‐species of the nominal genera, and the largest possible number of species, including cleared and stained specimens, available in zoological collections. Previous hypotheses about monophyly of the Ariidae are strongly corroborated by new synapomorphies discovered in the present study. The subfamily Galeichthyinae and the remaining ariids are strongly supported by new morphological characters. The monotypic subfamily Bagreinae is recognized as the sister group to all nongaleichthyin ariids, supported by a large series of exclusive synapomorphies. A new concept of Ariinae is presented: the subfamily is found to be unequivocally monophyletic and includes all ariid genera, except Galeichthys and Bagre. New data supporting the monophyly of the genera included in the Ariinae are introduced and previous hypotheses of monophyly, species composition, morphological definition, and relationships are reviewed and discussed.     

Phylogeny of the North American vaejovid scorpion subfamily Syntropinae Kraepelin, 1905, based on morphology,mitochondrial and nuclear DNA

The first rigorous analysis of the phylogeny of the North American vaejovid scorpion subfamily Syntropinae is presented. The analysis is based on 250 morphological characters and 4221 aligned DNA nucleotides from three mitochondrial and two nuclear gene markers, for 145 terminal taxa, representing 47 species in 11 ingroup genera, and 15 species in eight outgroup genera. The monophyly and composition of Syntropinae and its component genera, as proposed by Soleglad and Fet, are tested. The following taxa are demonstrated to be para‐ or polyphyletic: Smeringurinae; Syntropinae; Vaejovinae; Stahnkeini; Syntropini; Syntropina; Thorelliina; Hoffmannius; Kochius; and Thorellius. The spinose (hooked or toothed) margin of the distal barb of the sclerotized hemi‐mating plug is demonstrated to be a unique, unambiguous synapomorphy for Syntropinae, uniting taxa previously assigned to different subfamilies. Results of the analysis demonstrate a novel phylogenetic relationship for the subfamily, comprising six major clades and 11 genera, justify the establishment of six new genera, and they offer new insights about the systematics and historical biogeography of the subfamily, and the information content of morphological character systems.     

A modern classification of the family Caliscelidae Amyot et Serville (Homoptera,Fulgoroidea)

Recent changes in the classification of the family Caliscelidae Amyot et Serville are summarized. All the genera are attributed to tribes and subfamilies. Data on the distribution and the number of genera and species for each tribe are presented. The subfamily Ommatidiotinae is shown to be absent in the recent New World fauna, and only the tribe Peltonotellini represents the subfamily Caliscelinae in the Nearctic and Neotropical regions.     

Diverse oxyporine rove beetles from the Early Cretaceous of China (Coleoptera: Staphylinidae)

Two new genera and species, Protoxyporus grandis gen. et sp.n. and Cretoxyporus extraneus gen. et sp.n. , are described and figured on the basis of two well‐preserved impression fossils from the Lower Cretaceous Yixian Formation of China. The two genera display a combination of traits that are diagnostic of the extant staphylinid subfamily Oxyporinae: large and prominent mandibles, distinctly separated mesocoxae, abdominal tergites III–VII each with a pair of basolateral ridges, and abdominal intersegmental membranes with ‘brick‐wall’ pattern. Protoxyporus differs from extant Oxyporus in retaining the following plesiomorphic features: an elongate intercoxal process (or longitudinal carina) between metacoxae, relatively narrowly separated mesocoxae, less developed metaventral anterior process, and long infraorbital ridges. Cretoxyporus is morphologically very similar to Oxyporus, but retains distinct elongate infraorbital ridges. The discovery of two new oxyporine genera suggests that the early oxyporines were more diverse and their evolutionary history more complicated than previously documented. This published work has been registered in ZooBank, http://zoobank.org/urn:lsid:zoobank.org:pub:11503976‐D577‐45D2‐ACCB‐8226782EF817 .     

Revised systematics and higher classification of pierid butterflies (Lepidoptera: Pieridae) based on molecular data

The butterfly family Pieridae comprises approximately 1000 described species placed in 85 genera, but the higher classification has not yet been settled. We used molecular data from eight gene regions (one mitochondrial and seven nuclear protein‐coding genes) comprising a total of ~6700 bp from 96 taxa to infer a well‐supported phylogenetic hypothesis for the family. Based on this hypothesis, we revise the higher classification for all pierid genera. We resurrect the tribe Teracolini stat. rev. in the subfamily Pierinae to include the genera Teracolus, Pinacopteryx, Gideona, Ixias, Eronia, Colotis and most likely Calopieris. We transfer Hebomoia to the tribe Anthocharidini and assign the previously unplaced genera Belenois and Dixeia to the subtribe Aporiina. Three lineages near the base of Pierinae (Leptosia, Elodina and Nepheronia + Pareronia) remain unplaced. For each of these, we describe and delineate new tribes: Elodinini Braby tribus nova, Leptosiaini Braby tribus nova and Nepheroniini Braby tribus nova. The proposed higher classification is based on well‐supported monophyletic groups and is likely to remain stable even with the addition of more data.     

Taxonomic classification of the reef coral family Mussidae (Cnidaria: Anthozoa: Scleractinia)

Molecular analyses are transforming our understanding of the evolution of scleractinian corals and conflict with traditional classification, which is based on skeletal morphology. A new classification system, which integrates molecular and morphological data, is essential for documenting patterns of biodiversity and establishing priorities for marine conservation, as well as providing the morphological characters needed for linking present‐day corals with fossil species. The present monograph is the first in a series whose goal is to develop such an integrated system. It addresses the taxonomic relationships of 55 Recent zooxanthellate genera (one new) in seven families (one new), which were previously assigned to the suborder Faviina (eight genera are transferred to incertae sedis). The present monograph has two objectives. First, we introduce the higher‐level classification system for the 46 genera whose relationships are clear. Second, we formally revise the taxonomy of those corals belonging to the newly discovered family‐level clade (restricted today to the western Atlantic and Caribbean regions); this revised family Mussidae consists of ten genera (one of which is new) and 26 species that were previously assigned to the ‘traditional’ families Faviidae and Mussidae. To guide in discovering morphologic characters diagnostic of higher‐level taxa, we mapped a total of 38 morphologic characters [19 macromorphology, eight micromorphology, 11 microstructure] onto a molecular tree consisting of 67 species [22 Indo‐Pacific and seven Atlantic species in the traditional family Faviidae; 13 Indo‐Pacific and ten Atlantic species in the traditional family Mussidae; 13 species in the traditional families Merulinidae (5), Pectiniidae (7), and Trachyphylliidae (1); two Atlantic species of traditional Montastraea], and trace character histories using parsimony. To evaluate the overall effectiveness of morphological data in phylogeny reconstruction, we performed morphology‐based phylogenetic analyses using 27 (80 states) of the 38 characters, and compared morphological trees with molecular trees. The results of the ancestral state reconstructions revealed extensive homoplasy in almost all morphological characters. Family‐ and subfamily‐level molecular clades [previously identified as XVII?XXI] are best distinguished on the basis of the shapes of septal teeth and corresponding microstructure. The newly revised family Mussidae (XXI) has septal teeth with regular pointed tips (a symplesiomorphy) and a stout blocky appearance. It has two subfamilies, Mussinae and Faviinae. The subfamily Mussinae is distinguished by spine‐shaped teeth and widely spaced costoseptal clusters of calcification centres. The subfamily Faviinae is distinguished by blocky, pointed tricorne or paddle‐shaped teeth with elliptical bases, transverse structures such as carinae that cross the septal plane, and well‐developed aligned granules. Defining diagnostic characters for the broader data set is more challenging. In analyses of taxonomic subsets of the data set that were defined by clade, morphological phylogenetic analyses clearly distinguished the families Mussidae (XXI) and Lobophylliidae (XIX), as well as the two subfamilies of Mussidae (Mussinae, Faviinae), with one exception (Homophyllia australis). However, analyses of the entire 67‐species data set distinguished the family Lobophylliidae (XIX), but not the Merulinidae (XVII) and not the newly defined Mussidae (XXI), although the subfamily Mussinae was recovered as monophyletic. Some lower‐level relationships within the Merulinidae (XVII) agree with molecular results, but this particular family is especially problematic and requires additional molecular and morphological study. Future work including fossils will not only allow estimation of divergence times but also facilitate examination of the relationship between these divergences and changes in the environment and biogeography. Published 2012. This article is a U.S. Government work and is in the public domain in the USA. Zoological Journal of the Linnean Society, 2012, 166 , 465–529.     

Loxocaudinae: A new subfamily in the ostracod family loxoconchidae

A new Ostracoda subfamily, Loxocaudinae subfam. nov., separated from the subfamily Loxoconchinae of the family Loxoconchidae with five genera (Loxocauda Schornikov, 1969, Glacioloxoconcha Hartmann, 1990, Phlyctocythere Keij, 1958, Pseudoloxoconcha Müller, 1894 and Sarmatina Stancheva, 1984) is described in the paper. The new subfamily differs from Loxoconchinae Sars, 1926 s. str. by the presence of a compact eye, the absence of an eye tubercle, an originally adont hinge with a tendency to formation of an anterior tooth on the left valve and a posterior one on the right valve, a pronounced caudal process and reduction of fossa-mural sculpture. The volume of the genera that are included in the subfamily is determined. A review of 55 various species and forms mentioned in the literature in open nomenclature, whose taxonomic position has not been determined yet is presented. A total of 45 species close to Loxocauda are attributed to the new subfamily, but proper morphological investigations are necessary for their classification. Ten species that were referred earlier to the genera included in Loxocaudinae are excluded from this subfamily. Issues of functional morphology of Loxocaudinae shells and morphological evolution of their sculpture are considered.     

Further revision of the caddisfly family Uenoidae (Trichoptera): evidence for inclusion of Neophylacinae and Thremmatidae

Abstract. A survey of characters within the Limnephilidae was made to test the cladistic hypothesis underlying traditional classification placing Neophylacinae as a subfamily of the Limnephilidae. Characters emerged which support an alternative hypothesis of phylogeny and classification, linking the Neophylacinae with the Thremmatidae and with the Uenoidae. The Uenoidae as defined here contain two monophyletic groups: subfamily Uenoinae containing the four genera Uenoa, Sericostriata, Neothremma and Farula previously constituting Uenoidae ( sensu Wiggins et al. , 1985), and subfamily Thremmatinae consisting of Thremma, Neophylax and Oligophlebodes. The new classification is shown to have predictive value for behaviour and ecology of the taxa.     

Flightless scaly‐tailed squirrels never learned how to fly: A reappraisal of Anomaluridae phylogeny

Anomaluroidea, commonly known as the “scaly‐tailed squirrels,” are an emblematic group of tropical African mammals that includes gliding forms. The family Anomaluridae was until recently represented by three genera: the flying scaly‐tailed squirrels (Anomalurus), the flying mouse (Idiurus) and the flightless scaly‐tailed squirrels (Zenkerella). Idiurus and Zenkerella have long been grouped into the Zenkerellinae subfamily, and Zenkerella was interpreted as a rare case of evolutionary reversal to non‐gliding lifestyle. Recent studies have demonstrated that Zenkerella is sister to all other modern anomalurids, and represents in fact the monogeneric family Zenkerellidae. The Anomalurus genus was split into Anomalurus and Anomalurops, but no study has ever considered all Anomalurus species together in a phylogeny to test the status of Anomalurops. Here, we used mitogenomic next‐generation sequencing to infer the phylogenetic relationships among all extant anomalurids and to estimate their divergence ages. We found that the arboreal Zenkerella is the sister group of all extant gliding anomalurids (Idiurus and Anomalurus). We confirmed that Anomaluroidea only evolved the gliding adaptation once. A comparison based on morphological traits indicates that Zenkerella harbours several unique morphological features. We propose new morphological characters for the novel classification of modern Anomaluroidea, which includes the families Zenkerellidae and Anomaluridae. Using different calibration schemes, we demonstrated that classical dating methods relying only on mitogenomes provide rather young Miocene estimates between Zenkerellidae and the Anomaluridae. The use of published nuclear genes, internal calibrations and tip dating converged towards an Eocene split between gliding and non‐gliding scaly‐tailed squirrels, which is in agreement with the African fossil record. Finally, we provide the first exhaustive species‐level molecular phylogenetic inference for the genus Anomalurus. We found that Anomalurus beecrofti is the sister group of all other species of Anomalurus and branched off during the Miocene.     

Pollen morphology of the Dimorphandra group (Leguminosae,Caesalpinioideae)

The Dimorphandra group, as traditionally circumscribed, is a rather diverse assemblage of genera in Leguminosae subfamily Caesalpinioideae that share certain morphological characteristics with the basally branching lineages of subfamily Mimosoideae. It currently comprises 51 species in seven genera: Burkea (1 species), Dimorphandra (26 species), Erythrophleum (10 species), Mora (6 species), Pachyelasma (1 species), Stachyothyrsus (2 species) and Sympetalandra (5 species). This study investigates the pollen morphology of 25 samples from 19 species of all seven genera. Pollen of the Dimorphandra group is small, isopolar, trizonocolporate and released in monads. Apertures are almost equal to polar length, with correspondingly small apocolpial areas. The shape of the aperture apices varies from acute to wide and rounded. Surface ornamentation is psilate, perforate, microreticulate, or perforate‐rugulate. The wall structure is usually columellate with a well developed foot layer. The pollen is small and unspecialised, agreeing with a previously noted pattern of more fixed and homogenous pollen structure in the more derived clades in subfamily Caesalpinioideae, compared with the great diversity of pollen types found in the basally branching lineages.