The marine midge Pontomyia (Chironomidae) with a description of females of P.oceana Tokunaga

A key is provided to both sexes of the four species of Pontomyia; cottoni Womersley, natans Edwards, oceana Tokunaga, pacifica Tokunaga. This genus is widespread in the western Pacific Ocean. Adult females are vermiform. Adult males have normal halteres but reduced wings; they are unusual among Chironomidae in having bare, not plumose antennae.     

Resurrection in Carapa (Meliaceae): a reassessment of morphological variation and species boundaries using multivariate methods in a phylogenetic context

The taxonomy of the amphi‐Atlantic tree genus Carapa (Meliaceae) has long been controversial. Of the three species currently recognized in the genus, two are known to present substantial morphological variation that has been used in the past to distinguish several taxa, most of which are currently placed in synonymy. Here, a combination of field observations, univariate analyses of leaf, floral and seed characters and principal coordinate analyses of floral characters in the context of a molecular phylogenetic analysis was used to investigate the patterns of variation and delimit morphological species anew in the genus. These results support the recognition of 27 species in Carapa, of which 16 are previously described and 11 are new. In general, phylogenetically related species occurred in the same geographical area, but were morphologically distinct. © 2011 The Linnean Society of London, Botanical Journal of the Linnean Society, 2011, 165 , 186–221.     

Systematics of a widespread Southeast Asian frog,Rana chalconota (Amphibia: Anura: Ranidae)

The abundant Sundaland forest frog, Rana chalconota, has long been considered a single widespread species, although some authors have recommended its division into regional subspecies. The discovery of co‐occurring pairs of morphologically distinct populations in three widely separated parts of the range led to a morphological and molecular analysis of populations from all parts of the known range. The results suggest that R. chalconota consists of at least seven species from Thailand through Borneo and Java. Existing names are applied to three of these species, R. chalconota (Schlegel), R. raniceps (Peters) and R. labialis Boulenger. We describe four others as new species and suggest the existence of one or two additional, unnamed species. © 2009 The Linnean Society of London, Zoological Journal of the Linnean Society, 2009, 155 , 123–147.     

Systematics of the Miocene–Recent bryozoan genus Pentapora (Cheilostomata)

The four existing species of the ascophoran bryozoan Pentapora Fisher, 1807 are revised, and two new fossil species are introduced: Pentapora lacryma sp. nov. from the Pliocene Coralline Crag Formation of Suffolk, and Pentapora clipeus sp. nov. from the Pliocene of Emilia, Italy. The Arctic species Pentapora boreale Kuklinski & Hayward possesses a lyrula, does not belong in Pentapora, and is a junior synonym of Raymondcia rigida (Lorenz). The morphology of the autozooids is relatively uniform within the genus, and the main distinguishing characters are those of the ovicells and, particularly, the giant avicularia that are developed sporadically in all species apart from Pentapora foliacea, popularly known as ‘Ross coral’. A phylogenetic analysis based on skeletal characters returned a single shortest tree in which the three species of Pentapora from the North Atlantic (P. foliacea, Pentapora pertusa, and P. lacryma sp. nov. ) form a clade crownward of the three basal species from the Mediterranean (Pentapora ottomulleriana, Pentapora fascialis, and P. clipeus sp. nov. ). © 2010 The Linnean Society of London, Zoological Journal of the Linnean Society, 2010, 160 , 17–39.     

Systematics of the scimitar babblers (Pomatorhinus: Timaliidae): phylogeny,biogeography, and species‐limits of four species complexes

Monophyly of the scimitar babblers (Pomatorhinus, Xiphirhynchus: Timaliidae), traditionally defined by the characteristic of having long, curved bills, has been questioned by recent molecular phylogenetic results. We examined representatives of all scimitar babbler species complexes, including all distinct lineages of four complexes as well as several potential relatives, and corroborate that Xiphirhynchus and some species of Stachyris group within Pomatorhinus. Pomatorhinus species comprise three separate clades: larger scimitar babblers (Pomatorhinus hypoleucos, Pomatorhinus erythrogenys complexes); orange‐ and coral‐billed scimitar babblers (Pomatorhinus ochraceiceps, Pomatorhinus ferruginosus complexes); and small scimitar babblers (Pomatorhinus schisticeps, Pomatorhinus ruficollis, Pomatorhinus horsfieldii, Pomatorhinus montanus complexes). Additionally, at least two of the traditional species complexes are not monophyletic. Lineages of the ruficollis and schisticeps groups are intertwined, and P. montanus and P. horsfieldi group within the schisticeps complex. Upon revision of four traditional species complexes, P. hypoluecos, P. ferruginosus, P. schisticeps, and P. ruficollis (with 41 subspecies described in total), 27 distinct, independent lineages or phylogenetic species were distinguished. Two contrasting biogeographical patterns are evident in these groups: Sino‐Himalayan areas are either sister to south‐east Asian areas or are embedded within a clade of other Asian areas. The present study demonstrates the need for unraveling the confusion in traditional taxonomy to allow the study of complex biodiversity patterns in tropical Asia. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 102 , 846–869.     

Genetic diversity,endemism and phylogeny of lampreys within the genus Lampetra sensu stricto (Petromyzontiformes: Petromyzontidae) in western North America

Phylogenetic structure of four Lampetra species from the Pacific drainage of North America (western brook lamprey Lampetra richardsoni, Pacific brook lamprey Lampetra pacifica, river lamprey Lampetra ayresii and Kern brook lamprey Lampetra hubbsi) and unidentified Lampetra specimens (referred to as Lampetra sp.) from 36 locations was estimated using the mitochondrial cytochrome b gene. Maximum parsimony and Bayesian inferences did not correspond with any taxonomic scheme proposed to date. Rather, although L. richardsoni (from Alaska to California) and L. ayresii (from British Columbia to California) together constituted a well‐supported clade distinct from several genetically divergent Lampetra populations in Oregon and California, these two species were not reciprocally monophyletic. The genetically divergent populations included L. pacifica (from the Columbia River basin) and L. hubbsi (from the Kern River basin) and four Lampetra sp. populations in Oregon (Siuslaw River and Fourmile Creek) and California (Kelsey and Mark West Creeks). These four Lampetra sp. populations showed genetic divergence between 2·3 and 5·7% from any known species (and up to 8·0% from each other), and may represent morphologically cryptic and thus previously undescribed species. A fifth population (from Paynes Creek, California) may represent a range extension of L. hubbsi into the Upper Sacramento River.     

Osteology of the South Asian Genus Psilorhynchus McClelland, 1839 (Teleostei: Ostariophysi: Psilorhynchidae), with investigation of its phylogenetic relationships within the order Cypriniformes

Members of the genus Psilorhynchus are small benthic fishes, commonly referred to as torrent minnows, which inhabit the fast to swift flowing water bodies of the Indo‐Burma region and the Western Ghats of Peninsular India. Despite being described scientifically in the mid 18^th century, the morphology of Psilorhynchus remains poorly known and its phylogenetic placement within the order Cypriniformes is a matter of considerable debate. In this paper the osteology of Psilorhynchus sucatio is described and illustrated in detail. Notes and/or illustrations on the osteology of 12 other species of Psilorhynchus are also provided for the first time. A phylogenetic investigation of the position of Psilorhynchus within the order Cypriniformes is also conducted. Analysis of 127 morphological characters scored for 52 ingroup taxa (including 12 species of Psilorhynchus) and four outgroup taxa resulted in 14 equally parsimonious cladograms (287 steps long; consitency index, CI = 0.48; retention index, RI = 0.88). Psilorhynchus is recovered as the sister group to the family Cyprinidae, and is regarded as a member of the superfamily Cyprinoidea, which forms the sister group to the Cobitoidea (including all other cypriniform families). The sistergroup relationship between Psilorhynchus and Cyprinidae is supported by eight derived characters (five of which are homoplastic within the order Cypriniformes). The monophyly of Psilorhynchus is supported by 16 derived characters (eight of which are homoplastic within Cypriniformes). Three species groups of Psilorhynchus are proposed, the Psilorhynchus balitora group (including P. amplicephalus, P. balitora, P. breviminor, P. nepalensis, P. rahmani, P. pavimentatus, and P. brachyrhynchus), the Psilorhynchus gracilis group (including P. gracilis, P. melissa, P. robustus, and P. tenura), and the Psilorhynchus homaloptera group (including P. arunachalensis, P. homaloptera, P. microphthalmus, and P. pseudecheneis). The continued use of the family group name Psilorhynchidae is recommended. Comments on the interrelationships of the Cypriniformes are also provided. © 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011.     

A multivariate morphometric analysis and systematic review of Pseudonaja (Serpentes,Elapidae, Hydrophiinae)

Pseudonaja is a clade of seven nominal species of elapid snakes distributed throughout Australia and in southern New Guinea. The species‐level systematics of this group is generally considered to be problematic. A recent phylogenetic analysis of mitochondrial DNA sequences for a geographically extensive series of Pseudonaja specimens revealed nine major clades, of which six largely coincide with nominal species (P. affinis, P. guttata, P. inframacula, P. ingrami, P. modesta and P. textilis). The three remaining clades are composed of specimens currently referred to P. nuchalis. This paper presents a multivariate analysis of 30 morphometric variables recorded for 220 specimens, representing the P. affinis, P. inframacula, P. textilis and three P. nuchalis clades (P. guttata, P. ingrami and P. modesta are well‐demarcated species and, accordingly, were not considered). The morphometric data readily separate these putative lineages, affording compelling evidence that they constitute evolutionary species. The names aspidorhyncha and mengdeni are resurrected for two of the three species presently recognized as P. nuchalis. These species, P. affinis, P. inframacula, P. nuchalis and P. textilis are redescribed. © 2009 The Linnean Society of London, Zoological Journal of the Linnean Society, 2009, 155 , 171–197.     

An integrative approach to the taxonomy of the pigmented European Pseudomonocelis Meixner, 1943 (Platyhelminthes: Proseriata)

Only a few species belonging to the Proseriata (Platyhelminthes) show a parenchymatic pigmentation, which may aid identification. Among these, Pseudomonocelis agilis has a yellowish body and is provided with a reddish–brown girdle in front of the statocyst. The species is known for limited areas of northern Europe and the Mediterranean. The present study was conducted to assess both the taxonomic status of populations attributed to the species across the unusually wide range for an interstitial flatworm, which lacks an obvious means of dispersal, and the levels of genetic variability within and among populations, by employing an integrative approach that included the analyses, on six populations, of three molecular markers (small subunit ribosomal 18S‐like gene, inter‐simple sequence repeat, allozymes), karyotypes, and 11 morphological characters. Furthermore, crossbreeding experiments were carried out on the Mediterranean populations. The results obtained revealed the existence of four highly divergent genotypic clusters, accompanied by karyological differences, with complete intersterility among the clusters tested. The combination of approaches adopted strongly supports the conclusion that the wide‐ranging European pigmented species P. agilis is actually composed of four species: P. agilis in the Baltic area; Pseudomonocelis cetinae in the Adriatic; and Pseudomonocelis sp. nov. A and Pseudomonocelis sp. nov. B in the western and eastern Mediterranean, respectively. The latter two species are morphologically indistinguishable for the parameters essayed. Reconstruction of the phylogenetic relationships of these taxa, including congeneric and consubfamilial outgroups, showed that pigmentation is a plesiomorphic condition for the genus Pseudomonocelis and that Pseudomonocelis sp. nov. A shares a previously undetected, sister‐group relationship with species of the unpigmented P. ophiocephala complex. The present study thus depicts complex speciation processes in a mesopsammic species, which involves allopatric divergence operating on different scales and ecological shifts, and highlights that the contribution of microturbellarians to marine biodiversity may be seriously underestimated. © 2009 The Linnean Society of London, Biological Journal of the Linnean Society, 2009, 98 , 907–922.     

With eyes wide open: a revision of species within and closely related to the Pocillopora damicornis species complex (Scleractinia; Pocilloporidae) using morphology and genetics

Molecular studies have been instrumental for refining species boundaries in the coral genus Pocillopora and revealing hidden species diversity within the extensively studied global species Pocillopora damicornis. Here we formally revise the taxonomic status of species closely related to and within the P. damicornis species complex, taking into account both genetic evidence and new data on morphometrics, including fine‐scale corallite and coenosteum structure. We found that mitochondrial molecular phylogenies are congruent with groups based on gross‐morphology, therefore reflecting species‐level differentiation. However, high levels of gross morphological plasticity and shared morphological characteristics mask clear separation for some groups. Fine‐scale morphological variation, particularly the shape and type of columella, was useful for differentiating between clades and provides an excellent signature of the evolutionary relationships among genetic lineages. As introgressive hybridization and incomplete lineage sorting complicate the delineation of species within the genus on the basis of a single species concept, the Unified Species Concept may represent a suitable approach in revising Pocillopora taxonomy. Eight species are herein described (P. damicornis, P. acuta, P. aliciae, P. verrucosa, P. meandrina, P. eydouxi, P. cf. brevicornis), including a novel taxon – P ocillopora bairdi sp. nov. (Schmidt‐Roach, this study). Citation synonyms and type materials are presented. © 2014 The Linnean Society of London     

Discovery of cryptic species among North American pine‐feeding Chionaspis scale insects (Hemiptera: Diaspididae)

Cryptic species are present in many animal groups and they may be best detected through large sample sizes collected over broad geographic ranges. Fine‐scale local adaptation has been hypothesized to occur in armoured scale insects (Hemiptera: Diaspididae) and a consequence of this process may be multiple cryptic species. We estimate species diversity of pine‐feeding Chionaspis scale insects across North America by inferring species boundaries using genealogical concordance across allele genealogies of two nuclear loci and one mitochondrial locus. Our ingroup sample includes 366 individual insects from 320 localities and 51 host species within the Pinaceae. We also conducted a morphological survey of all insect specimens and assigned them as either one of the two currently recognized pine‐feeding species, Ch. heterophyllae, Ch. Pinifoliae, or with undescribed morphology. Using maximum likelihood allele genealogies in a majority‐rule consensus to assess congruence, we conservatively detect ten species in this group. Most of these species are robust to alternative methods of genealogical inference (Bayesian inference, maximum parsimony) and a few are robust to the use of strict consensus to assess congruence. Species show both narrow and more widespread ranges where almost half of the individuals sampled belong to a single very widespread polyphagous species. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 104 , 47–62.     

Revision and phylogenetic assessment of the rove beetle genus Pseudohesperus Hayashi,with broad reference to the subtribe Philonthina (Coleoptera: Staphylinidae: Staphylinini)

This paper studies the phylogeny of the rove beetle subtribe Philonthina, to test its hypothetical monophyly and to unravel the evolutionary relationships of the subtribe and its included genus‐level taxa, with emphasis on the genus Pseudohesperus and its close‐allied relatives. The phylogenetic analyses are based on 105 adult morphological characters and 66 terminal taxa, i.e., all six members of Pseudohesperus, 51 species to represent 29 other genera of the subtribe Philonthina, seven species to represent the other six subtribes of Staphylinini, one species of the tribes Arrowinini, and one of the Platyprosopini. According to the phylogenetic results obtained, the genus Erichsonius should move out from the hitherto‐defined subtribe Philonthina and thus the monophyly of this taxon is challenged. The phylogenetic tree suggests that the genera Hesperus and Belonuchus might not be monophyletic, but the monophyly of Pseudohesperus and the sister relationship between it and Bisnius are well supported. The species‐level phylogenetic relationships of the genus Pseudohesperus reveal a clear pattern of species diversification that can be correlated well with the species' zoogeographical patterns. The paper also revises the taxonomy of Pseudohesperus and describes five new species from China: Pseudohesperus luteus Li & Zhou sp. nov. , Pseudohesperus pedatiformis Li & Zhou sp. nov. , Pseudohesperus tripartitus Li & Zhou sp. nov. , Pseudohesperus sparsipunctatus Li & Zhou sp. nov. , and Bisnius lubricus Li & Zhou sp. nov. An identification key to the species of Pseudohesperus is provided and their geographical distributions are mapped. © 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 163 , 679–722.     

Phylogenetic interrelationships,taxonomy, and reductive evolution in the Neotropical electric fish genus Hypopygus (Teleostei,Ostariophysi, Gymnotiformes)

A phylogenetic reconstruction of the Neotropical electric fish genus Hypopygus based on 47 parsimony‐informative morphological characters is presented. A series of synapomorphies support the hypothesis of monophyly of Hypopygus, and partially resolve species‐level relationships within the genus. Hypopygus species are recognized here as miniaturized fishes based on two criteria; first, a derived condition of diminutive body size, and; second, the presence of a suite of reductive morphological characters, including partial or total losses, simplifications, and reductions of the anal‐fin rays, scales, cranial bones, and laterosensory canal system. Reductive characters associated with miniaturization comprise 45% of the total number of characters in the phylogenetic reconstruction of the genus. Miniaturization and reductive morphological evolution in Hypopygus are discussed here in the phylogenetic context. A taxonomic revision of Hypopygus is presented, in which five new species are described, two species previously assigned to the genus are redescribed, and a single known species of Stegostenopos is redescribed and included in Hypopygus as a junior synonym. Distribution maps and a key for all eight valid species of Hypopygus are provided, based on the examination of 5014 catalogued museum specimens. © 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 163 , 1096–1156.     

World revision of Dolichocolon Brauer & Bergenstamm (Diptera: Tachinidae: Exoristinae: Goniini)

The Palaeotropical goniine genus Dolichocolon Brauer & Bergenstamm is revised and analysed cladistically. Seventeen new species are described from Australia (Queensland, Northern Territory), Cameroon, China, the Democratic Republic of Congo, Ethiopia, Japan, Papua New Guinea, Senegal, Thailand, Uganda, Yemen, and Zimbabwe. The following new synonymy is proposed after direct comparison of primary types: Dolichocolon klapperichi Mesnil, 1967 = Dolichocolon orientale Townsend, 1927 syn. nov. A key to the 21 known species is presented. A cladistic analysis based on 36 morphological characters provides support for the monophyly of Dolichocolon. A sister‐group relationship is indicated between Dolichocolon and Kuwanimyia Townsend, whereas Dolichocolon chiangmaiensis sp. nov. from Thailand takes a position as sister group to all other Dolichocolon species. © 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 162 , 544–584.     

Revision of the New World genus Quichuana Knab, 1913 (Diptera: Syrphidae), including descriptions of 24 new species

The species of the New World syrphid genus, Quichuana Knab, 1913 (Diptera: Syrphidae: Eristalini), are revised and the genus is re‐diagnosed. Twenty‐four new species are described, bringing the total number considered valid to 48. New species were mostly reared from Bolivia, Brazil, Costa Rica, El Salvador, Mexico, Peru, Trinidad, and Venezuela. The male of Quichuana picadoi is described for the first time. Quichuana championi is proposed as a synonym of Quichuana cincta, Quichuana aurata as a synonym of Quichuana angustiventris, and Quichuana sepiapennis as a synonym of Quichuana calathea. Quichuana inca var. brevicera is raised to specific status. Lectotypes are designated for Helophilus auratus, Quichuana bezzii, Quichuana fasciata, and Quichuana parisii. A species‐level identification key for both males and females is provided. Male genitalia of 18 species are illustrated. © 2012 The Linnean Society of London, Zoological Journal of the Linnean Society, 2012, 166 , 72–131.     

The fern genus Elaphoglossum section Lepidoglossa (Dryopteridaceae) in Africa,Macaronesia, the mid‐Atlantic and southern Indian Ocean Islands

Elaphoglossum section Lepidoglossa in Africa, Macaronesia (Azores and Madeira), the mid‐Atlantic Ocean Islands (St Helena, Gough and Tristan da Cunha Island groups) and the southern Indian Ocean Islands (Marion and Prince Edward Islands) is reviewed. Fifteen Elaphoglossum species from this region are ascribed to the section. A new species, Elaphoglossum rivularum , confined to the Chimanimani Mountains in eastern Zimbabwe and formerly ascribed to E. kuhnii from West Africa, is described. © 2010 The Linnean Society of London, Botanical Journal of the Linnean Society, 2011, 165 , 20–63.     

Almost but not quite a subspecies: a case of genetic but not morphological diagnosability in Nicrophorus (Coleoptera: Silphidae)

Allopatric populations that show genetic differentiation but lack phenotypic diagnosability are difficult to classify. In 1946, Arnett described a new species of burying beetle (Silphidae: Nicrophorus) from Luzon Island, Philippines, Nicrophorus benguetensis. In 2002, Sikes et al., finding the species to be inconsistently diagnosable, synonymized N. benguetensis under Nicrophorus nepalensis Hope 1831. We rigorously and quantitatively test the validity of N. benguetensis using several different species delimitation criteria. We employed discrete and quantitative character‐based methods to test similarity criteria using linear morphometrics in a discriminant analysis framework, and percentage sequence divergences based on sequences from four mitochondrial DNA (mtDNA) genes (COI, COII, ND4 and ND5) and two nuclear genes [28S (D2 region) and CAD]. We also employed tree‐based methods to test phylogenetic criteria using mtDNA sequences and morphology with parsimony, Bayesian inference and maximum likelihood. The analysis of discrete phenotypic characters did not reliably diagnose N. benguetensis. The results of the discriminant analysis provided moderate support for the validity of N. benguetensis (71.4% of the specimens were properly classified). There is adequate genetic distance between N. benguetensis and its nearest neighbour for DNA barcoding to identify an unknown sequence, although this may be an artefact of the small sample size. Phylogenetic analyses of the morphological data, with and without the morphometric data, yielded unresolved trees. Molecular phylogenetic results found N. benguetensis to be monophyletic, but neither clearly rejected nor supported its validity. Two of our approaches using molecular data diagnosed N. benguetensis, but those that used phenotypic data did not exceed a 75% success rate. We conclude, therefore, that N. benguetensis, despite being a weakly distinct allopatric population, and deserving of additional study, should remain synonymized until further analysis suggests otherwise. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 102 , 311–333.     

Repeated evolution of sympatric,palaeoendemic species in closely related,co‐distributed lineages of Hemiphyllodactylus Bleeker, 1860 (Squamata: Gekkonidae) across a sky‐island archipelago in Peninsular Malaysia

A time‐calibrated phylogenetic tree indicates that the evolution of sympatric, montane, endemic species from closely related, co‐distributed lineages of the Hemiphyllodactylus harterti group were not the result of rapid, forest‐driven, climatic oscillations of the Last Glacial Maximum, but rather the result of infrequent episodes of environmental fluctuation during the Late Miocene. This hypothesis is supported by genetic divergences (based on the mitochondrial gene ND2) between the three major lineages of the H. harterti group (17.5–25.1%), their constituent species (9.4–14.3%), and the evolution of discrete, diagnostic, morphological, and colour pattern characteristics between each species. Sister species pairs from two of the three lineages occur in sympatry on mountain tops from opposite sides of the Thai–Malay Peninsula, but the lineages to which each pair belongs are not sister lineages. A newly discovered species from Gunung Tebu, Terengganu State, H emiphyllodactylus bintik sp. nov. , is described. © 2015 The Linnean Society of London