Species delimitation in the Andean grasshopper genus Orotettix Ronderos & Carbonell (Orthoptera: Melanoplinae): an integrative approach combining morphological,molecular and biogeographical data

The reciprocal illumination nature of integrative taxonomy through hypothesis testing, corroboration and revision is a powerful tool for species delimitation as more than one source has to support the hypothesis of a new species. In this study, we applied an integrative taxonomy approach combining molecular and morphological data sets with distributional patterns to examine the level of differentiation between and within the grasshopper Orotettix species. Orotettix was described based on five valid species distributed in the Andes of Peru. In our study, initially a molecular‐based hypothesis was postulated and tested against morphological data and geographical patterns of distribution. Results from molecular and morphological analyses showed agreement among the species delimitation in Orotettix, and were also consistent with the geographical distribution. The analyses allowed us to delimit five new species for the genus ( O. lunatus sp. nov. , O . astreptos sp. nov. , O. colcaensis sp. nov. , O . paucartambensis sp. nov. and O . dichrous sp. nov. ) from the Eastern and Western Cordilleras of Peru. We also provide critical knowledge on the phylogenetic relationships and distribution of the genus and conduct a revision of Orotettix. © 2015 The Linnean Society of London     

A tale of time and depth: intralacustrine radiation in endemic Gammarus species flock from the ancient Lake Ohrid

Tentatively dated, the Plio‐/Pleistocene origin of the ancient Lake Ohrid on the Balkan Peninsula makes it the oldest ancient lake in Europe. Given the surface area of the lake and the adjusted endemicity rate, it may be also defined as the most diverse of all the ancient lakes in the world. From all the animal groups endemic to this lake, gammarids are amongst the most scarcely known in terms of their diversity and phylogenetic relationships. Partial DNA sequences of two mitochondrial genes, cytochrome oxidase subunit I (cox1) and 16S ribosomal RNA (16S rRNA) of eight known endemic Gammarus species from the Lake Ohrid valley were analysed. Phylogenetic analyses showed that endemic Gammarus species comprise an ancient species flock, with Gammarus sketi from the feeder springs being their sister taxon outside the lake. Amongst the species inhabiting the lake, Gammarus solidus and Gammarus salemaai are morphologically and molecularly well defined. By contrast, Gammarus ochridensis, Gammarus parechiniformis, Gammarus lychnidensis, and Gammarus stankokaramani revealed high discrepancy between morphological and genetic data. None of these morphospecies form a monophyletic clade and a significant degree of apparent gene flow occurs between them. This could be caused by incomplete lineage sorting and/or hybridization events. Two novel mtDNA lineages were found within the lake, possibly constituting two new species (Gammarus sp. 1 and Gammarus sp. 2). Molecular clock analysis showed that the split between G. sketi and the Gammarus species flock from the lake occurred approximately 5–7 Mya, whereas within the flock there were at least two intralacustrine radiations: one estimated at 2–3 Mya and the second at less than 1 Mya. The first one could be associated with the origin of the lake and the second with the lake water‐level fluctuations during Pleistocene. © 2013 The Linnean Society of London     

First reports of Grania (Clitellata: Enchytraeidae) from Africa and South America: molecular phylogeny and descriptions of nine new species

In this article, our knowledge of the geographic distribution of Grania species is expanded by describing seven new species, G rania bekkouchei sp. nov. , G rania brasiliensis sp. nov. , G rania capensis sp. nov. , G rania chilensis sp. nov. , G rania cryptica sp. nov. , G rania hinojosai sp. nov. , and G rania simonae sp. nov. , from poorly investigated regions of the Southern Hemisphere, plus two new species, G rania carolinensis sp. nov. and G rania unitheca sp. nov. , from off the east coast of the USA. An immature achaetous specimen that we call Grania cf. levis was also included. The newly generated data were combined with a previously published data set in order to update the hypothesis of phylogenetic relationships among Grania species. All new species except G . cryptica sp. nov. are supported by both morphological and molecular data. In addition, we find that G . chilensis sp. nov. is structured in at least four distinct populations along the Chilean coast. The species described from South Africa form a monophyletic clade where two are morphologically indistinguishable but diverging in both mitochondrial and nuclear ribosomal genes, and thus we describe them as different species, G . bekkouchei sp. nov. and G . cryptica sp. nov. Among the North Carolinian species, G . unitheca sp. nov. is indicated as a close relative of Grania monospermatheca Erséus & Lasserre, 1976, and G . carolinensis sp. nov. is indicated as a close relative of G. cf. levis. The updated phylogeny is strongly concordant with geographical species distributions, thus supporting a low level of dispersal within this genus, as has previously been hypothesized. © 2015 The Linnean Society of London     

Three new species of deep‐sea Gromia (Protista,Rhizaria) from the bathyal and abyssal Weddell Sea,Antarctica

We describe three new species of the genus Gromia from bathyal and abyssal depths in the Weddell Sea. The new species are characterized by a combination of morphological and molecular criteria. All three species possess a distinct oral capsule and a layer of ‘honeycomb membranes’, which form the inner part of the organic test wall. Both these features are typical of gromiids. Their identification as gromiids is confirmed by analyses of partial small subunit ribosomal DNA (SSU rDNA) gene sequences. Gromia marmorea sp. nov. is a rounded species with a prominent oral capsule and a characteristically mottled appearance. In Gromia melinus sp. nov. , the test surface exhibits a polygonal pattern of ridges, with a layer of clay particles coating the surface between the ridges. Gromia winnetoui sp. nov. represents an elongate morphotype in which the organic test is enclosed within an agglutinated case, a feature previously unknown in gromiids. Phylogenetic analysis using the maximum‐likelihood method revealed that all three species form distinct clades, reflecting the morphological differences among Weddell Sea species, as well as between deep‐water Southern Ocean Gromia and previously described gromiids. © 2009 The Linnean Society of London, Zoological Journal of the Linnean Society, 2009, 157 , 451–469.     

An integrative approach to characterize cryptic species in the Thoracostoma trachygaster Hope, 1967 complex (Nematoda: Leptosomatidae)

Nematode diversity may seriously be underestimated when taking into account cryptic speciation. Thoracostoma trachygaster is commonly found in kelp holdfasts along the California coastline and was recently shown to consist of at least two distinct molecular clades (I and II). Here, we provide detailed morphological analysis of both clades, based on measurements taken from video vouchers of respectively eight and 16 individuals from the previous study, as well as 80 newly collected specimens from four Californian beaches. The latter were vouchered, measured, and then subjected to molecular analyses of the mitochondrial cytochrome oxidase c subunit I (COI) gene, and the ribosomal D2D3 and internal transcribed spacer (ITS) regions. This integrative approach shows that the three molecular clades are phylogenetically and morphologically distinct species, but a combination of morphological characters is needed to distinguish them. Two new species, Thoracostoma fatimae sp. nov. and Thoracostoma igniferum sp. nov. , are identified and described. The spicule length of T. fatimae sp. nov. is significantly shorter than that of T. trachygaster. Thoracostoma igniferum sp. nov. can be distinguished by the irregular posterior edge of the cephalic capsule and the two internal subdorsal tropis‐like projections in the wall of the cephalic capsule, which are lacking in T. fatimae sp. nov. and T. trachygaster. © 2012 The Linnean Society of London, Zoological Journal of the Linnean Society, 2012, 164 , 18–35.     

Systematics of the Platyrrhinus helleri species complex (Chiroptera: Phyllostomidae), with descriptions of two new species

Platyrrhinus is a diverse genus of small to large phyllostomid bats characterized by a comparatively narrow uropatagium thickly fringed with hair, a white dorsal stripe, comparatively large inner upper incisors that are convergent at the tips, and three upper and three lower molars. Eighteen species are currently recognized, the majority occurring in the Andes. Molecular, morphological, and morphometric analyses of specimens formerly identified as Platyrrhinus helleri support recognition of Platyrrhinus incarum as a separate species and reveal the presence of two species from western and northern South America that we describe herein as new ( Platyrrhinus angustirostris sp. nov. from eastern Colombia and Ecuador, north‐eastern Peru, and Venezuela and Platyrrhinus fusciventris sp. nov. from Guyana, Suriname, French Guiana, Trinidad and Tobago, northern Brazil, eastern Ecuador, and southern Venezuela). These two new species are sister taxa and, in turn, sister to Platyrrhinus incarum. © 2010 The Linnean Society of London, Zoological Journal of the Linnean Society, 2010, 159 , 785–812.     

Morphological and molecular taxonomy of a new Daptonema (Nematoda,Xyalidae) with comments on the systematics of some related taxa

A new species of Daptonema is described based upon morphological characters and 18S rRNA sequence. Daptonema matrona sp. nov. was collected in Pina Basin (north‐eastern Brazil). It differs from all other species of the genus by the presence of reduced cephalic setae and straight spicules. These features require an adaptation of the generic diagnosis. Moreover, the females are characterized by intra‐uterine development of the offspring, considered herein as their major autapomorphic feature. Molecular systematic analyses supported Daptonema matrona sp. nov. as a distinct genetic and evolutionary lineage. The data also indicate hypotheses of taxonomic synonymies amongst some related taxa from Xyalidae as well as the paraphyly of Daptonema. © 2010 The Linnean Society of London, Zoological Journal of the Linnean Society, 2010, 158 , 1–15.     

A reanalysis of Parabuthus (Scorpiones: Buthidae) phylogeny with descriptions of two new Parabuthus species endemic to the Central Namib gravel plains,Namibia

Two new thick‐tail scorpions in the genus Parabuthus Pocock, 1890 are described from the gravel plains of the Central Namib Desert, Namibia: Parabuthus glabrimanus sp. nov. ; Parabuthus setiventer sp. nov. The two new species occupy discrete distributional ranges, allopatric with the closely related species Parabuthus gracilis Lamoral, 1979 and Parabuthus nanus Lamoral, 1979. The distributions of the four species are mapped and a key provided for their identification. Revised diagnoses are provided for P. gracilis and P. nanus. The two new species are added to a previously published morphological character matrix for Parabuthus species and their phylogenetic positions determined in a reanalysis of Parabuthus phylogeny. Parabuthus setiventer sp. nov. is found to be the sister species of P. nanus, whereas P. glabrimanus sp. nov. is sister to a monophyletic group comprising P. gracilis, P. nanus, and P. setiventer sp. nov. The discovery of two new scorpion species endemic to the Central Namib gravel plains contributes to a growing body of evidence that this barren and desolate region is a hotspot of arachnid species richness and endemism. © 2010 The Linnean Society of London, Zoological Journal of the Linnean Society, 2010, 159 , 673–710.     

Diversification of low dispersal crustaceans through mountain uplift: a case study of Gammarus (Amphipoda: Gammaridae) with descriptions of four novel species

Lineages with low dispersal ability are geographically restricted. We used freshwater Gammarus to test this hypothesis. Sequences of two mitochondrial (cytochrome c oxidase subunit I and 16S) and two nuclear (28S and cytosolic heat‐shock protein) genes were obtained for seven species distributed in 28 localities along the Lüliang and Taihang mountains in China. Phylogenetic analyses showed that Gammarus species were grouped into two clades, one from the Lüliang range and the other from the Taihang range. Each clade was further divided into three or four species, showing a congruent pattern with geographical vicariance. Divergence time estimation indicated that the split between the two clades coincided with the uplift of the Taihang Mountains at the boundary of Oligocene/Miocene. Most speciation events may have been driven by massive uplifting of the Lüliang and Taihang mountains from the late Miocene to early Pliocene. Additionally, four new species are described: G ammarus incoercitus sp. nov. , G ammarus benignus sp. nov. , G ammarus monticellus sp. nov. , and G ammarus pisinnus sp. nov. The new species are compared with related species in this area and a key to these species is provided. © 2014 The Linnean Society of London     

New species of Hispanomys (Rodentia,Cricetodontinae) from the Upper Miocene of Batallones (Madrid,Spain)

Material of Hispanomys (Rodentia, Cricetodontinae) is described from various localities at Batallones (MN10) (Madrid, Spain). All of it belongs to a single species, which differs from the other known species of the genus and a new taxon, Hispanomys moralesi sp. nov. , is created for it. The samples from the various localities show differences interpreted as being the result of slight age disparities amongst the different sites. Although they were previously thought to be coeval, Batallones 10 is probably older than Batallones 1, which is possibly older than Batallones 3. Hispanomys moralesi sp. nov. is characterized by several morphological features such as the lack of cingula and mesolophs, the presence of well‐developed ectolophs, four‐ or five‐rooted M1, short or absent mesolophids, and reduced and simplified M3. Hispanomys moralesi sp. nov. is a relatively derived species, the evolutionary stage of which is comparable to those of other members of the genus from the Upper Vallesian. © 2010 The Linnean Society of London, Zoological Journal of the Linnean Society, 2010, 160 , 725–747.     

Three new species of Callulina (Amphibia: Anura: Brevicipitidae) highlight local endemism and conservation plight of Africa's Eastern Arc forests

Material ascribed to the genus Callulina from north‐east Tanzania and south‐east Kenya is assessed. Three new species of Callulina are described from the North ( Callulina laphami sp. nov. ) and South ( Callulina shengena sp. nov. and Callulina stanleyi sp. nov. ) Pare Mountains in Tanzania. The species are diagnosed based on morphological, acoustic, and molecular data. A new key to the species of Callulina is provided. Based on an interpretation of the International Union for Conservation of Nature (IUCN) red list, we suggest that the three species will qualify as critically endangered, because of their small distributions and the ongoing threat to their habitat. We reveal the high local endemism of Callulina in the northern part of the Eastern Arc Mountains, with each species restricted to no more than one mountain (fragment) block. This high local endemism in Callulina is probably widespread across the Eastern Arc, raising further conservation concern for this group of amphibians. Based on new molecular phylogenetic data for Callulina, we discuss biogeographical relationships among north‐east Tanzanian mountains, and evolutionary patterns in Eastern Arc breviciptids. © 2010 The Linnean Society of London, Zoological Journal of the Linnean Society, 2010, 160 , 496–514.     

Revision and cladistic analysis of the jumping spider genus Onomastus (Araneae: Salticidae)

The jumping spider genus Onomastus Simon, 1900 is revised. Four new species: Onomastus indra sp. nov. , Onomastus kaharian sp. nov. , Onomastus pethiyagodai sp. nov. , and Onomastus rattotensis sp. nov. are described. Parsimony analysis of 26 morphological characters supported the monophyly of Onomastus. Lyssomanes is sister to Onomastus. Onomastus separates into two clades: the widespread South‐East Asia clade and the South Asia clade. The South Asia clade is restricted to the Sri Lanka–Western Ghats biodiversity hotspot. Species of the South Asia clade appear to be spot endemics, highly in danger of extinction because of habitat loss and climate change. Male palps are complex and species‐specific, suggesting rapid divergent evolution. © 2010 The Linnean Society of London, Zoological Journal of the Linnean Society, 2010, 159 , 711–745.     

A revision of West Palaearctic species of the Micropsectra atrofasciata species group (Diptera: Chironomidae)

Phylogenetic relations among selected Micropsectra (Diptera: Chironomidae) species are inferred using mitochondrial COII gene sequences and morphological characters. Both maximum parsimony and Bayesian analyses indicate a monophyletic atrofasciata species group. However, some species morphologically similar to M. atrofasciata (Kieffer) in the adult male, i.e. M. appendica sp. nov. and M. logani (Johannsen), are placed outside the atrofasciata group in the gene trees. The phylogenetic hypotheses based on parsimony analyses of the combined molecular and morphological data sets weakly support a monophyletic atrofasciata group including M. appendica sp. nov. and M. logani if third positions in the COII gene sequences are excluded. In the taxonomic part of this study, adult males, pupae and larvae of Micropsectra aristata Pinder, M. atrofasciata, M. bavarica sp. nov., M. klinki sp. nov., M. logani, M. pallidula (Meigen) and M. schrankelae sp. nov. , and adult males and pupae of M. appendica sp. nov. and M. sofiae sp. nov. as well as adult males of M. andalusiaca Marcuzzi, M. calcifontis sp. nov., M. freyi Storå, M. longicrista sp. nov., M. robusta sp. nov. , and M. zernyi Marcuzzi are described and diagnosed. Micropsectra bidentata (Goetghebuer), M. jokaquarta Sasa & Ogata, M. miki Marcuzzi, M. pallida Goetghebuer, M. subnitens Goetghebuer, M. suecica (Kieffer) and M. shouharasima Sasa are new junior synonyms of M. atrofasciata; Micropsectra groenlandica Andersen and M. utonaitertia Sasa are new junior synonyms of M. logani; Micropsectra candida Chaudhuri & Datta, M. franzi Goetghebuer and M. viridiscutellata Goetghebuer are new junior synonyms of M. pallidula. Comments on the ecology, distribution and systematics of each species are provided. Keys to all known life stages are presented, as well as an annotated list of species morphologically similar to species in the atrofasciata group and nomina dubia in the atrofasciata group. Chironomus mimulus Holmgren is transferred to the genus Micropsectra. © 2006 The Linnean Society of London, Zoological Journal of the Linnean Society, 2006, 146 , 165–225.     

Systematics and biodiversity of Ophryotrocha (Annelida,Dorvilleidae) with descriptions of six new species from deep-sea whale-fall and wood-fall habitats in the north-east Pacific

Six new Ophryotrocha species are described from five whale-falls and two wood-falls off the southern Californian coast. Phylogenetic analyses based on the nuclear gene H3 and the mitochondrial genes COI and 16S using MrBayes and maximum likelihood analyses were performed on 40 dorvilleid taxa and one outgroup. Ophryotrocha batillus sp. nov. is morphologically identical to Ophryotrocha scutellus described from a shallow water whale-fall in the North Atlantic, although the two cryptic species differ genetically. Ophryotrocha langstrumpae sp. nov. is closely related in the molecular phylogenetic analyses to these two sibling species. Ophryotrocha flabella sp. nov. is similar to Ophryotrocha globopalpata, and although there are a few morphological differences, the genetic divergence is low between the two species. Ophryotrocha nauarchus sp. nov. is sexually dimorphic, with males having appendages on the first chaetiger. Ophryotrocha magnadentata sp. nov. and Ophryotrocha longicollaris sp. nov. are sister species in our molecular analyses, and together with O. nauarchus sp. nov. and O. flabella sp. nov. they fall within a clade that includes O. globopalpata and Exallopus jumarsi described from hydrothermal vents in the Pacific Ocean and Ophryotrocha longidentata from the shallow North Atlantic. Our results highlight the remarkable unknown diversity of deep-water habitats and the role of chemosynthetic ecosystems in the evolution of deep-sea life.     

A new genus of African Acrometopini (Tettigoniidae: Phaneropterinae) based on morphology,chromosomes, acoustics,distribution, and molecular data,and the description of a new species

A new genus, Altihoratosphaga, is erected for species formerly assigned to Horatosphaga Schaum, 1853, and a new species is described. Four species are included in Altihoratosphaga: Altihoratosphaga nomima (Karsch, 1896), Altihoratosphaga montivaga ( Sjöstedt, 1909 ), Altihoratosphaga nou (Hemp, 2007) and Altihoratosphaga hanangensis sp. nov. All four species are restricted to Tanzanian localities, and, except for A. nomima, for which no ecological data are available, are confined to montane forest habitats. Data on ecology, acoustics, chromosomes, and molecular relationships are provided, as well as a key to Altihoratosphaga species. The present‐day distribution of Altihoratosphaga species suggests former migration events at times when wetter and colder climatic fluctuations favoured connections between montane forest communities, which today are isolated, enabling flightless taxa such as Altihoratosphaga and Monticolaria to spread. © 2010 The Linnean Society of London, Zoological Journal of the Linnean Society, 2010, 158 , 66–82.     

DNA sequences corroborate Soesiladeepakius as a non‐salticoid genus of jumping spiders: placement with lapsiines,phylogeny, and description of six new species (Araneae,Salticidae)

Species of the Amazonian jumping spider genus Soesiladeepakius Makhan are confirmed as non‐salticoids. Sequences of nuclear (28S, Actin) and mitochondrial (16S through NADH dehydrogenase subunit I, ‘16S‐ND1’) gene regions, analysed under parsimony and maximum likelihood, placed the genus within the lapsiines, closely related to Galianora Maddison. Additionally, six new species of this genus are herein described, namely Soesiladeepakius lyra sp. nov. , Soesiladeepakius retroversus sp. nov. , Soesiladeepakius arthrostylus sp. nov. , Soesiladeepakius gasnieri sp. nov. , Soesiladeepakius biarmatus sp. nov. , and Soesiladeepakius uncinatus sp. nov. , all from the Amazon region in Brazil. To test the monophyly of Soesiladeepakius within lapsiines, a cladistic analysis was carried out using a data matrix comprising 24 morphological characters scored for 12 taxa. The analysis resulted in two equally parsimonious trees of 29 steps. One of these trees is used to discuss the relationships among the species of Soesiladeepakius and character evolution. © 2012 The Linnean Society of London, Zoological Journal of the Linnean Society, 2012, 165 , 274–295.