Intraspecific or interspecific variation: delimitation of species boundaries within the genus Gammarus (Crustacea,Amphipoda, Gammaridae), with description of four new species

The delineation of Gammarus species is controversial because of extensive intraspecific morphological variation. The current study examined DNA sequences from the mitochondrial cytochrome c oxidase subunit I and the nuclear 28S genes as well as morphological and ecological data to determine the species boundaries of Gammarus species from China. The results of molecular analyses showed that Gammarus sp1, G. sp2, G. sp3, and G. sp4 are monophyletic and deeply divergent from sister groups. Detailed morphological and ecological comparisons with closely related species were consistent with molecular analyses. Gammarus sp1, G. sp2, G. sp3, and G. sp4 were described as four new species: Gammarus illustris sp. nov. , Gammarus clarus sp. nov. , Gammarus hypolithicus sp. nov. , and Gammarus parvioculus sp. nov. We recommend that molecular detected species should be formally named and described for future research. © 2010 The Linnean Society of London, Zoological Journal of the Linnean Society, 2010, 160 , 215–253.     

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.     

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.     

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.     

The first fossil spider cricket (Orthoptera: Gryllidae: Phalangopsinae): 20 million years of troglobiomorphosis or exaptation in the dark?

A new spider cricket (Orthoptera: Gryllidae: Phalangopsinae) is described from an adult female preserved in Early Miocene (Burdigalian) amber from the Dominican Republic. Araneagryllus dylani gen. et sp. nov. represents the first fossil record of Phalangopsinae, and is assigned to the tribe Luzarini, subtribe Amphiacustina stat. nov. A cladistic analysis of Amphiacustina places Araneagryllus gen. nov. within a clade comprising Arachnopsita, Leptopedetes, Longuripes, Mayagryllus, Nemoricantor, and Prolonguripes. This clade is the sister group to a clade comprising Amphiacusta, Cantrallia, and Noctivox. The results of this analysis suggest that: (1) the common ancestor of all Amphiacustina was epigean, and was likely to have been cavicolous and/or straminicolous; (2) strict troglobitism evolved twice within Amphiacustina, once in the lineage leading to Noctivox and again in the clade comprising Mayagryllus, Arachnopsita, Longuripes, and Prolonguripes; and (3) Prolonguripes is secondarily epigean, having reverted to life above ground. The occurrence of Araneagryllus gen. nov. in amber indicates that it was not troglobitic, but was instead more likely to have been straminicolous, living on the ground and foraging amongst leaf litter. Araneagryllus gen. nov. possesses a number of characters that are usually considered to be adaptive to a troglobitic life history, suggesting that many so‐called troglobiomorphies are not adaptations to life in caves, but are instead likely to have been exaptive. © 2010 The Linnean Society of London, Zoological Journal of the Linnean Society, 2010, 158 , 56–65.     

First fossil Molinaranea Mello‐Leitão, 1940 (Araneae: Araneidae), from middle Miocene Dominican amber,with a phylogenetic and palaeobiogeographical analysis of the genus

The first fossil Molinaranea is described, from middle Miocene Dominican amber. This record extends the known range of the genus back 16 million years; it also extends the geographical range of the genus through time, with extant species known only from Chile, Argentina, the Falkland Islands, and Juan Fernandez Island. A parsimony‐based phylogenetic analysis was performed, which indicates that the fossil species, Molinaranea mitnickii sp. nov. , is nested with Molinaranea magellanica Walckenaer, 1847 and Molinaranea clymene Nicolet, 1849 . A modified Brooks parsimony analysis was conducted in order to examine the biogeography and origins of the fossil species in the Dominican Republic; the analysis suggests that M. mitnickii sp. nov. arrived in Hispaniola from South America as a result of a chance dispersal event. © 2010 The Linnean Society of London, Zoological Journal of the Linnean Society, 2010, 158 , 711–725.     

Descriptions and phylogenetic relationships of a new genus and two new species of Oligo‐Miocene cormorants (Aves: Phalacrocoracidae) from Australia

Tertiary cormorant fossils (Aves: Phalacrocoracidae) from Late Oligocene deposits in Australia are described. They derive from the Late Oligocene – Early Miocene (26–24 Mya) Etadunna and Namba Formations in the Lake Eyre and Lake Frome Basins, South Australia, respectively. A new genus, Nambashag gen. nov. , with two new species ( Nambashag billerooensis sp. nov. , 30 specimens; Nambashag microglaucus sp. nov. , 14 specimens), has been established. Phylogenetic analyses based on 113 morphological and two integumentary characters indicated that Nambashag is the sister taxon to the Early Miocene Nectornis miocaenus of Europe and all extant phalacrocoracids. As Nambashag, Nectornis, and extant phalacrocoracids constitute a strongly supported clade sister to Anhinga species, the fossil taxa have been referred to Phalacrocoracidae. Sulids and Fregata were successive sister taxa to the Phalacrocoracoidea, i.e. phalacrocoracids + Anhinga. As phalacrocoracids lived in both Europe and Australia during the Late Oligocene and no older phalacrocoracid taxa are known, the biogeographical origin of cormorants remains unanswered. The phylogenetic relationships of extant taxa were not wholly resolved, but contrary to previous morphological analyses, considerable concordance was found with relationships recovered by recent molecular analyses. Microcarbo is sister to all other extant phalacrocoracids, and all Leucocarbo species form a well‐supported clade. © 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 163 , 277–314.     

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.     

Phylogenetic analysis and revision of the linyphiid spider genus Solenysa (Araneae: Linyphiidae: Erigoninae)

In this paper we carry out a taxonomic revision and phylogenetic analysis of the linyphiid spider genus Solenysa Simon, 1894. A total of 12 species is treated here, including five new species collected from China and Japan: Solenysa akihisai Tu sp. nov., Solenysa lanyuensis Tu sp. nov., Solenysa retractilis Tu sp. nov., Solenysa tianmushana Tu sp. nov. , and Solenysa yangmingshana Tu sp. nov. Solenysa circularis Gao, Zhu & Sha, 1993 is a junior synonym of Solenysa protrudens Gao, Zhu & Sha, 1993. We have assembled two different character matrices to reconstruct the phylogenetic relationships of Solenysa. In the first matrix (Matrix 1), five representative species of Solenysa were added to the morphological dataset of Miller & Hormiga to test the monophyly of the genus and its placement within Linyphiidae. The genitalic structures and somatic morphology of Solenysa were studied by means of scanning electron microscopy for the first time. To infer the species‐level phylogenetic relationships of Solenysa we produced a second matrix (Matrix 2) that includes all 12 Solenysa species and six outgroup species chosen from the results of the analysis of the first matrix. The two most parsimonious trees from the analysis of Matrix 1 support the monophyly of Solenysa and its placement within the ‘Distal Erigonines’ clade. The single most parsimonious tree resulting from the analysis of the second matrix suggests that the Solenysa clade includes four monophyletic groups, each group represented by a distinct genitalic pattern. The morphology of Solenysa, both somatic and genitalic, is highly autapomorphic. © 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 161 , 484–530.     

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.     

Nothofagus subgenus Brassospora (Nothofagaceae) leaf fossils from New Zealand: a link to Australia and New Guinea?

N othofagus palustris sp. nov. is the first record of well‐preserved leaves of Nothofagus subgenus Brassospora in New Zealand, and is described from an Oligo–Miocene leaf bed in the Gore Lignite Measures of the South Island. Nothofagus palustris is represented by relatively small and variably toothed leaves with cuticular remains that possess all the characteristic features of the subgenus, including the presence of variably sized stomata that are randomly arranged within areoles, hydathodes along the major veins and ‘bulging cells’ within the areoles on the adaxial side. Phylogenetic assessment shows that the leaves are similar to those of Australian Oligocene and Miocene species and may belong to the same clade of Brassospora. Most notably, these species share the derived feature of abundant leaf wax, a feature that is now only well developed in two New Guinean species. This and other evidence allows the possibility that the ancestor of N. palustris reached New Zealand from Australia. However, it is improbable that N. palustris or a similar species was the common ancestor of the clade of Brassospora that is now confined to New Caledonia. Ecologically, N. palustris is unusual among extant and previously described macrofossil species of Brassospora in being found in a relatively open, swampy habitat. © 2014 The Linnean Society of London, Botanical Journal of the Linnean Society, 2014, 174 , 503–515.     

New representatives of the genus Ancorabolina George, 2006 (Copepoda,Harpacticoida, Ancorabolidae) including remarks on ancorabolid phylogeny

Four new species of Ancorabolina George, 2006 (Copepoda: Harpacticoida: Ancorabolidae) are described: Ancorabolina belgicae sp. nov. (from the Porcupine Seabight, north‐east Atlantic Ocean), Ancorabolina anaximenesi sp. nov. and Ancorabolina galeata sp. nov. (both from Anaximenes Mountain, eastern Mediterranean Sea), and Ancorabolina divasecunda sp. nov. (from Cape and Guinea Basins, south‐east Atlantic Ocean). Furthermore, the generic diagnosis for Ancorabolina is provided and the position of the taxon within Ancorabolidae is discussed. The characters originally employed to allocate Ancorabolina to the subfamily Ancorabolinae Sars, 1909, are critically evaluated and two true autapomorphies for a monophyletic Ancorabolinae (including Ancorabolina) are recognized. Two autapomorphies supporting Ancorabolina are proposed: (1) cephalothorax posteriorly with pair of ventrolateral cuticular processes turned backwards and (2) loss of the third setal element subapically on the antennary endopod. The urgent need for a phylogenetic re‐evaluation of the paraphyletic subfamily Laophontodinae Lang, 1944 is stressed and evidenced by three characters widespread in this subfamily but also present in Ancorabolina. This may be an indication of a closer relationship of Ancorabolina with certain members of Laophontodinae. © 2010 The Linnean Society of London, Zoological Journal of the Linnean Society, 2010, 158 , 16–55.     

A new Oligocene Ctenodactylinae (Rodentia: Mammalia) from Ulantatal (nei Mongol): new insight on the phylogenetic origins of the modern Ctenodactylidae

The four living genera of Ctenodactylidae (Rodentia) are the only survivors of a flourishing Tertiary group. In this paper, we describe a new Oligocene species from Ulantatal (Chinese Mongolia) that highlights the origins of crown ctenodactylids. This species, Helanshania deserta gen. et sp. nov. , is lophodont and displays semi‐hypsodont teeth, a dental pattern that is somewhat transitional between that of primitive Oligocene ctenodactylids and the later hypsodont genera. We perform here a cladistic assessment of the dental evidence for species produced by the successive radiations of the group. In order to get new data, to decipher homologies for the dental pattern of modern ctenodactylids, and to specify their dental replacement, we describe additional dental material of Ctenodactylus, Massoutiera, and Felovia. The phylogenetic analysis (using PAUP) considered 45 characters (mainly dental) and 31 species. The performed heuristic searches yielded 596 equally most parsimonious trees. Protataromys and Karakoromys are stem ctenodactylids and appear as the earliest offshoots of the Ctenodactylidae clade, which represents a well‐supported family rank. Within this family, the Tataromyinae appear paraphyletic, whereas the Ctenodactylinae sensu lato are a clade including the new taxon Helanshania. As such, a revision of the Tataromyinae is envisaged and a new subfamily is erected (Yindirtemyinae). Amongst the Ctenodactylinae, a tribe Ctenodactylini encompassing the crown ctenodactylines is proposed. © 2010 The Linnean Society of London, Zoological Journal of the Linnean Society, 2010, 160 , 531–550.     

The lace web spiders (Araneae,Phyxelididae) of Madagascar: phylogeny,biogeography and taxonomy

We briefly review the potential history of Madagascar as either a Darwinian or a Wallacean island, summarize the phylogenetic evidence regarding the biogeography of Madagascar spiders, examine the dispersal history of the Madagascar Phyxelididae, and monograph the family in Madagascar. Molecular phylogenetic analyses for 32 Malagasy phyxelidid exemplars, nine confamilial outgroup taxa, and seven other more distant outgroups are performed for three nuclear markers and one mitochondrial genetic marker (28S, 18S, H3 and COI) utilizing Bayesian, maximum‐likelihood and parsimony methods. These analyses suggest that there are 14 species of Phyxelididae that may be recognized from Madagascar, that these may be divided into three genera, and that the Malagasy phyxelidids form a monophyletic group, probably resulting from a single invasion of the island by an ancestor from Africa. Two new genera, ten new species, and two new combinations are proposed: Manampoka atsimo gen. nov., sp. nov. ; Rahavavy gen. nov. , including R. ida sp. nov. and R. fanivelona (Griswold, 1990) comb. nov. and R. malagasyana (Griswold, 1990) comb. nov. ; and Ambohima andrefana sp. nov. , A. antsinanana sp. nov. , A. avaratra sp. nov. , A. maizina sp. nov. , A. ranohira sp. nov. , A. vato sp. nov. , A. zandry sp. nov. and A. zoky sp. nov. © 2012 The Linnean Society of London, Zoological Journal of the Linnean Society, 2012, 164 , 728–810.     

Macrocyprididae (Ostracoda) from the Southern Ocean: taxonomic revision,macroecological patterns,and biogeographical implications

Animal communities inhabiting shallow marine benthic environments around Antarctica are quite well known, however, only a few studies deal with the deep‐water faunas in the Southern Ocean. In the present study 19 deep‐water samples and 14 shallow and mid‐water samples containing macrocypridids were studied. The first important finding of the present study is the fivefold increase in the diversity of Macrocyprididae in the Southern Ocean: the species count increased from six to 30. Additionally, statistical analyses showed that depth is the main factor influencing macrocypridid assemblages and that geography does not play an important role. Two faunal bathymetric boundaries were observed, one around 1250 m and the other around 2500 m depth. Furthermore, species previously believed to be widely distributed (i.e. Macroscapha opaca, Macroscapha turbida, and Macroscapha inaequata) are shown to be groups of sometimes conspicuously distinct species. Over 30 Macrocyprididae species were identified from over 1200 specimens studied. After the study of the soft part morphology and chaetotaxy of Mh. opaca under the electron microscope, the following has been observed: (1) The exopodite of antenna II is clearly inserted on the distal margin of podomere II; (2) several setae previously considered aesthetascs (i.e. the long, proximal setae of podomere IV of the antenna II; the two sexually dimorphic setae of the podomere IV of antenna II; the modified setae of the podomeres II and III of the male appendage V) lack any pore, and should therefore not be chemoreceptors. Sixteen new species are described ( Macromckenziea giambonini sp. nov. , Macropyxis alanlordi sp. nov. , Macropyxis cronini sp. nov. , Macropyxis ghartmanni sp. nov. , Macropyxis hornei sp. nov. , Macropyxis jeans sp. nov. , Macropyxis parajeans sp. nov. , Macrosarisa andeep sp. nov. , Macrosarisa fahrbachi sp. nov. , Macroscapha cactus sp. nov. , Macroscapha falcis sp. nov. , Macroscapha rehmi sp. nov. , Macroscapha scotia sp. nov. , Macroscapha solecavai sp. nov. , Macroscapha subhemispherica sp. nov. , Macroscapha walterae sp. nov. ). Five previously described species are recorded: Macromckenziea glacierae Maddocks 1990 , Macroscapha inaequalis ( G. W. Müller, 1908 ), Mh. inaequata Maddocks 1990 , Mh. opaca Maddocks, 1990 species complex, Mh. turbida ( G. W. Müller, 1908 ). At least nine species are left in open nomenclature. Moreover, Yemanja gen. nov. is described from shallow tropical waters of the Western Atlantic; Macrosarisa procera ( Jellinek & Swanson, 2003 ) comb. nov. is proposed; and Macroscapha tensa ( Müller, 1908 ) is herein considered a nomen dubium. © 2010 The Linnean Society of London, Zoological Journal of the Linnean Society, 2010, 159 , 567–672.     

A new arthropod from the Early Cambrian of North Greenland,with a ‘great appendage’‐like antennula

Kiisortoqia soperi gen. et sp. nov. is an arthropod species from the Early Cambrian Sirius Passet Lagerstätte of North Greenland. A head, incorporating four appendiferous segments and biramous limbs, with an anteroposteriorly compressed basipod with a spine bearing median edge, support the euarthropod affinities of K. soperi gen. et sp. nov. Similarities with ‘short great appendage’ arthropods, or megacheirans, like the nine‐segmented endopod, and the flap‐ or paddle‐like exopod, may be symplesiomorphies. The antennula, however, resembles in composition and size the anteroventral raptorial appendage of anomalocaridids. Thus, the morphology of K. soperi gen. et sp. nov. provides additional support for the homologization of the anomalocaridid ‘great appendage’ with the appendage of the antennular or deutocerebral segment of extant Euarthropoda. © 2010 The Linnean Society of London, Zoological Journal of the Linnean Society, 2010, 158 , 477–500.     

Systematic revision of the genus Everettia Godwin‐Austen, 1891 (Mollusca: Gastropoda: Dyakiidae) in Sabah,northern Borneo

The species of the snail genus Everettia in the Malaysian state of Sabah are superficially similar and difficult to distinguish by their shells. This paper presents new data on the taxonomy and distribution of Everettia in Sabah that have accumulated since the revision by Godwin‐Austen in 1891. By using morphological and molecular phylogenetic approaches, we reveal at least seventeen species of Everettia in Sabah, of which eleven are new to science, namely: Everettia layanglayang sp. nov. , Everettia lapidini sp. nov. , Everettia paulbasintali sp. nov. , Everettia occidentalis sp. nov. , Everettia jasilini sp. nov. , Everettia safriei sp. nov. , Everettia interior sp. nov. , Everettia jucundior sp. nov. , Everettia planispira sp. nov. , Everettia monticola sp. nov. , and Everettia dominiki sp. nov. , and one new subspecies, namely, Everettia corrugata williamsi ssp. nov. Phylogenetic analysis of mitochondrial COI and 16S, and nuclear ITS‐1 sequences demonstrates the monophyly of most of the morphologically well‐defined species. Our results show that certain aspects of classical morphology‐based taxonomy for Everettia species, especially with regard to the unique combination of shell surface sculptures, animal head colour, and mantle pigmentation, are solid. A dichotomous key to the Sabah species and subspecies of Everettia is provided. © 2009 The Linnean Society of London, Zoological Journal of the Linnean Society, 2009, 157 , 515–550.     

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.