Further observations on scorpion genera Hadrurus and Hoffmannihadrurus (Scorpiones, Caraboctonidae)

Multiple populations of Hadrurus pinteri from Baja California Sur, Mexico have been examined. It is demonstrated that the southern populations of this species have a larger number of accessory trichobothria (neobothriotaxy) than the northern populations, numbers exceeding the maximum currently recorded for the genus. Examination of carapace and chela coloration and its patterns show a close affinity between Hadrurus pinteri and the dark phase of Hadrurus concolorous. A new morphometric ratio of the carapace is defined that distinguishes Hadrurus from Hoffmannihadrurus, further supporting the monophyly of the latter genus.     

The Buproridae Thorell, 1859, a family of ascidicolous copepods (Copepoda: Cyclopoida)

The adult female of Buprorus loveni Thorell, 1859, a cyclopoid copepod inhabiting the branchial cavities of solitary ascidians, is redescribed in detail from new material collected in Scandinavian waters and from the southern part of the North Atlantic off Mauritania. The antennal segmentation pattern exhibited by all seven subfamilies of the family Ascidicolidae Thorell, 1860 is compared. The segmentation of the Buprorinae is different from that of all other subfamilies within the ascidicolid series. It is concluded that the antenna of Buprorus is not an appropriate model for interpreting the segmentation of other ascidicolids because the Buprorinae is the only subfamily in which the first and second endopodal segments are primitively fused and the third is distinct. The Buproridae Thorell, 1859 is here treated as a separate family within the Cyclopoida.     

Courtship and Mating in the Giant Hairy Desert Scorpion, Hadrurus arizonensis (Scorpionida, Iuridae)

Hadrurus arizonensis is a large, long-lived species of North American desert scorpion with lengthy, stereotyped courtship behaviors that lead to sperm transfer via an external spermatophore. Courtship and mating behaviors in H. arizonensis and other members of the Iuridae family have not been described. H. arizonensis has reproductive behavior similar to that of other scorpions, including the promenade a deux, but with some unique components described here for the first time. Courtship and mating behaviors of H. arizonensis are presented in a flowchart to emphasize its stereotypical nature and suitability for experimental manipulation in field and laboratory studies.     

Phylogeny of the spider genus Ixchela Huber, 2000 (Araneae: Pholcidae) based on morphological and molecular evidence (CO1 and 16S), with a hypothesized diversification in the Pleistocene

The genus Ixchela Huber is composed of 20 species distributed from north‐eastern Mexico to Central America, including the five new species described here from Mexico: I xchela azteca sp. nov. , I xchela jalisco sp. nov. , I xchela mendozai sp. nov. , I xchela purepecha sp. nov. and I xchela tlayuda sp. nov. We test the monophyly and investigate the phylogenetic relationships among species of the genus Ixchela using morphological and molecular data. Parsimony (PA) analysis of 24 taxa and 40 morphological characters with equal and implied weights supported the monophyly of Ixchela with eight morphological synapomorphies. The PA analyses with equal and implied weights, and separate Bayesian inference (BI) analyses for the CO1 gene (506 characters), concatenated gene fragments CO1 + 16S (885 characters), morphology + CO1 (546 characters) and the combined evidence data set (morphology + CO1 + 16S) (925 characters) support the monophyly of Ixchela. Our preferred topology shows two large clades; clade 1 has a natural distribution in the Mesoamerican biotic component, whereas clade 2 predominates in the Mexican Montane biotic component. The genus Ixchela diverged in the late Miocene, and the divergence between the internal clades in the genus occurred in the late Pliocene; by contrast, most of the speciation events seem to have occurred mainly during the Pleistocene, where climatic changes brought on by repeated glaciations played an important role in the diversification of the genus. © 2015 The Linnean Society of London     

Evidence of Mate Trailing in the Giant Hairy Desert Scorpion, Hadrurus arizonensis (Scorpionida, Iuridae)

During the mating season, male desert scorpions are often found wandering through dune environments, presumably in search of female conspecifics. For a male desert scorpion, finding a receptive female is a potentially difficult and hazardous task, suggesting that wandering by the male may actually be a form of guided mate searching. In this study we examined whether male giant hairy desert scorpions (Hadrurus arizonensis) were capable of trailing female conspecifics using a Y-maze choice test. In three separate experiments, a significant proportion of male scorpions preferred the arm of the maze that a reproductive female scorpion had walked down. Male scorpions did not prefer maze arms that male conspecifics had walked down, indicating that this response was sex specific. When the substrate was repeatedly exposed to a reproductive female, males took significantly longer to complete the Y-maze test, exhibited an increase in pausing behavior, and displayed precourtship behaviors and a novel lunging behavior resembling sex-specific mate seizing behavior. The results from this study suggest that male H. arizonensis can orient and respond to substrate-borne signals from female conspecifics that are likely chemical in origin.     

The genus Hormiops Fage, 1933 (Hormuridae,Scorpiones), a palaeoendemic of the South China Sea: Systematics and biogeography

The monotypic genus Hormiops Fage, 1933 is so far only known from two groups of granitic islands off the coasts of Peninsular Malaysia and Vietnam. Examination of newly collected material from both archipelagos and of the type series of Hormiops davidovi Fage, 1933 reveals previously disregarded morphological differences sufficient to assign the Malaysian specimens to a distinct species, described here as Hormiops infulcra sp. nov. An updated diagnosis of the genus, as well as a dichotomic key enabling the determination of Hormiops from its close relatives, Hormurus Thorell, 1876 and Liocheles Sundevall, 1833 are also provided. The phylogenetic position, distribution pattern, and ecology of these insular scorpions suggest that they are palaeoendemics, remnants of a previously more widely distributed lineage. A biogeographical model is proposed for the genus based on these observations and on a synthesis of palaeogeographical and palaeoenvironmental data currently available for Sundaland.     

Morphology‐based phylogenetic analysis and classification of the family Rhinocryptidae (Aves: Passeriformes)

The family Rhinocryptidae comprises an assemblage of 12 genera and 55 species confined to the Neotropical region. Here we present the first morphology‐based phylogenetic study of the Rhinocryptidae, using 90 anatomical characters (62 osteological, 28 syringeal) scored for all genera of the family and representatives of all families of the infraorder Furnariides. Parsimony analysis of this dataset recovered 7428 equally most‐parsimonious trees. The strict consensus of those trees was completely resolved at the genus level, with the topology (Liosceles (Psilorhamphus ((Eleoscytalopus + Merulaxis) (Acropternis ((Teledromas + Rhinocrypta) ((Pteroptochos + Scelorchilus) (Eugralla (Myornis + Scytalopus)))))))). The monophyly of the Rhinocryptidae as presently understood was recovered with strong support [eight synapomorphies and Bremer support (BS) = 6). Strongly supported internal arrangements included the basal position of the Amazonian genus Liosceles relative to the rest of the family (four synapomorphies, BS = 4), a clade containing Acropternis through Scytalopus (six synapomorphies, BS = 4), and other less inclusive nodes. The main points of congruence between the present morphological phylogeny and previous molecular phylogenetic work on the family were clades supported by six or more synapomorphies and Bremer values of 6–7: Eleoscytalopus + Merulaxis (eight synapomorphies, BS = 6), Scelorchilus + Pteroptochos (seven synapomorphies, BS = 7), Rhinocrypta + Teledromas (seven synapomorphies, BS = 7), and Eugralla + Myornis + Scytalopus (six synapomorphies, BS = 6). A classification derived from the morphological phylogeny is proposed, with new suprageneric taxa being named and diagnosed. © 2012 The Linnean Society of London, Zoological Journal of the Linnean Society, 2012, 166 , 377–432.     

Evidence for Eurogondwana: the roles of dispersal,extinction and vicariance in the evolution and biogeography of Indo‐Pacific Hormuridae (Scorpiones: Scorpionoidea)

Scorpions previously assigned to the genus Liocheles Sundevall, 1883, of the family Hormuridae Laurie, 1896, are widely distributed in the tropical forests of the Indo‐Pacific region. Revisionary systematics of these poorly known scorpions has revealed a tremendous diversity of species. As part of an ongoing investigation, the first analysis of Indo‐Pacific hormurid scorpion phylogeny based on morphological data scored for all currently recognized species of Hormiops Fage, 1933 , Hormurus Thorell, 1876, and Liocheles, is presented. The taxonomy of these scorpions is reassessed and their biogeography reinterpreted in the light of the phylogeny. Phylogenetic, morphological, and distributional data support the revalidation of Hormiops and Hormurus, previously synonymized with Liocheles. The phylogeny indicates that the Australasian hormurids are more closely related to the Afrotropical and Neotropical hormurids than to the Indian hormurids, as previously proposed, refuting the “out‐of‐India” origin of Asian hormurids. A recent paleogeographical hypothesis, the “Eurogondwana model”, is supported instead. According to this hypothesis, hormurid scorpions colonized Laurasia from Africa via the Apulia microplate (Europa terrane) in the Cretaceous, subsequently colonized the Australo‐Papuan archipelago in the early‐mid Cenozoic, and then went extinct in the Northern Hemisphere during the second half of the Cenozoic. These results suggest that, contrary to the traditional paradigm, dispersal and extinction may affect spatial and temporal biotic distributions as much as vicariance, even in animals with limited vagility, such as scorpions.     

A new species of Euscorpius Thorell, 1876 (Scorpiones, Euscorpiidae) from Turkey

A new species of the genus Euscorpius Thorell, 1876is described based on specimens collected from Dilek Peninsula (Davutlar, Aydın) in Turkey. It is characterized by an oligotrichous trichobothrial pattern (Pv= 7, et= 5/6, eb= 4) and small size. Euscorpius (Euscorpius) avcii sp. n. is the first named species of the subgenus Euscorpius from Turkey.     

Phylogeny of Grumichellini Morse, 1981 (Trichoptera: Leptoceridae) with the description of a new genus from southeastern Peru

Osflintia manu, new genus, new species, of long-horned caddisfly (Leptoceridae: Triplectidinae: Grumichellini) is described and illustrated from southeastern Peru. The phylogeny of Grumichellini Morse (Leptoceridae: Triplectidinae) is revisited and hypotheses of homology of some morphological characters are reconsidered. The monophyly of the tribe is corroborated and the phylogenetic relationships of its included genera are inferred to be (Triplexa (Gracilipsodes ((Grumichella, Amazonatolica) (Atanatolica, Osflintia, n. gen.)))) from adult and larval characters. Diagnostic characters of the new genus include the following: reduced tibial spur formula (2, 2, 2), loss of forewing crossvein sc-r1, hind wing discoidal cell closed, hind wing fork IV present, pair of long setae on tergum IX of the male genitalia, and pair of processes on the apex of segment X.     

Monophyletic groups within 'higher land birds'– comparison of morphological and molecular data

The relationships within the ‘higher land birds’ and putatively related taxa are analysed in a study using 89 morphological characters and DNA sequences of three nuclear, protein‐coding genes, c‐myc, RAG‐1, and myoglobin intron II. Separate analyses of the different data sets and a ‘total evidence’ analysis in which the data sets of the morphological and molecular analyses were combined are compared. All three analyses support the hitherto disputed sister group relationship between Pici (Ramphastidae, Indicatoridae and Picidae) and Galbulae (Galbulidae and Bucconidae). Previously unrecognized osteological synapomorphies of this clade are presented. All analyses further resulted in monophyly of the taxon [Aegothelidae + (Apodidae/Hemiprocnidae + Trochilidae)]. Analysis of the morphological data and of the combined data set also supported monophyly of the taxon [Strigiformes + (Falconidae + Accipitridae)]. The morphological data further support monophyly of the taxon (Upupidae + Bucerotidae). Other placements in the three analyses received either no or only weak bootstrap support.     

Phylogeny,classification and evolution of the water scavenger beetle tribe Hydrobiusini inferred from morphology and molecules (Coleoptera: Hydrophilidae: Hydrophilinae)

The water scavenger beetle tribe Hydrobiusini contains 47 species in eight genera distributed worldwide. Most species of the tribe are aquatic, although several species are known to occur in waterfalls or tree mosses. Some members of the tribe are known to communicate via underwater stridulation. While recent morphological and molecular‐based phylogenies have affirmed the monophyly of the tribe as currently circumscribed, doubts remain about the monophyly of included genera. Here we use morphological and molecular data to infer a species‐level phylogeny of the Hydrobiusini. The monophyly of the tribe is decisively supported, as is the monophyly of most genera. The genus Hydrobius was found to be polyphyletic, and as a result the genus Limnohydrobius stat. rev. is removed from synonymy with Hydrobius, yielding three new combinations: L. melaenus comb.n. , L. orientalis comb.n. , and L. tumbius comb.n. Recent changes to the species‐level taxonomy of Hydrobius are reviewed. The morphology of the stridulatory apparatus has undergone a single remarkable transformation within the lineage, from a simple, unmodified pars stridens to one that is highly organized and complex. We present an updated key to genera, revised generic diagnoses and a list of the known distributions for all species within the tribe.     

Wide phylogenetic distribution of Scorpine and long-chain beta-KTx-like peptides in scorpion venoms: identification of "orphan" components

Scorpine and toxins specific for potassium channels of the family beta (beta-Ktx) are two types of structurally related scorpion venom components, characterized by an unusually long extended N-terminal segment, followed by a Cys-rich domain with some resemblance to other scorpion toxins. In this communication, we report evidence supporting the ubiquitous presence of Scorpine and beta-KTx-like polypeptides and their precursors in scorpions of the genus Tityus of the family Buthidae, but also included is the first example of such peptides in scorpions from the family Iuridae. Seven new beta-KTxs or Scorpine-like peptides and precursors are reported: five from the genus Tityus (T. costatus, T. discrepans and T. trivittatus) and two from Hadrurus gertschi. The cDNA precursors for all of these peptides were obtained by molecular cloning and their presence in the venoms were confirmed for various peptides. Analysis of the sequences revealed the existence of at least three distinct groups: (1) beta-KTx-like peptides from buthids; (2) Scorpine-like peptides from scorpionid and iurid scorpions; (3) heterogeneous peptides similar to BmTXKbeta of buthids and iurids. The biological function for most of these peptides is not well known; that is why they are here considered "orphan" peptides.     

A preliminary cladistic analysis of Nipponoserica,with implications for phylogenetic relationships among sericine chafers (Coleoptera: Scarabaeidae: Sericini)

A cladistic analysis of the species of Nipponoserica Nomura, 1972, was executed using 36 morphological characters of adults. From the generated phylogenetic tree, the monophyly of the genus is supported, when Pseudosericania Kobayashi, 1980 is included. Consequently, Pseudosericania Kobayashi, 1980 is considered a junior subjective synonym of Nipponoserica Nomura, 1972. Interestingly, the analysis indicates the monophyly of the East Asian Nipponoserica and the North American Stilbolemma. A new genus, Xenoserica gen. nov., is established for the basal monophyletic clade found for the Himalayan species so far assigned to Nipponoserica including the first brachypterous species of Sericini from the Himalaya. Two new species are described: Xenoserica brachyptera sp. nov. from Nepal, and Nipponoserica dahongshanica sp. nov. from Hubei (China).     

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.     

Lyneborgimyia magnifica gen. et sp.n. (Diptera: Syrphidae) from Tanzania,with a phylogenetic analysis of the Eumerini using new morphological characters

Lyneborgimyia magnifica gen. et sp.n. is described and placed with Merodon Meigen and Platynochaetus Wiedemann in the Merodon genus group within the tribe Eumerini. New morphological characters and terms are introduced, and precise definitions proposed for characters applied inconsistently in the literature. The species possesses several unique autapomorphies, including an additional cross‐vein between R₄₊₅ and M₁, and a sclerotized connection between sternite 1 and the metepimeron, which are unknown otherwise within Diptera. A parsimony‐based phylogenetic analysis including 73 characters from the adult morphology provides strong support for the monophyly of the Eumerini (Eumerus Meigen, Azpeytia Walker, Lyneborgimyia, Merodon, Platynochaetus) and the Merodon genus group, whereas no support is found for the possible monophyly of the genus Eumerus. A sister‐group relationship is indicated between the Afrotropical Lyneborgimyia and the Palaearctic Platynochaetus.     

Phylogeny and systematics of the subfamily Bryocorinae based on morphology with emphasis on the tribe Dicyphini sensu Schuh,

The classification of the hyperdiverse true bug family Miridae is far from settled, and is particularly contentious for the cosmopolitan subfamily Bryocorinae. The morphological diversity within the subfamily is pronounced, and a lack of explicit character formulation hampers stability in the classification. Molecular partitions are few and only a handful of taxa have been sequenced. In this study the phylogeny of the subfamily Bryocorinae has been analysed based on morphological data alone, with an emphasis on evaluating the tribe Dicyphina sensu Schuh, 1976, within which distinct groups of taxa exist. A broad sample of taxa was examined from each of the bryocorine tribes. A broad range of outgroup taxa from most of the other mirid subfamilies was also examined to test for bryocorine monophyly, ingroup relationships and to determine character polarity. In total a matrix comprising 44 ingroup, 15 outgroup taxa and 111 morphological characters was constructed. The phylogenetic analysis resulted in a monophyletic subfamily Bryocorinae sensu Schuh (1976, 1995), except for the genus Palaucoris, which is nested within Cylapinae. The tribe Dicyphini sensu Schuh (1976, 1995) has been rejected. The subtribe Odoniellina is synonymized with the subtribe Monaloniina and the subtribes Dicyphina, Monaloniina and Eccritotarsina are now elevated to tribal level, with the Dicyphini now restricted in composition and definition. The genus Felisacus is highly autapomorphic and a new tribe – the Felisacini – is erected for the included taxa. This phylogeny of the tribes of the Bryocorinae comprises the following sister‐group relationships: Dicyphini ((Bryocorini + Eccritotarsini)(Felisicini + Monaloniini)).     

Structural features of the mitogenome of the leafhopper genus Cladolidia (Hemiptera: Cicadellidae: Coelidiinae) and phylogenetic implications in Cicadellidae

The first two complete mitogenomes of the leafhopper genus Cladolidia (C. biungulata and C. robusta) were sequenced and annotated to further explore the phylogeny of Cladolidia. Both the newly sequenced mitogenomes have a typical circular structure, with lengths of 15,247 and 15,376 bp and A + T contents of 78.2% and 78%, respectively. We identified a highly conserved genome organization in the two Cladolidia spp. through comparative analysis that included the following assessments: genome content, gene order, nucleotide composition, codon usage, amino acid composition, and tRNA secondary structure. Moreover, we detected the base heterogeneity of Cicadellidae mitogenomic data and constructed phylogenetic trees using the nucleotide alignments of 12 subfamilies of 58 leafhopper species. We noted a weak heterogeneity in the base composition among the Cicadellidae mitogenomes. Phylogenetic analyses showed that the monophyly of each subfamily was generally well supported in the family Cicadellidae; the main topology was as follows: (Deltocephalinae + (Treehoppers + ((Megophthalminae + (Macropsinae + (Hylicinae + (Coelidiinae +Iassinae)) + (Idiocerinae + (Cicadellinae + (Typhlocybinae + (Mileewinae + (Evacanthinae +Ledrinae)))))))))). Within Coelidiinae, phylogenetic analyses revealed that C. biungulata and C. robusta belong to Coelidiinae and the monophyly of Cladolidia is well supported. In addition, on the basis of complete mitogenome phylogenetic analysis and the comparison of morphological characteristics, we further confirm the genus Olidiana as a paraphyletic group, suggesting that the genus may need taxonomic revisions.     

Phylogenetic analysis of Trechitae (Coleoptera: Carabidae) based on larval morphology, with a description of first-instar Phrypeus and a key to genera

Abstract. Sixty-nine characters of larval structure of twenty-eight genera of the supertribe Trechitae (Coleoptera: Carabidae) were analysed phylogenetically. The monophyly of Trechitae is strongly supported with five unique synapomorphies. The monophyly of Zolini + Bembidiini + Pogonini is supported with two synapomorphies. We propose that the tribe Trechini is a sister group to them and its monophyly is supported with two unique synapomorphies. The inferred branching pattern of Trechini genera is (Perileptus + Thalassophilus) + (Amblystogenium + (Trechimorphus + (Trechus + Epaphius + Aepopsis + Trechisibus))); Perileptus is a member of Trechodina rather than Trechina. The monophyly of Zolini is not supported. The monophyly of Pogonini is supported with two unique synapomorphies; its sister group relationships remain obscure; the branching pattern of pogonine genera is (((Pogonus + Pogonistes) + Cardiaderus) + Thalassotrechus). No evidence for monophyly of the tribe Bembidiini (s. lato; including subtribes Bembidiina, Tachyina, Xystosomina, and Anillina) was found. The relationships of Phrypeus are obscure; no evidence could be found linking it with Bembidiina. Without Phrypeus, Bembidiina might be a monophylum with a single synapomorphy. Sinechostictus branches basal of (Bembidion + Asaphidion) and therefore should be treated as a separate genus. Tachyina and Xystosomina form a monophylum based on two unique synapomorphies; a close relationship with a monophyletic Anillina is suggested. Reduction of the number of claws from two to one in Trechitae has taken place twice: within Trechina (Trechus, Epaphius, Aepopsis and Trechisibus) and in (Zolini + Bembidiini + Pogonini). The previously unknown larvae of the isolated genus Phrypeus are described and illustrated. A key to all twenty-eight analysed Trechitae genera based on characters of larvae and a list of larval autapomorphies for each genus are provided.