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Ochoa J.A.,Scorpion Systematics Research Group | Ochoa J.A.,University of Sao Paulo | Ojanguren Affilastro A.A.,Museo Argentino de Ciencias Naturales Bernardino Rivadavia | Mattoni C.I.,National University of Crdoba | Prendini L.,Scorpion Systematics Research Group
Bulletin of the American Museum of Natural History | Year: 2011

The systematics of the Andean scorpion genus, Orobothriurus Maury, 1976 (Bothriuridae Simon, 1880), is revised. New locality records, obtained during recent field expeditions, distribution maps, and a key to identification of the 15 known species, are provided. Six new species are described: Orobothriurus calchaqui, n. sp., from northwestern Argentina; Orobothriurus compagnuccii, n. sp., from the central Andes of Argentina; Orobothriurus huascaran, n. sp., from central Peru; Orobothriurus quewerukana, n. sp., from southern Peru and northern Chile; Orobothriurus ramirezi, n. sp., from central Chile; and Orobothriurus tamarugal, n. sp., from northern Chile. The known distribution of Orobothriurus and the altitude record for scorpions are discussed. The world's altitude record for a scorpion, previously reported as 5550 m, is demonstrated to be 4910 m. © 2011 American Museum of Natural History.

Mattoni C.I.,National University of Cordoba | Ochoa J.A.,University of Sao Paulo | Ojanguren Affilastro A.A.,Museo Argentino de Ciencias Naturales Bernardino Rivadavia | Prendini L.,Scorpion Systematics Research Group
Zoologica Scripta | Year: 2012

The genus Orobothriurus Maury, 1976 (Bothriuridae Simon, 1880) displays an Andean pattern of distribution, most of its species occurring at high altitudes (over 2000-2500m to a maximum altitude record of 4910m) from central Peru to Argentina. The recent discovery of several new species and the uncertain phylogenetic position of Orobothriurus lourencoi Ojanguren Affilastro, 2003, required a reanalysis of Orobothriurus phylogeny. Thirty bothriurid taxa, including all species of Orobothriurus and Pachakutej Ochoa, 2004, were scored for 65 morphological characters and analysed with parsimony under equal and implied weighting. The resulting topology justifies the establishment of a new genus, RumikiruOjanguren Affilastro , in press, for O. lourencoi and a closely related, new species, Rumikiru atacamaOjanguren Affilastro , in press. It also offers new insights about the phylogeny and biogeography of Orobothriurus and related genera. Characters from the male genitalia (i.e. hemispermatophore), comprising approximately 26% of the morphological matrix, were found to be less homoplastic than those from somatic morphology, contradicting suggestions that genitalia are uninformative or potentially misleading in phylogenetic studies. © 2012 The Authors. Zoologica Scripta © 2012 The Norwegian Academy of Science and Letters.

Loria S.F.,American Museum of Natural History | Loria S.F.,Scorpion Systematics Research Group | Prendini L.,Scorpion Systematics Research Group
PLoS ONE | Year: 2014

Scorpions possess two types of visual organs, the median and lateral eyes. Both eyes consist of simple ocelli with biconvex lenses that differ in structure and function. There is little variation in the number of median ocelli across the order. Except for a few troglomorphic species in which the median ocelli are absent, all scorpions possess a single pair. In contrast, the number of pairs of lateral ocelli varies from zero to five across Scorpiones and may vary within species. No attempt has been made to homologize lateral ocelli across the order, and their utility in scorpion systematics has been questioned, due to the variation in number. A recent study examined the number of lateral ocelli among various Asian Buthidae C.L. Koch, 1837 and proposed a "five-eye model" for the family. This model has not been examined more broadly within Buthidae, however, nor compared with the patterns of variation observed among other scorpion families. An eyespot, referred to as an accessory lateral eye, situated ventral or posteroventral to the lateral ocelli, has also been reported in some scorpions. Analysis of its structure suggests it serves a nonvisual function. We present the first comparative study of variation in the lateral ocelli across the order Scorpiones, based on examination of a broad range of exemplar species, representing all families, 160 genera (78%), 196 species (9%), and up to 12 individuals per species. We propose a six-ocellus model for Recent scorpions with four accessory ocelli observed in various taxa, homologize the individual ocelli, and correct erroneous counts in the recent literature. We also investigate the presence of the eyespot across scorpions and discover that it is more widespread than previously recognized. Future work should investigate the genetic and developmental mechanisms underlying the formation of the lateral ocelli to test the hypotheses proposed here. Copyright: © 2014 Loria, Prendini.

Bird T.L.,National Museum of Namibia | Bird T.L.,Colorado State University | Wharton R.A.,Texas A&M University | Prendini L.,Scorpion Systematics Research Group
Bulletin of the American Museum of Natural History | Year: 2015

Arachnids of the order Solifugae (solifuges, false spiders, sun spiders, camel spiders, Walzenspinne, wind spiders) possess the largest jaws for body size among the Chelicerata. The chelicerae provide the most important character systems for solifuge systematics, including dentition and the male cheliceral flagellum, both used extensively for species delimitation and diagnosis. However, the terminology used for cheliceral characters is not standardized and often contradictory, in part because it fails to represent homologous structures among taxa. Misinterpretation of character homology may introduce errors in phylogenetic analyses concerning relationships within Solifugae and among the orders of Chelicerata. This contribution presents the first comprehensive analysis of cheliceral morphology across the order Solifugae, the aims of which were to provide a broad survey of cheliceral characters for solifuge systematics, to identify and reinterpret structures based on primary homology, to revise the terminology to be consistent with homology hypotheses, and to provide a guide to terminological synonyms and character interpretations in the literature. Chelicerae were studied in 188 exemplar species (17% of the total), representing all 12 solifuge families, 17 of the 19 subfamilies, 64 genera (46% of the total), and the full range of variation in cheliceral morphology across the order. In total, 157 species representing 49 genera and 17 subfamilies are illustrated. Hypotheses of character transformation, particularly concerning the male flagellum, and a standardized terminology, are presented. The functional morphology of the chelicerae is discussed and the role of sexually dimorphic modifications to the male chelicerae in mating behavior emphasized. The revised terminology, based on hypotheses of primary homology, will facilitate solifuge revisionary systematics and provide a stronger basis for reconstructing phylogenetic relationships within the order Solifugae and testing the phylogenetic position of the order within Chelicerata. © 2015 American Museum of Natural History.

Gonzalez-Santillan E.,City University of New York | Gonzalez-Santillan E.,Scorpion Systematics Research Group | Gonzalez-Santillan E.,National Polytechnic Institute of Mexico | Gonzalez-Santillan E.,National Autonomous University of Mexico | Prendini L.,Scorpion Systematics Research Group
Cladistics | Year: 2015

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. © The Willi Hennig Society 2014.

Gonzalez-Santillan E.,City University of New York | Gonzalez-Santillan E.,Scorpion Systematics Research Group | Gonzalez-Santillan E.,CINVESTAV | Gonzalez-Santillan E.,National University of Costa Rica | Prendini L.,Scorpion Systematics Research Group
Bulletin of the American Museum of Natural History | Year: 2015

The Mexican vaejovid scorpion genus Konentontli González-Santillán and Prendini, 2013, was created to accommodate five species united, among other characters, by a subaculear tubercle on the telson. Species of Konetontli are among the smallest vaejovid scorpions. Their very small size, cryptic coloration, and apparently seasonal surface activity may explain their rarity in collections and it is likely that more undescribed species await discovery. In the present contribution, we describe four new species (Konetontli ignes, sp. nov.; Konetontli ilitchi, sp. nov.; Konetontli juxtlahuaca, sp. nov.; Konetontli migrus, sp. nov.) and revalidate Konetontli zihuatanejensis (Baldazo-Monsivaiz, 2003), comb. nov., previously synonymized with Konetontli acapulco (Armas and Martín-Friás, 2001), raising to 10 the number of species in the genus; redescribe previously described species, including the first description of the female of Konetontli nayarit (Armas and Martín-Friás, 2001); and present new records, comprehensive distribution maps, and a key to the identification of the species. © Copyright © American Museum of Natural History 2015.

Klussmann-Fricke B.-J.,University of Rostock | Prendini L.,Scorpion Systematics Research Group | Wirkner C.S.,University of Rostock
Arthropod Structure and Development | Year: 2012

Phylogenetically informative characters from the internal anatomy of scorpions were first reported more than 150 years ago, but the subject received little attention after the mid-1920s. Recent investigations, using traditional dissection, illustration and histological sectioning, microscopy, and innovative new methods for investigating complex soft tissue anatomy identified a new wealth of variation. Additionally, these investigations confirmed the phylogenetic significance of previously identified structures. Building on earlier work we present a more detailed description of the hemolymph vascular system (HVS) in scorpions, based on comparison of the vascular structures of the heart and the branching pattern of the prosomal arteries among 45 exemplar species representing most of the major scorpion lineages. Using corrosion-casting, MicroCT in combination with computer-aided 3D-reconstruction, and scanning electron microscopy, we conceptualize a series of phylogenetically informative characters for the anterior aorta system and characters of the heart and associated structures (e.g. arrangement of the ostia) of scorpions. Furthermore, we optimize the possible evolution of these new characters on a previous hypothesis of scorpion phylogeny, and discuss alternative character state transformations, their evolutionary consequences, and possible underlying evolutionary mechanisms acting on the HVS. © 2012 Elsevier Ltd.

Prendini L.,Scorpion Systematics Research Group
Bulletin of the American Museum of Natural History | Year: 2016

The scorpion fauna of East Africa, encompassing Burundi, Kenya, Tanzania, Rwanda, and Uganda, is more diverse than those of West and Central Africa, but a systematic survey has never been conducted and the distributions of its species remain poorly understood. A recent opportunity to examine two extensive collections of East African, and predominantly Kenyan, scorpions and new material acquired by the author permitted a reassessment of the fauna of the region. The present contribution, the first of several emanating from this research, comprises two parts. The first part presents a redefinition and revision of the scorpionid genus Pandinoides Fet, 1997, with a redescription of the type species, Pandinoides cavimanus (Pocock, 1888), a revalidation and redescription of Pandinoides militaris (Pocock, 1900), and a description of Pandinoides duffmackayi, sp. nov. Pending reassessment of the genera and subgenera of Pandinus, sensu lato, based on quantitative phylogenetic analysis, Pandinoides is restricted to the three species with a marked concave depression in the retrodorsal surface of the pedipalp chela manus of the adult male, and Pandinus platycheles Werner, 1916, transferred to Pandinus subgenus Pandinoriens Rossi, 2015, creating a new combination: Pandinus (Pandinoriens) platycheles (Werner, 1916), comb. nov. The availability of large series comprising both sexes and all stages of the three Pandinoides species covered herein revealed considerable variation in counts of pedipalp trichobothria, spiniform macrosetae of the leg telotarsi, and pectinal teeth, among and even within individual conspecifics, calling into question the widespread practice of defining species and supraspecific taxa almost exclusively on trivial meristic differences between small samples of material (often singletons, female or immature). Furthermore, whereas neobothriotaxic patterns with low counts may provide appropriate diagnostic characters for genera and species, in combination with other characters, this is generally inadvisable when trichobothrial counts are high, due to the greater instability of the patterns. The second part of this contribution assesses the validity of several putative species of Pandinus, sensu lato, recently described or revalidated, in light of data presented in the first part, and presents 10 new synonyms: Heterometrus roeseli Simon, 1872 = Pandinus (P.) imperator (C.L. Koch, 1841), syn. nov.; Pandinus (P.) camerounensis Lourenço, 2014 = Pandinus (P.) imperator (C.L. Koch, 1841), syn. nov.; Pandinurus (P.) prendinii Rossi, 2015 = Pandinurus (P.) sudanicus (Hirst, 1911), syn. nov.; Pandinurus (Pandicaporiaccous) Rossi, 2015 = Pandinurus (Pandiborellius) Rossi, 2015, syn. nov.; Pandinurus (Pandicaporiaccous) janae Rossi, 2015 = Pandinurus (Pandiborellius) percivali (Pocock, 1902), syn. nov.; Pandinurus (Pandipalpus) bartolozii Rossi, 2015 = Pandinurus (Pandipalpus) viatoris (Pocock, 1890), syn. nov.; Pandinurus (Pandipalpus) flagellicauda Rossi, 2015 = Pandinurus (Pandipalpus) viatoris (Pocock, 1890), syn. nov.; Pandinurus (Pandipalpus) lorenzoi Rossi, 2015 = Pandinurus (Pandipalpus) viatoris (Pocock, 1890), syn. nov.; Pandinurus (Pandipalpus) pantinii Rossi, 2015 = Pandinurus (Pandipalpus) viatoris (Pocock, 1890), syn. nov.; Pandinurus (Pandipalpus) pygmaeus Rossi, 2015 = Pandinurus (Pandipalpus) viatoris (Pocock, 1890), syn. nov. Copyright © 2016 American Museum of Natural History.

Gonzalez-Santillan E.,New York University | Gonzalez-Santillan E.,Scorpion Systematics Research Group | Prendini L.,Scorpion Systematics Research Group
Bulletin of the American Museum of Natural History | Year: 2013

The endemic North American vaejovid scorpion subfamily Syntropinae Kraepelin, 1905, is redefined and its component genera revised, based on a simultaneous phylogenetic analysis of 250 morphological characters and 4221 aligned DNA nucleotides from three mitochondrial and two nuclear gene markers. Tribe Stahnkeini Soleglad and Fet, 2006, is removed from Syntropinae. Tribe Paravaejovini Soleglad and Fet, 2008, and subtribe Thorelliina Soleglad and Fet, 2008, are abolished: Paravaejovini Soleglad and Fet, 2008 = Syntropinae Kraepelin, 1905, syn. nov.; Thorelliina Soleglad and Fet, 2008 = Syntropinae Kraepelin, 1905, syn. nov. Eleven genera, six newly described, are recognized within Syntropinae: Balsateres, gen. nov.; Chihuahuanus, gen. nov.; Kochius Soleglad and Fet, 2008; Konetontli, gen. nov.; Kuarapu Francke and Ponce-Saavedra, 2010; Maaykuyak, gen. nov.; Mesomexovis, gen. nov.; Paravaejovis Williams, 1980; Syntropis Kraepelin, 1900; Thorellius Soleglad and Fet, 2008; Vizcaino, gen. nov. Hoffmannius Soleglad and Fet, 2008, is abolished: Hoffmannius Soleglad and Fet, 2008 = Paravaejovis Williams, 1980, syn. nov. Lissovaejovis Ponce-Saavedra and Beutelspacher, 2001 nomen nudum = Paravaejovis Williams, 1980, syn. nov. Ten species, formerly placed in Hoffmannius, are transferred to Paravaejovis: Paravaejovis confusus (Stahnke, 1940), comb. nov.; Paravaejovis diazi (Williams, 1970), comb. nov.; Paravaejovis eusthenura (Wood, 1863), comb. nov.; Paravaejovis flavus (Banks, 1900), comb. nov. nomen dubium Paravaejovis galbus (Williams, 1970), comb. nov.; Paravaejovis gravicaudus (Williams, 1970), comb. nov.; Paravaejovis hoffmanni (Williams, 1970), comb. nov.; Paravaejovis puritanus (Gertsch, 1958), comb. nov.; Paravaejovis spinigerus (Wood, 1863), comb. nov.; Paravaejovis waeringi (Williams, 1970), comb. nov. Paravaejovis schwenkmeyeri (Williams, 1970), comb. nov., is removed from synonymy. Four species, formerly placed in Kochius, are transferred to Chihuahuanus, gen. nov.: Chihuahuanus cazieri (Williams, 1968), comb. nov.; Chihuahuanus crassimanus (Pocock, 1898), comb. nov.; Chihuahuanus kovariki (Soleglad and Fet, 2008), comb. nov.; Chihuahuanus russelli (Williams, 1971), comb. nov. Four species, formerly placed in Kochius, Thorellius, or Vaejovis C.L. Koch, 1836, are transferred to Mesomexovis, gen. nov.: Mesomexovis atenango (Francke and González-Santillán, 2007), comb. nov.; Mesomexovis oaxaca (Santibáñez-López and Sissom, 2010), comb. nov.; Mesomexovis occidentalis (Hoffmann, 1931), comb. nov.; Mesomexovis subcristatus (Pocock, 1898), comb. nov. Mesomexovis variegatus (Pocock, 1898), comb. nov., is reinstated to its original rank as species. Four subspecies are newly elevated to species: Kochius barbatus (Williams, 1971), stat. nov.; Kochius cerralvensis (Williams, 1971), stat. nov.; Kochius villosus (Williams, 1971), stat. nov.; Mesomexovis spadix (Hoffmann, 1931), comb. et stat. nov. Three subspecies are synonymized: Vaejovis diazi transmontanus Williams, 1970 = Paravaejovis diazi (Williams, 1970), syn. nov.; Vaejovis bruneus loretoensis Williams, 1971 = Kochius villosus (Williams, 1971), syn. nov.; Vaejovis hoffmanni fuscus Williams, 1970 = Paravaejovis hoffmanni (Williams, 1970), syn. nov. © 2013 American Museum of Natural History.

Prendini L.,Scorpion Systematics Research Group
African Invertebrates | Year: 2015

The scorpion fauna of southern Africa is very diverse, especially in the arid western half of the subcontinent. New species continue to be discovered as the region is surveyed with ultraviolet light detection methods. The present contribution describes Uroplectes ansiedippenaarae sp. n., which is endemic to the Succulent Karoo Biome in the Northern Cape and Western Cape provinces of South Africa. The new species appears to be most closely related to U. variegatus (C.L. Koch, 1844), which is endemic to the Fynbos Biome in the Western Cape Province. Uroplectes ansiedippenaarae sp. n. is the smallest species of Uroplectes Peters, 1861, and among the smallest scorpion species in southern Africa, with adults ranging from 16–20 mm in total length. The addition of this new species raises the number of Uroplectes species and subspecies in South Africa to 19, and the number of endemics to 10. © FUNPEC-RP.

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