Evolution of Scorpion Orthobothriotaxy: A Cladistic Approach

This study presents a cladistic analysis of the derivation of orthobothriotaxic patterns in scorpions. Included in this analysis are the original three orthobothriotaxic patterns defined by Vachon (1972, 1974), the pattern of the unique scorpion Pseudochactas ovchinnikovi Gromov, 1998, and two trichobothrial patterns of fossil scorpions, the Upper Carboniferous palaeopisthacanthids and the Lower Cretaceous archaeobuthids. An overview of all fossil scorpion material where trichobothria are reported is presented in detail. The approach used in this analysis is to model the existence of an individual trichobothrium, adopting the ‘absence of’, ‘petite size’ and ‘full size’ as incremental stages of a trichobothrium’s development. Of particular interest is the phylogenetic placement of Pseudochactas within Recent scorpions, for which the results of this study provide preliminary insight. Phylogenetic results of this analysis, based entirely on the derivation of orthobothriotaxic patterns, show that Archaeobuthus is the plesiomorphic sister group of all Recent scorpions, placed between the ancient palaeopisthacanthids and Recent scorpions. Within Recent scorpions, the clades ‘buthids + pseudochactids’ and ‘chaerilids + Type C’ are strongly endorsed by this analysis. Formal orthobothriotaxic types are defined for the palaeopisthacanthids (Type P), representing the earliest known complete fundamental trichobothrial pattern, and the pseudochactids (Type D), the fourth fundamental pattern for Recent scorpions

Contributions to Scorpion Systematics. I. On Recent Changes in High-Level Taxonomy

Prendini & Wheeler (2005) criticized the methods of phylogenetic analyses by Soleglad, Fet, and their coauthors, and executed an unprecedented taxonomic action: without analyzing any of these taxa, they performed a wholesale synonymization of four parvorders, eight superfamilies, one family, 11 subfamilies, eight tribes, two subtribes, and three genera (in total, 37 taxa) of scorpions, and made other taxonomic changes. No alternative new classification has been proposed (instead, they revert to a previous classification), and no results of original work on this subject by Prendini & Wheeler (2005) have been presented. Here, we reverse all taxonomic changes performed by Prendini & Wheeler (2005) since we do not consider these actions justified. We comment on a few issues pertaining to the International Code of Zoological Nomenclature

A new scorpion genus (Scorpiones: Vaejovidae) from Mexico

A new vaejovid genus from Mexico, Franckeus, gen. nov., is described based on unique neobothriotaxy. Species placed in this genus are from the Vaejovis “nigrescens” group (previously called the “nitidulus” group). Six species comprise this new genus, distributed throughout Mexico (mainland as well as Baja California Sur): Franckeus nitidulus, F. rubrimanus, F. platnicki, F. minckleyi, F. kochi and F. peninsularis. A new species of the Vaejovis “nigrescens” group, Vaejovis davidi, sp. nov., is also described

The Scorpion Sternum: Structure and Phylogeny (Scorpiones: Orthosterni)

The structure of the sternum of all major Recent scorpion groups is analyzed in detail. Based on this analysis, two fundamental sternum types are identified, described and illustrated, type 1 and type 2. These sternum types are distinguished by criteria based on external and internal structural features. The sternum types described herein are offered as a replacement for the various characterizations used throughout the last 140 years which emphasize only gross overall shape and proportions. Phylogenetic and taxonomic ramifications of these new sternal types are discussed. The Carboniferous fossil scorpion Palaeopisthacanthus schucherti Petrunkevitch is assigned to sternum type 1. The type 1 sternum is also assigned to the three primitive Recent scorpion groups, the pseudochactids, the buthoids, and the chaerilids. Sternum type 2 is defined for the remaining scorpion groups, those complying with Type C orthobothriotaxy. Based on these assignments, sternum type 1 is considered primitive. Within these two basic sternal types a hypothesis of horizontal and vertical compression is offered as a cause-effect for the unique sterna of the buthoid and bothriurid scorpions

Morphology analysis supports presence of more than one species in the \u3ci\u3e“Euscorpius carpathicus”\u3c/i\u3e complex (Scorpiones: Euscorpiidae)

We investigate a number of scorpion populations from southern and central Europe, commonly classified under a “catch-all” name of Euscorpius carpathicus (L., 1767). This species includes a high number of described subspecies but its composition is not resolved. The detailed morphology analysis in the present paper includes a number of new characters, in particular individually mapped external patellar accessory trichobothria. It suggests that several clearly separated lineages are present. E. carpathicus (L.) is restricted here to geographically marginal populations from Romania (terra typica), which exhibit loss of one trichobothrium in the patellar series em (= 3). Another lineage (Austria, Croatia, Italy, France, Slovenia) is characterized here as E. tergestinus (C.L. Koch, 1837); it has a “standard” trichobothrial number in the patellar series eb (= 4), eba (= 4) and em (= 4) and exhibits only variation in the ventral and et series. This species includes as new synonyms the following seven subspecies described by Caporiacco (1950): E. c. apuanus, E. c. concinnus, E. c. niciensis, E. c. aquilejensis, E. c. picenus, E. c. oglasae, and E. c. corsicanus. A very distinct Balkan lineage is delineated based on unique trichobothrial numbers in patellar series eb (= 5) and eba (= 7); it is elevated here to the species status as E. hadzii Caporiacco, 1950 (Albania, Bosnia, Bulgaria, Croatia, Greece, Macedonia, Yugoslavia). This species includes as a new synonym E. c. lagostae Caporiacco, 1950. The fourth species-rank taxon confirmed here is E. koschewnikowi Birula, 1900 (Greece), with “standard” trichobothrial number in the patellar eb (= 4), eba (= 4) and em (= 4) series but with other unique morphological features. We fix neotypes of E. tergestinus and E. hadzii, and a lectotype of E. koschewnikowi. These four species and E. balearicus Caporiaccebao, another member of this complex, are contrasted in detail using trichobothrial patterns, morphometric ratios and carinal development trends as diagnostic characters

EDITORIAL: \u3cem\u3eEuscorpius\u3c/em\u3e at 100

With the publication of Graeme Lowe on Vachoniolus from Oman, our journal Euscorpius celebrates its 100th issue. Euscorpius is the first and only research publication completely devoted to scorpions (Arachnida: Scorpiones). Since its inception in 2001, Euscorpius published 100 issues authored by 79 zoologists from 23 countries (USA, Argentina, China, Colombia, Cuba, Czech Republic, France, Germany, Greece, Guadeloupe, India, Iran, Italy, Kazakhstan, Mexico, the Netherlands, Norway, Pakistan, Russia, Switzerland, Turkey, United Kingdom, and Venezuela.

Contributions to scorpion systematics. III. Subfamilies Smeringurinae and Syntropinae (Scorpiones: Vaejovidae)

Stockwell (1989), in an unpublished revision, suggested splitting the family Vaejovidae in two subfamilies, introducing also four tribes and four new genera. These results have not been reevaluated for 20 years. Here, we establish a new subfamily Smeringurinae, subfam. nov., with two tribes, four genera, and 40 species and subspecies. The subfamily Smeringurinae includes two tribes: Smeringurini, trib. nov. and Paravaejovini, trib. nov. The tribe Smeringurini includes three genera: Paruroctonus Werner, 1934; Smeringurus Haradon, 1983; and Vejovoidus Stahnke, 1974. The tribe Paravaejovini includes the genus Paravaejovis Williams, 1980. We restore from synonymy the subfamily Syntropinae Kraepelin, 1905, and significantly expand its scope to include two tribes, two subtribes, eight genera, and 75 species and subspecies. The subfamily Syntropinae includes two tribes: Stahnkeini Soleglad et Fet, 2006, and Syntropini Kraepelin, 1905. The tribe Stahnkeini includes four genera: Gertschius Graham et Soleglad, 2007; Serradigitus Stahnke, 1974; Stahnkeus Soleglad et Fet, 2006; and Wernerius, gen. nov. The tribe Syntropini includes two subtribes, Syntropina Kraepelin, 1905, and Thorelliina, subtrib. nov. The subtribe Syntropina includes two genera, Syntropis Kraepelin, 1900 and Hoffmannius, gen. nov. The subtribe Thorelliina includes two new genera, Thorellius, gen. nov. and Kochius, gen. nov. Four new genera accommodate species from the former “eusthenura,” “intrepidus,” “punctipalpi,” and “spicatus” groups of Vaejovis. One new species from Durango, Mexico is described, Kochius kovariki Soleglad et Fet, sp. nov. Thorellius cristimanus, stat. nov. and T. atrox, stat. nov. are elevated to species from subspecies level. Our results confirm, in part, the phylogeny proposed by Stockwell (1989), with its further splitting of the overly inflated genus Vaejovis. The remaining default nominotypic subfamily, Vaejovinae Thorell, 1876, still unrevised at this time, is reduced to 66 species and subspecies, grouped in four genera, Franckeus, Pseudouroctonus, Uroctonites, and Vaejovis (the latter being divided into the “mexicanus” and “nigrescens” groups). Phylogenetic relationships within subfamilies Smeringurinae, Syntropinae, and Vaejovinae are discussed. As a result of the current revision, the family Vaejovidae now includes three subfamilies, four tribes, two subtribes, and 15 genera, with a total of 181 species-group taxa

\u3ci\u3eProceedings of the 3rd Scorpiology Symposium\u3c/i\u3e (American Arachnological Society, 28th Annual Meeting)

Table of Contents Victor Fet and Douglas D. Gaffin. Preface p. iKaren C. Bost and Douglas D. Gaffin. Sand scorpion home burrow navigation in the laboratory p. 1Douglas D. Gaffin and Mark E. Walvoord. Scorpion peg sensilla: are they the same or are they different? p. 7J. Zachary Porterfield, Douglas D. Gaffin, Caitlin Porterfield and Curtis Johnston. Screening for scorpions: A non-invasive approach to tracking the movements of arachnids in sand p. 17Tsunemi Yamashita. Surface activity, biomass, and phenology of the striped scorpion, Centruroides vittatus (Scorpiones: Buthidae) in Arkansas, USA p. 25C. Neal McReynolds. Temporal patterns in microhabitat use for the scorpion Centruroides vittatus (Scorpiones: Buthidae) p. 35Victor Fet, Michael E. Soleglad and Benjamin Gantenbein. The Euroscorpion: Taxonomy and systematics of the genus Euscorpius Thorell, 1876 (Scorpiones: Euscorpiidae) p.47Victor Fet, Michael E. Soleglad and Alexander Gromov. The platypus of a scorpion: the genus Pseudochactas Thorell, 1876 (Scorpiones: Pseudochactidae) p. 6

High-level systematics and phylogeny of the extant scorpions (Scorpiones: Orthosterni)

A number of authors (e. g. Birula, 1917a, 1917b; Mello-Leitão, 1945; Stockwell, 1989) addressed above-level systematics of extant scorpions, and accepted the grouping of scorpion families in several superfamilies. At the same time, Kjellesvig-Waering (1986) classified all extant scorpions under the same superfamily, Scorpionoidea. Sissom (1990) and Fet et al. (2000) did not list any superfamilies, considering the systematic situation above family (and often at the family level as well) unresolved. Most recently, Lourenço (2000a) listed six superfamilies, largely following the unpublished but important study of Stockwell (1989). The goal of this paper is to address scorpion systematics and phylogeny above genus level. We conducted a comprehensive, cladistic morphological analysis of 90 extant genera (over 150 species) of scorpions belonging to all recognized families. We especially concentrated on relationships among so-called “chactoid” scorpions, where subfamilies, tribes, and subtribes were revised and/or established. The family Chactidae was given a special attention due to the number of phylogenetic and taxonomic issues that were revised. In addition, we addressed the status of a recently discovered, unique relict family Pseudochactidae, and the systematic relationships within Iuridae. As a result of intensive study, we propose a number of sweeping changes in current scorpion taxonomy; the results of analyses leading to these changes are discussed in detail. The category of parvorder, subordinate to infraorder, is introduced for the first time in arachnid systematics. Four extant parvorders are recognized within the scorpion infraorder Orthosterni: Buthida, Chaerilida, Pseudochactida, and Iurida. Six extant superfamilies are recognized: Buthoidea, Chactoidea (=Vaejovoidea, syn. n), Chaeriloidea, Iuroidea (new), Pseudochactoidea (new), and Scorpionoidea (=Bothriuroidea, syn. n). Parvorders Buthida, Chaerilida and Pseudochactida are monotypic, each including a single superfamily; parvorder Iurida includes three superfamilies (Chactoidea, Iuroidea, and Scorpionoidea). We recognize 14 extant scorpion families: Bothriuridae, Buthidae, Caraboctonidae, Chactidae, Chaerilidae, Euscorpiidae, Iuridae, Liochelidae, Microcharmidae, Pseudochactidae, Scorpionidae, Superstitioniidae, Urodacidae, and Vaejovidae. Superfamilies Chaeriloidea and Pseudochactoidea are monotypic; superfamily Buthoidea includes two families (Buthidae and Microcharmidae). Superfamily Iuroidea includes two families (Caraboctonidae and Iuridae); subfamily Caraboctoninae (formerly in Iuridae) is elevated to the family rank. Superfamily Chactoidea includes four families: Chactidae, Euscorpiidae, Superstitioniidae (=Troglotayosicidae, syn. n), and Vaejovidae. Within Chactidae, three subfamilies are established: Chactinae, Brotheinae, and Uroctoninae. Within Chactinae, two tribes are established: Chactini and Nullibrotheini, new tribe (monotypic). Within new subfamily Brotheinae, two tribes are established: Brotheini and Belisariini (monotypic). Within Brotheini, two subtribes are established: Brotheina and Neochactina, new subtribe; the latter is based on a new genus, Neochactas, gen. n. Within Brotheina, genera Cayooca, Guyanochactas, and Taurepania are synonymized with Broteochactas. Subfamily Uroctoninae is restored from synonymy under Vaejovidae and transferred to Chactidae; it includes genera Uroctonus and Anuroctonus (the latter transferred from the erstwhile Iuridae). Family Troglotayosicidae is abolished, and its two genera are transferred to other families: Troglotayosicus, to Superstitioniidae; and Belisarius, to Chactidae. Subfamily Belisariinae is downgraded to the tribe rank and transferred to Chactidae (subfamily Brotheinae). Superfamily Scorpionoidea includes four families: Bothriuridae, Liochelidae (=Hemiscorpiidae, syn. n.), Scorpionidae (=Diplocentridae, syn. n.), and Urodacidae (=Heteroscorpionidae, syn. n). Family Diplocentridae is downgraded to the subfamily rank in Scorpionidae. Subfamily Nebinae is downgraded to the tribe rank in Diplocentrinae. Family Hemiscorpiidae is downgraded to the subfamily rank in Liochelidae. Family Heteroscorpionidae is downgraded to the subfamily rank in Urodacidae. We provide detailed classification, taxonomic history, and diagnoses of all recognized scorpion taxa above genus level. The phylogeny and biogeographic implications are discussed. As an addition, we present, among other materials, results of the first pilot high-level scorpion DNA phylogeny, including representatives of seven families spanning all four parvorders. Both morphological analysis and DNA sequence analysis support the primitive nature of parvorders Pseudochactida, Buthida, and Chaerilida, as opposed to the derived position of parvorder Iurida. Especially remarkable is the parvorder Pseudochactida, which exhibits many primitive features. Within Iurida, the superfamily Iuroidea is firmly established as a basal group, and Scorpionoidea, as the most derived group. Phylogeny within Chactoidea shows ancient nature of many clades, as our analysis reveals hitherto unexpected relationships between a number of genera and tribes

Contributions to scorpion systematics. II. Stahnkeini, a new tribe in scorpion family Vaejovidae (Scorpiones: Chactoidea)

The diagnostic characters originally established by Herbert L. Stahnke (1940a, 1940b, 1974) in his description of genus Serradigitus are studied in detail from several new perspectives. A new genus, Stahnkeus, gen. nov., is described based on the presence of inner accessory (IAD) denticles on the chelal fingers, unprecedented in family Vaejovidae. Five species of Serradigitus are transferred to Stahnkeus: Stahnkeus harbisoni (Williams, 1970), comb. nov. (=Serradigitus harbisoni); Stahnkeus deserticola (Williams, 1970), comb. nov. (=Serradigitus deserticola); Stahnkeus subtilimanus (Soleglad, 1972), comb. nov. (=Serradigitus subtilimanus); Stahnkeus allredi (Sissom et Stockwell, 1991), comb. nov. (=Serradigitus allredi); and Stahnkeus polisi (Sissom et Stockwell, 1991), comb. nov. (=Serradigitus polisi). In this revision, a new tribe, Stahnkeini, trib. nov. (= Serradigitus + Stahnkeus), is formally described based on three unambiguous synapomorphies. Issues involving the taxonomic placement of species Serradigitus baueri, S. pacificus, S. bechteli and S. littoralis are discussed