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Evolution of Scorpion Orthobothriotaxy: a Cladistic Approach

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Evolution of Scorpion Orthobothriotaxy: a Cladistic Approach Euscorpius Occasional Publications in Scorpiology EDITOR: Victor Fet, Marshall University, ‘[email protected]’ ASSOCIATE EDITOR: Michael E. Soleglad, ‘[email protected]’ Euscorpius is the first research publication completely devoted to scorpions (Arachnida: Scorpiones). Euscorpius takes advantage of the rapidly evolving medium of quick online publication, at the same time maintaining high research standards for the burgeoning field of scorpion science (scorpiology). Euscorpius is an expedient and viable medium for the publication of serious papers in scorpiology, including (but not limited to): systematics, evolution, ecology, biogeography, and general biology of scorpions. Review papers, descriptions of new taxa, faunistic surveys, lists of museum collections, and book reviews are welcome. Derivatio Nominis The name Euscorpius Thorell, 1876 refers to the most common genus of scorpions in the Mediterranean region and southern Europe (family Euscorpiidae). Euscorpius is located on Website ‘http://cos-server.marshall.edu/euscorpius’ at Marshall University, Huntington, WV 25755-2510, USA. The International Code of Zoological Nomenclature (ICZN, 4th Edition, 1999) does not accept online texts as published work (Article 9.8); however, it accepts CD-ROM publications (Article 8). Euscorpius is produced in two identical versions: online (ISSN 1536-9307) and CD-ROM (ISSN 1536-9293). Only copies distributed on a CD-ROM from Euscorpius are considered published work in compliance with the ICZN, i.e. for the purposes of new names and new nomenclatural acts. All Euscorpius publications are distributed on a CD-ROM medium to the following museums/libraries: • USNM, United States National Museum of Natural History (Smithsonian Institution), Washington, DC, USA • AMNH, American Museum of Natural History, New York, USA • CAS, California Academy of Sciences, San Francisco, USA • FMNH, Field Museum of Natural History, Chicago, USA • MCZ, Museum of Comparative Zoology, Cambridge, Massachusetts, USA • MNHN, Museum National d’Histoire Naturelle, Paris, France • NMW, Naturhistorisches Museum Wien, Vienna, Austria • BMNH, British Museum of Natural History, London, England, UK • MZUC, Museo Zoologico “La Specola” dell’Universita de Firenze, Florence, Italy • ZISP, Zoological Institute, Russian Academy of Sciences, St. Petersburg, Russia • WAM, Western Australian Museum, Perth, Australia Publication date: 14 December 2001 Euscorpius — Occasional Publications in Scorpiology. 2001, No. 1 Evolution of Scorpion Orthobothriotaxy: A Cladistic Approach Michael E. Soleglad 1 and Victor Fet 2 1 P. O. Box 250, Borrego Springs, California 92004, USA 2 Department of Biological Sciences, Marshall University, Huntington, West Virginia 25755-2510, USA Summary 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 plesiomor- phic 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 ear- liest known complete fundamental trichobothrial pattern, and the pseudochactids (Type D), the fourth fundamental pattern for Recent scorpions. Introduction other character analyses in the aforementioned compan- ion paper. As stated by Gromov (1998, p. 1003) ‘It is The goal of this study is to investigate the evolution of possible that the representative of this new family scorpion orthobothriotaxic patterns using cladistic tech- (Pseudochactidae) stands close to the common ancestor niques. Our secondary goal, in support of a companion of all these families (the Recent scorpions)’ --- Pseudo- paper (Soleglad & Fet, in progress), is to determine the chactas is certainly a basal member of Recent scorpions phylogenetic placement of the recently described and and, based on our current research, also shows signifi- unique scorpion Pseudochactas ovchinnikovi from Cen- cant affinity in some characters with the Carboniferous tral Asia (Gromov, 1998). The satisfaction of these goals fossil scorpion family Palaeopisthacanthidae. Again, is further enhanced by two events: (1) the analysis of these comparisons are only made possible due to fossil scorpion trichobothria patterns conducted by Jeram’s (1994) comprehensive study of this group, Jeram (1994) and Lourenço (2001); and (2) the obtain- Gromov’s thorough description and excellent illustra- ment of actual specimens of P. ovchinnikovi for study. tions of Pseudochactas, and the availability of speci- With Jeram and Lourenço’s important contributions we mens of the latter. are able for the first time to investigate the evolution of the basic trichobothrial patterns originally established by Use of trichobothria in scorpion systematics Vachon (1972, 1974) by incorporating fossil patterns. This effort is of particular interest since Pseudochactas Max Vachon, in his monumental treatise on the formal itself represents another basic orthobothriotaxic pattern presentation of scorpion trichobothrial patterns (Vachon, type for Recent scorpions (in addition to the three basic 1972, 1974), established the importance of trichobothria patterns defined by Vachon (1972, 1974)). In satisfac- in upper level systematics by showing significant con- tion of the second goal, the phylogeny of Pseudochac- sistency across large numbers of genera in their basic tas, the trichobothria homologies established in this pa- patterns (i.e., orthobothriotaxy, as opposed to neoboth- per will become major characters that are combined with riotaxy, a comparatively rare deviation from one of these 2 Euscorpius — 2001, No. 1 patterns). In this analysis, Vachon defined three funda- Trichobothria Homology mental orthobothriotaxic patterns that accounted for all recent scorpions. For example, the buthids, the largest In order to conduct a meaningful cladistic analysis, scorpion aggregate, exhibited a pattern (Type A) not character homologies must be carefully established found in any other scorpion group. Similarly, a distinct across all taxa involved in the analysis. This aspect of pattern (Type B) was observed in all species of the ge- cladistics is by far the most important and also the most nus Chaerilus, a taxon considered disjoint from all other difficult since final results are totally dependent on these scorpions and constituting a monotypic family Chaerili- established homologies. In this section, we present the dae. All other genera, spanning widely diverse groups ‘existence’ approach to trichobothria evolution, an ap- conformed to a third pattern (Type C), a pattern consid- proach that directly addresses individual trichobothria. erably different from the other two. The significance of We present in detail proposed homologies across all Vachon’s three trichobothria patterns was quite appeal- fundamental orthobothriotaxic pattern types for each pe- ing since they also reflected, in general terms, the cur- dipalp segment --- the chela, patella and femur. In those rent breakdown in upper-level scorpion systematics. situations where we established designations different Sissom (1990) also considered these three fundamental from those of the original authors, we exercised the ap- patterns of key importance in scorpion systematics, il- propriate cladistic sequences to determine whether the lustrating the three patterns in detail as one of the im- alternative designations were consistent with overall portant characters sets used in his overall diagnoses of parsimony (i.e., if any alternative designation increased Recent scorpion families. These illustrations were also the number of steps, it was excluded from considera- accompanied by detailed trichobothria statistical data tion). In order to take advantage of the important tricho- based on segment and surface (see Sissom, 1990, Table bothria information now available on fossil scorpions 3.1). Furthermore, Sissom (1990) also used trichoboth- (e.g., Jeram, 1994, Lourenço, 2001) we provide a de- rial patterns as his primary couplet in his extensive key tailed overview of our current knowledge of all reported to the buthid genera, segregating genera by the femoral trichobothria in fossil scorpions. This data will be used alpha and beta patterns originally defined by Vachon in establishing homologies for scorpion families Palae- (1975a). Fet & Rechkin (1990) presented the first quan- opisthacanthidae and Archaeobuthidae, two components titative (non-cladistic) analysis of all known variations
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