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On the Phylogenetic Placement of the Spider Genus Atimiosa Simon, 1895, and the Circumscription of Dolichognatha OP-Cambridge

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On the Phylogenetic Placement of the Spider Genus Atimiosa Simon, 1895, and the Circumscription of Dolichognatha OP-Cambridge PUBLISHED BY THE AMERICAN MUSEUM OF NATURAL HISTORY CENTRAL PARK WEST AT 79TH STREET, NEW YORK, NY 10024 Number 3683, 19 pp., 12 figures March 4, 2010 On the Phylogenetic Placement of the Spider Genus Atimiosa Simon, 1895, and the Circumscription of Dolichognatha O.P.-Cambridge, 1869 (Tetragnathidae, Araneae) DIMITAR DIMITROV,1 FERNANDO A´ LVAREZ-PADILLA,2,* AND GUSTAVO HORMIGA3 ABSTRACT The genus Atimiosa Simon, 1895, is a junior synonym of Dolichognatha O. P.-Cambridge, 1869. This synonymy is strongly supported by cladistic analyses of morphological characters and examination of types of all known Atimiosa species. Two new combinations resulted from this nomenclatural change, Dolichognatha comorensis (Schmidt and Krause, 1993), new combination, and Dolichognatha quinquemucronata (Simon, 1895), new combination. New illustrations and photographs of these two species and of the poorly known Dolichognatha longiceps (Thorell, 1895) are provided. We also describe for the first time the web architecture of D. longiceps. INTRODUCTION species diversity is concentrated in the humid tropical regions of the world. More than 900 Tetragnathidae is a family of orbweaving valid species of tetragnathids have been spiders found worldwide; however, most of its described, classified in 47 genera (Platnick, 1 Department of Biological Sciences, the George Washington University, Washington, D.C. 20052, USA (dimitard. [email protected]). 2 California Academy of Sciences, Department of Entomology, 55 Music Concourse Drive, Golden Gate Park, San Francisco, CA 94118, USA. 3 Division of Invertebrate Zoology, American Museum of Natural History, New York. Department of Biological Sciences, the George Washington University, Washington, D.C. 20052, USA ([email protected]). *Current address: Universidad Nacional Auto´noma de Me´xico, Facultad de Ciencias, Departamento de Biologı´a Comparada, Laboratorio de Acarologı´a, s/n Ciudad Universitaria, Me´xico D. F., Del. Coyoaca´n, C.P. 04510, Me´xico ([email protected]). Copyright E American Museum of Natural History 2010 ISSN 0003-0082 2 AMERICAN MUSEUM NOVITATES NO. 3683 2010). Many tetragnathid genera are associat- known tetragnathid genera from Australasia ed with ecosystems neighboring freshwater and Africa have produced six generic synony- bodies, such as lakes or rivers, where their mies for Dolichognatha (Levi, 1981; Smith, population density is often very high, making 2008). In addition, they have also provided them the dominating orbweavers in the illustrated descriptions of several Dolicho- habitat. Members of the family are also gnatha species. We extend these efforts to common in humid tropical forests where they facilitate identification and future taxonomic are found from the ground level to the top of work by providing illustrations and photo- the canopy. As with many other tropical graphs of all species formerly included in arthropods, most of the diversity is still Atimiosa. Specimens of the poorly known undescribed. Inadequate descriptions and species Dolichognatha longiceps (Thorell, poor knowledge of many of the species further 1895) are also illustrated and included in the complicates any systematic work, hence the analyses to better test the monophyly of the need for generic systematic revisions to genus; that species was also selected because of address this problem. In recent years, several its unusual morphological features, such as the authors have addressed different aspects of absence of PME. Here we provide additional tetragnathid systematics (e.g., Tanikawa, 2001; illustrations of D. longiceps and document for Gillespie, 2003a, 2003b; A´ lvarez-Padilla, 2007; the first time its web architecture. Dimitrov et al., 2008; Smith, 2008; Dimitrov ´ and Hormiga, 2009; Alvarez-Padilla et al., MATERIALS AND METHODS 2009), but due to the large size of the group many genera still require systematic revisions. Type specimens for this study were bor- The present paper is a continuation of our rowed from the museum collections listed at efforts to update tetragnathid systematics. It the end of this section. Specimens from focuses on the South Asian genus Atimiosa Dolichognatha longiceps were collected by Simon, 1895. We have examined the type GH in Thailand. Digital photographs of the specimens of the two known species of type in the Muse´um National d’Histoire Atimiosa and studied their phylogenetic posi- Naturelle, Paris (MNHN) were taken by tion within Tetragnathidae using morpholog- FAP with a Nikon Coolpix 995 camera while ical characters. The type species, Atimiosa visiting the collections. The morphological quinquemucronata Simon, 1895, was originally methods of our study follow those previously described by Simon (1895) based on a described in Hormiga (2002). Other specimens subadult female specimen (fig. 1A–E) from were examined and illustrated using Leica Sri Lanka that is conspicuously similar to MZ16 or Leica MZ16A stereoscopic micro- some Dolichognatha species. Schmidt and scopes with a camera lucida and Leica Krause (1993) subsequently described another DMRM compound microscope with a draw- species, Atimiosa comorensis Schmidt and ing tube. Drawings were prepared with graph- Krause, 1993, from the Comoro Islands. It is ite pencils on acid-free cotton paper. Hairs in fact puzzling how Schmidt and Krause and macrosetae are not depicted in the final (1993) may have decided on this generic drawings. For male palp illustrations the placement given that Simon’s description left palp was used. Epigyna were treated with mentions very few diagnostic traits, all of them SIGMA Pancreatin LP 1750 enzyme complex consistent with Dolichognatha. A rationale for (A´ lvarez-Padilla and Hormiga, 2008) and the generic placement is absent from Schmidt transferred to methyl salicylate solution for and Krause’s paper. In this paper we propose examination and illustration. the synonymization of the genus Atimiosa with All pencil drawings were scanned and Dolichognatha based on the examination of the further improved with the help of the Gimp type specimens of both species and on the 2.4 and Adobe Photoshop CS2 programs. results of cladistic analyses that include Digital images of the specimens were taken in all known Atimiosa species and a broad alcohol with Nikon DXM1200F digital cam- sample of tetragnathids. Recent treatments of era mounted on a Leica MZ16A stereoscopic Dolichognatha and several small and poorly microscope. Final plate layout and editing was 2010 DIMITROV ET AL.: SPIDER GENUS ATIMIOSA SIMON, 1895 3 Fig. 1. Dolichognatha quinquemucronata (Simon, 1895), new combination. Subadult female holotype from Sri Lanka (MNHM 16140): ventral (A), dorsal (B). Cephalothorax: dorsal (C), frontal (D). Abdomen and genital area, ventral (E). done with Adobe Illustrator CS2. All mea- MCZ Museum of Comparative Zoology, surements are in millimeters. Cambridge, USA MNHN Muse´um National d’Histoire Nat- urelle, Paris, France ABBREVIATIONS USED IN TEXT AND FIGURES NRM Naturhistoriska riksmuseet, Stock- ALE anterior lateral eyes holm, Sweden C conductor RMCA Muse´e royal de l’Afrique centrale, CB cymbium Tervuren, Belgium CEBP cymbial ecto-basal process CD copulatory duct PHYLOGENETICS E embolus FD fertilization duct Morphological characters for Atimiosa co- MEA metine embolic apophysis morensis, A. quinquemucronata, Dolichognatha MPT maximum parsimony tree incaensis (Simon, 1895), D. longiceps, and P paracymbium Cyrtognatha insolita (Chickering, 1956) were PLE posterior lateral eyes scored (Appendix 1) and added to the PME posterior median eyes morphological matrix of A´ lvarez-Padilla et S spermatheca ´ ST subtegulum al. (2009). That work (Alvarez-Padilla et al., T tegulum 2009) provides the largest and most complete morphological data matrix for Tetragnathidae to date, along with a large number of out- MUSEUM COLLECTIONS group representatives. This relatively extensive AMNH American Museum of Natural His- taxon sample, coupled with the large number tory, New York, USA of characters (213 morphological and behav- 4 AMERICAN MUSEUM NOVITATES NO. 3683 ioral characters), allows robust testing of are based on direct examination of the holo- phylogenetic hypotheses for the relationships type (male from Panama, Barro Colorado of tetragnathid genera. Here we use that Island, in MCZ 21668) in addition to the SEM dataset to test the phylogenetic position of images, drawings, and descriptions provided by the tetragnathid genus Atimiosa. We retain the Dimitrov and Hormiga (2009). One character- original taxon sample of A´ lvarez-Padilla et al. istic of C. insolita, the presence of a ‘‘metine (2009) and add to the matrix the taxa listed embolic apophysis’’ (MEA), is particularly above and in Appendix 1. relevant as most of the species of Cyrtognatha Very few characters were scored for A. have this apophysis (Dimitrov and Hormiga, quinquemucronata because the only specimen 2009), which is absent in C. espanola (Bryant, available for study was the subadult type. 1945). The latter species was used in the phylo- Nonetheless, we have added this species to the genetic analyses of A´ lvarez-Padilla et al. (2009). matrix because it is the type species of the genus Dolichognatha has an MEA and the proper Atimiosa. Several somatic characters reflecting representation of the dominant embolic mor- traits such as eye size and distribution, which do phology in Cyrtognatha is important for the not show sexual dimorphism, were scored from correct optimization of this character. To reflect the male type of A. comorensis. Dolichognatha
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