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The Placement of the Spider Genus Periegops and the Phylogeny of Scytodoidea (Araneae: Araneomorphae)

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The Placement of the Spider Genus Periegops and the Phylogeny of Scytodoidea (Araneae: Araneomorphae) Zootaxa 3312: 1–44 (2012) ISSN 1175-5326 (print edition) www.mapress.com/zootaxa/ Article ZOOTAXA Copyright © 2012 · Magnolia Press ISSN 1175-5334 (online edition) The placement of the spider genus Periegops and the phylogeny of Scytodoidea (Araneae: Araneomorphae) FACUNDO M. LABARQUE1 & MARTÍN J. RAMÍREZ1 1Museo Argentino de Ciencias Naturales “Bernardino Rivadavia”, Consejo Nacional de Investigaciones Científicas y Técnicas (CONICET), Av. Ángel Gallardo 470, C1405DJR, Buenos Aires, Argentina. [email protected] / [email protected] Abstract The relationships of Scytodoidea, including the families Drymusidae, Periegopidae, Scytodidae and Sicariidae, have been con- tentious for a long time. Here we present a reviewed phylogenetic analysis of scytodoid spiders, emphasizing Periegops, the only genus in the family Periegopidae. In our analysis the Scytodoidea are united by the fusion of the third abdominal entapo- physes into a median lobe, the presence of female palpal femoral thorns and associated cheliceral stridulatory ridges, a mem- branous lobe on the cheliceral promargin, and the loss of minor ampullate gland spigots. A basal split within Scytodoidea defines two monophyletic groups: Sicariidae and a group formed by Scytodidae as the sister group of Periegopidae plus Dry- musidae, all united by having bipectinate prolateral claws on tarsi I–II, one major ampullate spigot accompanied by a nubbin, and the posterior median spinnerets with a mesal field of spicules. Periegops is the sister group of Drymusidae, united by the regain of promarginal cheliceral teeth and a triangular cheliceral lamina, which is continuous with the paturon margin. Key words: Drymusa, Drymusidae, Haplogyne, morphology, Scytodes, Stedocys, Scytodidae, Sicariidae, Sicarius, Loxosceles Introduction The family Periegopidae currently comprises only the genus Periegops, with two species: the type species Perie- gops suteri (Urquhart) from the Banks Peninsula on the South Island of New Zealand (Vink 2006), and Periegops australia Forster, from southeastern Queensland (Forster 1995). A single female of an unidentified Periegops spe- cies from the East Cape region of the North Island of New Zealand was also mentioned by Forster (1995); at pres- ent, the delimitation of species within Periegops is dependent on the detailed structure of the male copulatory bulb, hence the identification of this specimen should wait for new data, although mitochondrial COI molecular evidence suggests that it may be a distinct species (unpublished data mentioned in Vink 2006). The genus Periegops was first included in the subfamily Periegopinae by Simon (1893), and placed in the Sicariidae, along with Drymusinae, Loxoscelinae, Plectreurinae, Scytodinae and Sicariinae. Simon was the first to discover that Periegops and Scytodes had a bipectinate “external superior” tarsal claw on the anterior tarsi. Simon included Diguetia and Pertica (later synonymized with Segestrioides; Brescovit & Rheims 2005) in Periegopinae, but Petrunkevitch (1928) removed them and placed them into Diguetinae. Bryant (1935a, b) suggested a family rank for Periegops that included also Plectreurys and Diguetia, but she took no formal action. Lehtinen (1986) studied only the female holotype of P. suteri. He mentioned that Periegopidae, Drymusidae, Scytodidae and Ochyroceratidae share a particular dentition pattern in the tarsal claws, but he never explained or described this character in detail, so his observations were not conclusive. He also suggested that Periegops could be a strongly apomorphic derivative of the classical (non-monophyletic, Platnick et al. 1991), Scytodoidea, com- posed of the Caponiidae, Diguetidae, Loxoscelidae, Ochyroceratidae, Plectreuridae, Pholcidae, Scytodidae, Sicari- idae and Tetrablemmidae (Brignoli 1975, 1978). Platnick et al. (1991) investigated the spinneret morphology of haplogyne araneomorph spiders (Haplogynae), and produced a cladistic analysis with all haplogyne families except Periegopidae. They obtained a restricted Scyt- odoidea, composed of Sicariidae, Scytodidae, and Drymusidae, in which Scytodidae and Drymusidae were united Accepted by W. Fannes: 29 Dec. 2011; published: 14 May 2012 1 by having a field of spicules on the median surface of the posterior median spinnerets. Ramírez (2000) added a few characters from the respiratory system to the same dataset, further strengthening Platnick et al.’s hypothesis. Gris- wold et al. (2005) reviewed most of the character systems used in those sources, but only scored two families of Haplogynae (Filistatidae and Segestriidae) as outgroups for their analysis. Forster (1995) was the first to examine the tracheal system of Periegops, and their cuticular structures using scanning electron microscopy. He hypothesized that the Periegopidae were related to the ecribellate, haplogyne families Diguetidae, Drymusidae, Plectreuridae, Scytodidae and Sicariidae (that is, Simon’s Sicariidae, plus Diguetidae). Within this group, Forster suggested that Periegopidae was most closely related to the Scytodidae, based on two characters: (a) posterior tracheal system composed of a short median apodemal lobe and a distinct pair of simple lateral tracheae, a configuration also shared with Drymusidae; and (b) a double row of teeth on the ventral surface of the proclaws, and a single row on the retroclaws of leg I and II (later found in Drymusa as well, see below). Lamy (1902) was the first to describe the tracheae of Scytodes and Loxosceles, which both share a fused median apodeme. Ramírez (2000) confirmed that at least in Loxosceles and Sicarius the median lobe retained the muscle insertions corresponding to the paired third abdominal entapophyses, proposing their fusion on a median structure as a further synapomorphy for a clade that included Drymusa, Loxosceles, Scytodes and Sicarius (plus Periegops, although this genus was not included in the analysis). In the cladistic analysis of Ramírez (2000), the elongation of the median apodeme is a synapomorphy for Sicariidae, thus a short median apodeme is ple- siomorphic for Scytodes, Drymusa, and Periegops. Labarque & Ramírez (2007a) reported that the bipectinate pro- claws are also present in Drymusa, and that Periegops also have a field of spicules on the median surface of the posterior median spinnerets (PMS), along with Drymusa and Scytodes. In summary, the more promising characters that may help resolve the closest relationships of Periegops (the bipectinate proclaws, the spicules on the PMS), are shared both with Dymusa and Scytodes. Here we intend to clar- ify the relationships of Periegops and its relatives performing a cladistic analysis of Scytodoid spiders, reviewing and expanding on the previous analyses (Platnick et al. 1991; Ramírez 2000; Griswold et al. 2005). Material and methods Specimen preparation Clean museum specimens were chosen when available. Specimens in 80% alcohol were imaged with a Nikon DXM1200 digital camera mounted on a stereoscopic microscope Nikon SMZ1500 and the focal planes composed with Helicon Focus 3.10.3 and 4.01 Pro (Khmelik et al. 2006). Preparations were carefully cleaned using fine brushes, a thin jet of alcohol from a thinned pipette, or ultrasonic cleaner; some setae were removed to expose structures, especially those on legs and palp, spinnerets and chelicerae. For scanning electron microscope (SEM), all preparations were dehydrated in a series of increasing concentrations of ethanol (80%, 90%, 95%, 100%), and critical-point dried. After drying and brushing, they were mounted on adhesive copper tape (Electron Microscopy Sciences, EMS 77802) affixed to a stub and secured with a conductive paint of colloidal graphite on isopropyl alco- hol base (EMS 12660). Prior to SEM examination under high vacuum with a FEI XL30 TMP, the structures were sputter-coated with Au-Pd. Material examined The following specimens were examined to score the phylogenetic dataset. See Acknowledgements for a list of institutional acronyms and curators. Hypochilus pococki Platnick: 1 ♀, USA, North Carolina, Swain Co., Great Smokey Mountains, Deep Creek Campground, 0.3 Km along Stone Pile Trial, 585 m, N 35°27.848', W 83°26.078', J. Bond, F. Coyle coll., 19.X.2002, MACN-Ar 28173 (ARAMR-000641). 1 ♂, same data, MACN-Ar 29171 (ARAMR-000642). Kukulcania hibernalis (Hentz): 1 ♀, ARGENTINA, Capital Federal, Museo Argentino de Ciencias Naturales “Bernardino Rivadavia”, M. Ramírez, C. Grismado, A. Ojanguren-Afilastro coll., 6.I.2004, MACN-Ar 10427 (ARAMR-000701). 1 ♂, same locality, M. Ramírez col., 16.I.2003, MACN-Ar 10422 (ARAMR-000075). Ariadna boesenbergi Keyserling: 1 ♀, ARGENTINA, Buenos Aires, Ruta Pergamino, Arrecifes Km 210, S. Rodríguez Gil col., XI.2007, MACN-Ar 28170 (ARAMR-000966). 1 ♂, ARGENTINA, Capital Federal, M. Miranda col., I.1989, MACN-Ar 10177 (ARAMR-000965). 2 · Zootaxa 3312 © 2012 Magnolia Press FACUNDO M. LABARQUE Pholcus phalangioides (Fuesslin): 1 ♂, 1 ♀ and 3 immatures, ARGENTINA, Misiones, Santa María, De Carlo, Schiapelli, Viana, Galiano, coll., XI.1954, MACN-Ar 4337. Sicarius rupestris (Holmberg): 2 ♀, ARGENTINA, San Juan, Ruta 150, camino a Paso de Agua Negra, 6 km Oeste de Arrequintín, Maury col., 23.I.1982, MACN-Ar 28169 (ARAMR-000881) and MACN-Ar 28167 (ARAMR-000889), respectively. 1 ♂, same data, MACN-Ar 28168 (ARAMR-000882). Loxosceles rufescens (Dufour): 1 ♀, SPAIN, Granada, Cueva GL-10, Calahonda, M. Ramírez col., 8.XII.1983, UB, Col Aràcnids Reg. Nº 2067 Fl. 83 (ARAMR-000618). 1 ♂, same data (ARAMR-000617). Scytodes globula Nicolet: 2
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