Article. The spider genus Segestria Latreille, 1804 in South America (Araneae: Segestriidae)

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1 Zootaxa 3046: (2011) Copyright 2011 Magnolia Press Article ISSN (print edition) ZOOTAXA ISSN (online edition) The spider genus Segestria Latreille, 1804 in South America (Araneae: Segestriidae) ANDRÉ MARSOLA GIROTI & ANTONIO DOMINGOS BRESCOVIT Laboratório de Artrópodes Peçonhentos, Instituto Butantan. Avenida Vital Brasil 1500, São Paulo, São Paulo, Brazil. Abstract The genus Segestria Latreille, 1804 includes 21 species distributed mainly in the Palearctic region. Only two species have been described from South America: Segestria ruficeps Guérin, 1832 (known from Brazil, Uruguay and Argentina) and Segestria pusilla Nicolet, 1849 (known only from Chile). A study of the female genitalia shows that S. ruficeps is a junior synonym of the Paleartcic species Segestria florentina (Rossi 1790), which is redescribed and illustrated. S. pusilla is transferred to the genus Ariadna as a nomen dubium. Key words: taxonomy, Haplogynae, synonymy Introduction The genus Segestria Latreille, 1804 currently includes 21 species distributed mainly in the Palearctic region (Platnick 2011). The species are very similar to those of Ariadna Audouin, 1826 and Gippsicola Hogg, 1900, but can be distinguished from those genera by the three anterior and two posterior triangular cheliceral teeth, the shape of the labium, which is long, narrow, and nearly parallel-sided, with the greatest width near the middle, and the female spination pattern, with tibiae and metatarsi I II with two ventral rows of long and slender spines, and legs IV with many spines (Beatty 1970). Only two species of Segestria have been described from South America, S. pusilla from Chile by Nicolet (1849) and S. ruficeps from Brazil by Guérin (1832); the latter species was later recorded from Uruguay and Argentina (Keyserling 1877; Mello-Leitão 1933, 1944; Capocasale 1990, 1993, 1998; Platnick 2011). Segestria ruficeps is a synanthropic species. In Brazil this species is commonly found in and near houses in urban areas, in crevices or under rocks or stones. The female genitalia of S. ruficeps were never described in detail. Recently we examined a large number of specimens from Brazilian collections and studied the internal genitalia of this species. To our surprise, the genitalia strongly resembled those of the European species Segestria florentina (Rossi 1790) as illustrated by Brignoli (1976) and Kraus (1978). A careful comparison of specimens of S. ruficeps and S. florentina (kindly provided by Norman I. Platnick, American Museum of Natural History) confirmed the morphological resemblance. S. ruficeps is therefore synonymized with S. florentina. Presumably, S. florentina is an old introduction in South America. The type of the second Neotropical species, Segestria pusilla from Chile, is considered lost. However, the original description by Nicolet (1849) suggests that this species was based on a juvenile specimen of Ariadna. Consequently, S. pusilla is here transferred to the genus Ariadna as a nomen dubium. Material and methods The specimens studied are deposited in the Instituto Butantan, São Paulo (IBSP; I. Knysak), the Museu de Ciências Naturais, Fundação Zoobotânica do Rio Grande do Sul, Porto Alegre (MCN; E. H. Buckup) and the American Museum of Natural History, New York (AMNH; N. I. Platnick). The types of Latreille, Walckenaer and Guérin are lost (information provided by the curator of the Muséum National d Histoire Naturelle, MNHN, Paris). Accepted by W. Fannes: 29 Aug. 2011; published: 30 Sep

2 Format of descriptions, measurements and spination follow Grismado (2008). To study the female internal genitalia, the anterior portion of the abdomen was dissected, cleaned and digested in 85% lactic acid, following von Wirth (2006). All measurements are in millimeters. Coloration patterns are described based on specimens preserved in 70% ethanol. Pictures were taken with a Leica DFC 500 digital camera mounted on a MZ 16A. Extended focal range images were composed with Leica Application Suite version SEM images were taken under high vacuum in a JSM 5200 Scanning Electron Microscope in the Museu de Ciencias Naturais, FZB/RS. Geographical coordinates of collection localities are given as longitude and latitude degrees (DMS) in parenthesis. They were obtained with Google Earth (Lat/Lon-WGS84). Abbreviations: d dorsal; p prolateral; v ventral; r retrolateral Taxonomy Family Segestriidae Simon 1893 Genus Segestria Latreille 1804 Segestria florentina (Rossi 1790) Figs 1 23 Aranea florentina Rossi, 1790: 133, pl. 9, fig. 3 (Holotype, probably 1, from Tuscany, Italy, lost). Aranea perfida Walckenaer, 1802: 223 (Holotype, probably 1, from France, lost). Aranea cellaria Latreille, 1804: 217 (Holotype, probably 1, from France, lost). Segestria perfida Walckenaer, 1805: 48, pl. 5, figs (Holotype, probably 1, from France, lost); Hahn, 1820: 9 (Syn.) Segestria cellaria Latreille, 1806: 88 (Holotype, probably 1 and probably collected in France, lost); Hahn, 1820: 9 (Syn.). Segestria florentina: Hahn, 1831: 1, fig. 1; Brignoli 1976: 40, figs Segestria gracilis Lucas, 1838: 24, pl. 6, fig. 1 (Holotype 1 from Canary Islands, lost); Schmidt, 1990: 10 (Syn.). Segestria senoculata Lucas, 1846: 100, pl. 1, fig. 9 (misidentification). Segestria ruficeps Guérin, 1832: 8, pl. 1 (Holotype 1 from unknown locality in the state of Santa Catarina, Brazil, lost); Keyserling 1877: 221; Capocasale 1998: 2, figs NEW SYNONYMY. Segestria holmbergi Mello-Leitão, 1944: 322, figs 2 3 (holotype 1 from La Plata (34 55 S, W), Buenos Aires, Argentina, deposited in Museo de La Plata, La Plata, number 15905, not examined); Capocasale 1998: 2 (Syn.). Material examined. BRAZIL: São Paulo: São Paulo (23 32 S, W) (Bairro Pinheiros), 1, September 1978, E.C. Marceno leg. (IBSP 4010); Itapira (22 25 S, W), 2, 25 February 1975, S. Nougués leg. (IBSP 3009); Rio Grande do Sul: Rio Grande (32 01 S, W) (Campus da Fundação Universidade de Rio Grande), 1, 16 December 1997 (IBSP 15359); Pelotas (31 46 S, W), 1, 14 September 1962, R. Krebs leg. (MCN 1191); URUGUAY: Canelones: Canelones (34 27 S, W) (San Jose de Caruso), 1 1juv., 4 December 1997, A.D. Brescovit leg. (IBSP 14435); Lavalleja: Minas (34 22 S, W) (Complexo Gruta Arequita), 1juv., 5 December 2005, A.D. Brescovit et al. leg. (IBSP 59585); Montevideo: Montevideo (34º S, 56º W) (Parque Rivera), 1 2juv., 15 October 2009, V. Penna leg. (IBSP ); ARGENTINA: Buenos Aires Province: Buenos Aires (34 35 S, W), 1, September 2005, G. Ruiz leg. (IBSP 72053). ENG- LAND: Greater London: London (51 30 N, 0 7 W) Westminster, Dean s Yard, 1, 24 September 1955 (AMNH); FRANCE: Alpes: Cote d Azur, Menton (43 46 N, 7 30 E), 2, July 1934, J. Rossi leg. (AMNH). Diagnosis. Segestria florentina resembles S. senoculata (Linnaeus, 1758) in having a curved embolic tip and female genitalia with an elongated dorsal lobe and expanded ventral lobe (see Brignoli, 1976, figs 18 19), but differs from this species by the shorter embolic tip (Figs 15 18), the folded dorsal lobe and the ventral lobe with a larger glandular region (Figs 19 23). Synonymy. Keyserling (1877) and Brignoli (1976) already noted that S. ruficeps and S. florentina are morphologically very similar. After studying the available (non type) specimens listed above of S. ruficeps and S. florentina, and comparing their genitalia, especially the internal female structures, we conclude that these species are synonyms. Redescription. Male (IBSP 72053): Carapace orange brown, darker laterally and at eye area with darker thoracic striae. Chelicerae reddish brown, with iridescent shine (Fig. 1). Endites and labium yellowish brown. Sternum 60 Zootaxa Magnolia Press GIROTI & BRESCOVIT

3 orange brown, with dark reddish margins. Male palp yellowish brown, with cymbium lighter; legs orange brown, with pairs I and II darker and distal segments lighter (Figs 4 5). Abdomen pale yellowish brown with grey spots and a dorsal pattern composed of 5 6 brown triangular marks. Total length 12.3; carapace, 6.1 long, 4.2 wide. Chelicerae projected, with conspicuous frontal thickened setae (Fig. 2). Chelicerae with three promarginal and two retromarginal triangular teeth and many transversal grooves on the posterior region (Fig. 3). Carapace flattened. Labium twice as long as wide. Endites porrect, constricted in the middle and a posterior conspicuous conical projection. Sternum truncated anteriorly and pointed posteriorly, not projected between coxae IV. Legs with conspicuous ventral scopula on metatarsus and tarsus I IV. Leg formula: II-I-III-IV. Leg measurements: I, femur 6.6, patella 2.25, tibia 6.95, metatarsus 6.7, tarsus 1.6, total 24.1; II, 6.4, 2.25, 7.4, 6.65, 1.55, 24.25; III, 5.35, 2.15, 6.05, 5.9, 1.5, 20.95; IV, 5.55, 2.2, 5.8, 4.9, 1.6, Spination: leg I, femur d , p , v0, r0-0-1/ ; patella d0, p1/2-1, v0, r1; tibia d , p1/0-1/0-1-1/ /0-0-0, v3/ /1-3/2-0-2, r2/ /0-0-0; metatarsus d0, p1/ , v ; r ; II, femur d /1-1-0, p0-0-1, / 0, v0, r0-0-1/0-0/1-1-0/1-1-0; patella d0, p1-0/1, v0, r1; tibia d , p1-0/1-1/ /2-0, v3-1/ /1-1-1, r1/2-1-0/1-1-0/ ; metatarsus d0, p , v , r1-0/1-0-0; metatarsus IV with distal prolateral preening comb with 4 macrosetae. Tarsal organ leg I not found; tarsal organs legs II, III and IV slightly projected, with two very small sensillae (Figs 11 12). Palp with femur twice as long as tibia. Cymbium long, with the distal two-thirds much narrower than the basal third. Bulb inserted at the proximal third of the cymbium, with a slender apex and embolus curved at tip. FIGURES 1 5. Segestria florentina, male (Buenos Aires, Argentina, IBSP 72053). 1 Habitus, dorsal view; 2 Same, lateral view; 3 Chelicerae, ventral view; 4 Right leg I, prolateral view; 5 Same, retrolateral view. Abbreviations: CT = cheliceral teeth; TG = transversal grooves. SEGESTRIA IN SOUTH AMERICA Zootaxa Magnolia Press 61

4 FIGURES Segestria florentina, female (Canelones, Uruguay, IBSP 14435): 6 7 Habitus (6 dorsal view, 7 lateral view); 8 Labium and endites; 9 10 Right leg I, (9 prolateral, 10 retrolateral view). Abbreviations: CP = conical projection; L = labium. Female (IBSP ): Coloration as in male, except carapace and chelicerae darker and abdomen with brown triangular marks dorsally (Fig. 6). Total length 15.8; carapace, 6.8 long, 4.6 wide. Chelicearae without conspicuous frontal thickened setae. Cephalic region more elevated and convex than in the male (Fig. 7). Labium, endites and sternum as in male (Fig. 8). Palpal tarsus, with femur half the tibia length and tarsus with one prolateral macrosetae and one claw without teeth. Legs as in males, differing in size and number of macrosetae (Figs 9 10). Leg formula: I-II-IV-III. Leg measurements: I, femur 5.8, patella 2.5, tibia 5.6, metatarsus 5.1, tarsus 1.3, total 20.3; II, 5.5, 2.5, 5.6, 4.9, 1.4, 19.9; III, 4.55, 2.2, 4.35, 4.1, 1.25, 16.45; IV, 4.9, 2.3, 5.05, 3.85, 1.4, Spination: leg I, femur d , p , v0, r0; patella d0, p0, v0, r0; tibia d0, p1-0-1/2-0-1/0-0, v2/ /1-0-2, r1-0/ /1-0; metatarsus d0, p , v , r ; II, femur d , p , v0, r0; patella d0, p0, v0, r0; tibia d0, p , v , r0; metatarsus d0, p , v , r ; preening comb on metatarsus IV as in male. Tarsal organs as in male (Figs 13 14). Internal genitalia with anterior receptaculum composed of two lobes, with a glandular region between them: the dorsal lobe is very curved, with the distal portion expanded and with a pointed end; the ventral one is ventrally expanded, with muscular insertions. Variation. Males (n = 2): total length Females (n = 6): total length ; carapace ; femur I Distribution. Europe, western Asia, northern Africa (Slawson et al. 2009), the southeast and south of Brazil, Uruguay, and eastern Argentina. 62 Zootaxa Magnolia Press GIROTI & BRESCOVIT

5 FIGURES Segestria florentina, tarsal organs: Male (11 leg III, 12 leg IV); Female (13 leg III, 14 leg IV). Ariadna pusilla (Nicolet 1849), comb. nov. Segestria pusilla Nicolet, 1849: 345, pl. 2, fig. 9 (Holotype 1 juv. from Chile, should be deposited in the MNHN, lost). Remarks. Brignoli (1976) provided comments on S. pusilla, writing that the only available figure [ ] is of a female, more similar to Ariadna than to Segestria. Nicolet (1849) s description and drawing of S. pusilla suggests that he based his new species on a juvenile of Ariadna (the labium resembles that of Ariadna species, being short and wide and narrowed at each end; see Beatty 1970). S. pusilla is here transferred to the genus Ariadna as a nomen dubium. SEGESTRIA IN SOUTH AMERICA Zootaxa Magnolia Press 63

6 FIGURES Segestria florentina (male, Buenos Aires, Argentina, IBSP 72053; female, Canelones, Uruguay, IBSP 14435): Male left palp (15 prolateral view, 16 same, detail, 17 retrolateral view, 18 same, detail); female genitalia (19 lateral view, 20 anterior receptaculum in detail, 21 ventral view, 22 frontal view). Abbreviations: PR = posterior receptaculum; AR = anterior receptaculum; DL = dorsal lobe; GR = glandular region; MI = muscular insertions; IF= interpulmonary fold; UE = uterus externus; VL = ventral lobe. Acknowledgments We would like to thank Cristina Rheims, Cristian J. Grismado and Miguel Arnedo for reviewing the manuscript, Norman I. Platnick for loaning specimens for comparison, and Ricardo Ott for assistance with scanning micrographs. This study was supported by PAP (Programa de Aprimoramento Profissional Instituto Butantan, AMG) and CNPq (no. # / ADB). 64 Zootaxa Magnolia Press GIROTI & BRESCOVIT

7 FIGURE 23. Segestria florentina (female, Canelones, Uruguay, IBSP 14435): Illustration of the genitalia in lateral view. Abbreviations: PR = posterior receptaculum; AR = anterior receptaculum; DL = dorsal lobe; GR = glandular region; MI = muscular insertions; IF= interpulmonary fold; UE = uterus externus; VL = ventral lobe. References Audouin, V. (1826) Explication sommaire des planches d'arachnides de l'egypte et de la Syrie publiées... in "Description de l'egypte...". Histoire Naturelle 1, (arachnids, pp ). Beatty, J.A. (1970) The spider genus Ariadna in the Americas (Araneae, Dysderidae). Bulletin of the Museum of Comparative Zoology 139, Brignoli, P.M. (1976) Ragni d'italia XXIV. Note sulla morfologia dei genitalia interni dei Segestriidae e cenni sulle specie italiane. Fragmenta Entomológica, 12, Capocasale, R.M. (1990) An annotated checklist of uruguayan spiders. Aracnología, 11/12, Capocasale, R.M. (1993) Arañas del Uruguay, VI. Las familias Amaurobiidae, Corinnidae, Clubionidae, Dictynidae, Hahniidae, Mecicobothiidae, Miturgidae, Pisauridae, Segestriidae, Selenopidae y Trechaleidae. Aracnología Suplementos, 7, 1 8 Capocasale, R.M. (1998) Arañas del Uruguay, VIII. Contribución al conocimiento de Segestria ruficeps Guérin (Araneae, Segestriidae). Comunicaciones zoologicas del Museo de Historia Natural de Montevideo, 190, Grismado, C.J. (2008) A taxonomic revision of the spider genus Ariadna Audouin, 1826 in Argentina and Chile, with the description of five new species (Arachnida, Araneae, Segestriidae). Zoosystema, 30, Guérin-Méneville, F.E. (1832) [Description of Segestria ruficeps]. Magazine de Zoologie, de Guérin-Méneville, 2, 8. Hahn, C.W. (1820) Monographie der Spinnen. Nürnberg, Heft 1, pp. 1 16, 4 pls. Hahn, C.W. (1831) Monographie der Spinnen. Nürnberg, Heft 6, 1 p. Hogg, H.R. (1900) A contribution to our knowledge of the spiders of Victoria: including some new species and genera. Proceedings of the Royal Society of Victoria (N.S.) 13, Keyserling, E. (1877) Amerikanische Spinnenarten aus den Familien der Pholcoidae, Scytodoidae und Dysderoidae. Verhandlungen der k.k. Zoologisch-Botanische Gesellschaft in Wien, 27, Kraus, O. (1978) Liphistius and the evolution of spider genitalia. Symposia of the Zoological Society of London, 42, Latreille, P.A. (1804) Histoire naturelle générale et particulière des Crustacés et des Insectes. Paris, 7, Latreille, P.A. (1806) Genera crustaceorum et insectorum. Paris, tome 1, 302 pp. (Araneae, pp ). SEGESTRIA IN SOUTH AMERICA Zootaxa Magnolia Press 65

8 Lucas, H. (1838) Arachnides, Myriapodes et Thysanoures. In Barker-Webb, P. & S. Berthelot (eds.), Histoire naturelle des îles Canaries. Paris, 2, Lucas, H. (1846) Histoire naturelle des animaux articules. In: Exploration scientifique de l'algerie pendant les annees 1840, 1841, 1842 publiee par ordre du Gouvernement et avec le concours d'une commission academique. Paris, Sciences physiques, Zoologie, 1, Mello-Leitão, C.F. (1933) Catálogo das aranhas argentinas. Archivos da Escola Superior de Agricultura e Medicina Veterinaria, 10, Mello-Leitão, C.F. de. (1944) Arañas de la provincia de Buenos Aires. Revista del Museo de La Plata, 3, Nicolet, A.C. (1849) Aracnidos. In: Gay, C. (ed.), Historia física y política de Chile. Zoología, 3, Platnick, N.I. (2011) The world spider catalog, version American Museum of Natural History, online at research.amnh.org/entomology/spiders/catalog/index.html (date of access: 18 July 2011) Rossi, P. (1790) Fauna etrusca: sistens insecta quae in Provinciis Florentina et Pisana praesertim collegit. Liburni, 2, Simon, E. (1893) Histoire naturelle des araignées. Paris, 1, Slawson, C., Askins, M., Spilling, C. (2009) British Spiders. British Arachnological Society, online at: (date of access: 23 November 2010) von Wirth, V. (2006) Präparationstechniken von (Vogel-) Spinnenspermatheken. Arachne, 11, Walckenaer, C.A Faune parisienne. Insectes. ou Histoire abrégée des insectes de environs de Paris. Paris 2, Walckenaer, C.A. (1805) Tableau des aranéides ou caractères essentiels des tribus, genres, familles et races que renferme le genre Aranea de Linné, avec la désignation des espèces comprises dans chacune de ces divisions. Paris, 88 pp. 66 Zootaxa Magnolia Press GIROTI & BRESCOVIT