A new species of the spider genus Naevius Roth from Argentina (Araneae, Amaurobiidae, Macrobuninae)

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1 Stud Neotrop Fauna & Environm 35: (2000) /00/ $15.00 Swets & Zeitlinger A new species of the spider genus Naevius Roth from Argentina (Araneae, Amaurobiidae, Macrobuninae) Luis A. Compagnucci 1 and Martín J. Ramírez 1, 2 1 Argentinian Museum of Natural History, Buenos Aires, Argentina 2 Buenos Aires University, Department of Biology, Buenos Aires, Argentina Abstract Naevius calilegua spec. nov. is described from yungas in Jujuy Province, Argentina. Naevius species bear a conspicuous tibial gland, discharging through a movable dorsal tibial apophysis. This character relates Naevius to the macrobunine genera Emmenomma and Anisacate. Keywords: Naevius calilegua spec. nov., Amaurobiidae, Macrobuninae, subtropical Argentina. Correspondence: M. J. Ramírez, Museo Argentino de Ciencias Naturales, Av. Angel Gallardo 470, C 1405 DJR Buenos Aires, Argentina. Fax: (05411) ramirezm@muanbe.gov.ar INTRODUCTION The genus Naevius Roth 1967 currently contains 3 species from the yungas in Peru and Bolivia (Roth, 1967; Brescovit & Bonaldo, 1996). Formerly described as an agelenid, it was subsequently transferred to Amaurobiidae, Desinae by Lehtinen (1967). Forster (1970) elevated the Desinae to family, and since then, Naevius has been listed as Desidae incertae sedis. Forster (1970) included Desidae in the superfamily Dictynoidea, defined by having branched median tracheae. Brescovit & Bonaldo (1996: Fig. 6) found branched median tracheae in N. zongo, and thus agreed with the placement of Naevius in Dictynoidea. However, recent studies of the tracheal system of spiders (Ramírez, 1995, 1996; Griswold et al., 1999) indicate that Dictynoidea is polyphyletic, and that there is extensive homoplasy in the development of median tracheae. The male palps of Naevius species bear bizarre structures that help to resolve their relationship. Differing from most known spiders, the dorsal tibial apophysis is extremely elongated and movable, articulated at its base (Figs. 1, 2). But an even more striking structure was overlooked by previous workers: the globose palpal tibia contains a voluminous gland, discharging, through a long duct, at the tip of the dorsal apophysis (Fig. 11). We observed similar glands in certain genera of Macrobuninae, such as Emmenomma and Anisacate. In Emmenomma, as in Naevius, the tibial apophysis is articulated at its base, while in Anisacate it is fused to the rest of the tibia. Several members of Macrobuninae, such as Macrobunus and Emmenomma, also have branched median tracheae (Ramírez, 1996). In the light of these characters, it seems best to place Naevius in Macrobuninae. Similar orifices and ducts in the male palpal tibial apophysis were reported by Wanless (1979, 1984, 1987) for a few genera of Spartaeinae, which is considered the most basal group of the family Salticidae. Some of them also have movable apophyses, arising from a membranous base, and the duct is connected to a flask shaped vacuole. Wanless suggested that its function might be pheromone secretion or plugging the female copulatory openings, and that a ducted apophysis might be primitive for salticids. MATERIAL AND METHODS Female genitalia and male palps were cleared with clove oil, and the spermathecae, ducts, and tibial glands were observed through a compound microscope. Additional mounts of male palps were observed after digestion with KOH and trypsin solutions. Tracheae were observed after digestion of non-cuticular tissues with KOH 10-20% solution at 100 C in a double boiler. Measurements are given in mm. Spine distribution is described with standard notation, with slight modifications as in Ramírez & Grismado (1997). The specimens are deposited in

2 204 L.A. COMPAGNUCCI AND M.J. RAMÍREZ Figs Naevius calilegua spec. nov., left 么 palpus. Fig. 1. retrolateral view; Fig. 2. base of dorsal tibial apophysis; Fig. 3. apical view; Fig. 4. apex of dorsal apophysis (tip broken).

3 NEW NAEVIUS FROM ARGENTINA 205 Museo Argentino de Ciencias Naturales (MACN, Cristina Scioscia) and Instituto Miguel Lillo (IML, Pablo Goloboff). RESULTS Naevius calilegua spec. nov. (Figs. 1-13) Types holotype and 1 paratype from Argentina, Provincia de Jujuy, Parque Nacional Calilegua, 19.7 km up from park entrance, 1600m, IX.1995, M. Ramírez, P. Goloboff, C. Szumik (MACN 9651). Diagnosis Males are very similar to those of N. manu, but differ by the less globose palpal tibia, the distally wider ventral apophysis, and the basally wider and distally bifid dorsal apophysis. Description Male holotype: Total lenth Carapace length 1.46, width 1.12, height 0.82, posteriorly elevated (Fig. 6), thoracic fovea on posterior slope, very long, slightly sunken. Anterior median eyes smaller than others (Fig. 7). Sternum length 0.76, width Abdomen length 1.66, width Chelicerae with 3 teeth on promargin, 5 on retromargin: apical tooth isolated, smaller, 4 basals contiguous, in crescent series to apex. Palp: Patella lacking apophysis. Tibia globose, ventral apophysis bifid, apex as wide as base. Dorsal apophysis very long, curved, articulated at base, as long as cymbium; base wide, bilobate (Fig. 2), tip bifid (Fig. 4). Tibial gland voluminous, duct passing into dorsal apophysis, discharging through oblique opening (Fig. 11). Cymbium with apical, retrolateral longitudinal groove where apex of dorsal apophysis rests. Embolus arising from prolateral side of tegulum, not articulated. Only one tegular articulate sclerite, probably a conductor, with furrow where embolus fits (Figs. 3, 10). Spines: Femora I-IV: d 0-1-p1; p 0-0-d1; r 0-0d1. Patellae I-IV: d Tibiae I: v p1-2-2 or 2-2-0, p 1-d1-1, r 0-d1-1, d r II: v 0-2-0, p 0-d1-1, r 0-d1-1, d r III: v 0-p1-2, p 0-d1-d1, r 0-d1-1, d r IV: v 0-p1-2, p 0-d1-1, r 0-d1-1, d r Metatarsi: I: v 2-2-1; p d1-d1-1; r d1-d1-1; d 0-p1-2. II: v 2-2-1; p d1-d1-1; r d1-d1-1; d 1-p1-2. III: v 2-2-1; p d1-d1-1; r d1-d1-1; d 1-p1-2. IV: v 2-2-1; p d1-d1-1; r d1-d1-1; d 1- p1-2. Dorsal spines on metatarsus III very large (Fig. 8). Colour: Yellow with greenish gray spots. Carapace with median dark stripe, chelicerae with anterior and lateral spots. Sternum with sinuous dark bands on sides, legs with diffuse irregular dark spots. Abdomen with dark greenish spots on yellow background, white guanine patches (Fig. 5) on dorsum, venter paler, with two lateral lines of white guanine patches and median, diffuse, greenish gray strip. Female paratype: Total length Carapace length 1.58, width 1.26, height Sternum length 0.80, width Abdomen length 1.92, width Epigynum (Fig. 12) slightly sclerotized, median area posteriorly notched, membranous, folded. Lateral lobes posteriorly projected in sclerotized mounds, at sides of posterior atrium. Copulatory ducts long, passing through posterior margin of lateral lobes, copulatory openings hidden inside atrium (Fig. 13). Spines: Femora I-IV, d 0-1-1; p d1a; r d1a. Patellae I-IV, d Tibiae: I, v 2-0-2; p d1-d1-1; r d1-d1; d r1-1. II, v 2-2-0; p 0-d1-1; r d1-d1; d r1-1. III, v 0- p1-2;p d1-1;r d1-d1;d r1-1. IV, v 0-p1-2; p d1-d1; r d1-d1; d r1-1. Metatarsi: I, v 2-2-1; p d1-d1-2; r d1- d1-2; d 0-p1-0. II, v 2-2-1; p d1-d1-2; r d1-d1-2; d III, v 2-2-1; p d1-d1-1; r d1-d1-1; d p1-p1-0. IV, v 2-2-1; p d1-d1-1; r d1-d1-1; d p1-p1-p1. Colour as in female. Tracheal system as in N. zongo (Brescovit & Bonaldo, 1996: Fig. 6). Variability The number of retromarginal cheliceral teeth varies between 5 and 6, even in the same specimen. The size of the two apical teeth are quite homogenous among specimens. Natural history All specimens were collected beating epiphytic mosses and lichens in moderately moist yungas. Etymology The specific name is a noun in apposition, taken from the type locality. Other material examined Same data as types, 10, 1, 28 immatures (MACN), 5 (IML); same, 11.3 km up from Park entrance, 1 (MACN); same, 20.7 km up from Park entrance, 7, 2, 4 immatures (MACN); same, near Monolito, 1700m, 9.VIII.1997, M. Ramírez, L. Compagnucci, 1, 4 immatures (MACN).

4 206 L.A. COMPAGNUCCI AND M.J. RAMÍREZ Figs Naevius calilegua spec. nov. Fig. 5., dorsal; Fig. 6., lateral; Fig. 7., eyes, anterior view; Fig. 8., left metatarsus III, prolateral view; Fig. 9., left palp, ventral view; Fig. 10. the same, retrolateral view; Fig. 11. the same, cleared; Fig. 12. E epigynum, ventral view; Fig. 13. the same, cleared.

5 NEW NAEVIUS FROM ARGENTINA 207 ACKNOWLEDGEMENTS Helpful comments on the manuscript were provided by María Elena Galiano, Valerie Todd Davies, Charles Griswold, Norman Platnick and an anonymous reviewer. Gerardo Lamas and Janine Duarez (Museo de Historia Natural de la Universidad Nacional Mayor de San Marcos, Lima) sent the type of N. manu. The Administración de Parques Nacionales helped during field work. Patricia Sarmiento (Servicio de Microscopía Electrónica, Museo de la Plata) made the scanning micrographs. Lucrecia Nieto and Pablo Goloboff collaborated with the morphological studies of tibial glands. Specimens were reared in laboratory by courtesy of María Elena Galiano and Susana Ledesma (MACN). Support for this project was provided by a graduate fellowship and EXO085 fund from the Universidad de Buenos Aires to MJR. REFERENCES Brescovit AD, Bonaldo AB (1996): Two new species of the genus Naevius Roth (Arachnida, Araneae, Desidae). Spixiana 19: Forster RR (1970): The spiders of New Zealand, Part III: Desidae, Dictynidae, Hahniidae, Amaurobioididae, Nicodamidae. Otago Mus Zool Bull 3: Griswold CE, Coddington JA, Platnick NI, Forster RR (1999): Towards a phylogeny of entelegyne spiders (Araneae, Araneomorphae, Entelegynae). J Arachnol 27: Lehtinen PT (1967): Classification of the cribellate spiders and some allied families, with notes on the evolution of the suborder Araneomorphae. Ann zool fennici 4: Ramírez MJ (1995): A phylogenetic analysis of the subfamilies of Anyphaenidae (Arachnida, Araneae). Ent scandinavica 26: Ramírez MJ (1996): On the genus Callevopsis (Araneae, Dictynoidea). Revta Mus Arg Cien Nat (NS) 144: 1-8. Ramírez MJ, Grismado CJ (1997): A review of the spider family Filistatidae in Argentina (Arachnida, Araneae), with a cladistic reanalysis of filistatid genera. Ent scandinavica 28: Roth VD (1967): A review of the South American spiders of the family Agelenidae (Arachnida, Araneae). Bull Amer Mus nat Hist 134: Wanless FR (1979): A revision of the spider genus Brettus (Araneae: Salticidae). Bull British Mus Nat Hist (Zool) 35: Wanless FR (1984): A review of the spider subfamily Spartaeinae nom. n. (Araneae: Salticidae), with descriptions of six new genera. Bull British Mus Nat Hist (Zool) 46: Wanless FR (1987): Notes on spiders of the family Salticidae, 1: The genera Spartaeus, Mintonia and Taraxella. Bull British Mus Nat Hist (Zool) 52: Received: 2 November 1999 Accepted: 29 May 2000

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