Stud Neotrop Fauna & Environm 35: 203Ð207 (2000) 0165-0521/00/3503-0203$15.00 © Swets & Zeitlinger

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

Luis A. Compagnucci1 and Martín J. Ramírez1, 2

1Argentinian Museum of Natural History, Buenos Aires, Argentina 2Buenos 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) 4982-4494. E-mail: [email protected]

INTRODUCTION while in Anisacate it is fused to the rest of the tibia. Several members of Macrobuninae, such as Mac- The genus Naevius Roth 1967 currently contains 3 robunus and Emmenomma, also have branched me- species from the yungas in Peru and Bolivia (Roth, dian tracheae (Ramírez, 1996). In the light of these 1967; Brescovit & Bonaldo, 1996). Formerly de- characters, it seems best to place Naevius in Mac- scribed as an agelenid, it was subsequently trans- robuninae. Similar orifices and ducts in the male ferred to Amaurobiidae, Desinae by Lehtinen (1967). palpal tibial apophysis were reported by Wanless Forster (1970) elevated the Desinae to family, and (1979, 1984, 1987) for a few genera of , since then, Naevius has been listed as Desidae incer- which is considered the most basal group of the fam- tae sedis. Forster (1970) included Desidae in the su- ily Salticidae. Some of them also have movable apo- perfamily Dictynoidea, defined by having branched physes, arising from a membranous base, and the median tracheae. Brescovit & Bonaldo (1996: Fig. 6) duct is connected to a flask shaped vacuole. Wanless found branched median tracheae in N. zongo, and suggested that its function might be pheromone se- thus agreed with the placement of Naevius in Dicty- cretion or plugging the female copulatory openings, noidea. However, recent studies of the tracheal sys- and that a ducted apophysis might be primitive for tem of (Ramírez, 1995, 1996; Griswold et al., salticids. 1999) indicate that Dictynoidea is polyphyletic, and that there is extensive homoplasy in the development of median tracheae. MATERIAL AND METHODS The male palps of Naevius species bear bizarre structures that help to resolve their relationship. Dif- Female genitalia and male palps were cleared with fering from most known spiders, the dorsal tibial ap- clove oil, and the spermathecae, ducts, and tibial ophysis is extremely elongated and movable, articu- glands were observed through a compound micro- lated at its base (Figs. 1, 2). But an even more scope. Additional mounts of male palps were ob- striking structure was overlooked by previous work- served after digestion with KOH and trypsin solu- ers: the globose palpal tibia contains a voluminous tions. Tracheae were observed after digestion of gland, discharging, through a long duct, at the tip of non-cuticular tissues with KOH 10-20% solution at the dorsal apophysis (Fig. 11). We observed similar 100°C in a double boiler. Measurements are given in glands in certain genera of Macrobuninae, such as mm. Spine distribution is described with standard Emmenomma and Anisacate. In Emmenomma, as in notation, with slight modifications as in Ramírez & Naevius, the tibial apophysis is articulated at its base, Grismado (1997). The specimens are deposited in 204 L.A. COMPAGNUCCI AND M.J. RAMêREZ

Figs. 1-4. Naevius calilegua spec. nov., left ( palpus. Fig. 1. retrolateral view; Fig. 2. base of dorsal tibial apophysis; Fig. 3. apical view; Fig. 4. apex of dorsal apophysis (tip broken). NEW NAEVIUS FROM ARGENTINA 205

Museo Argentino de Ciencias Naturales (MACN, p1-2. Dorsal spines on metatarsus III very large (Fig. Cristina Scioscia) and Instituto Miguel Lillo (IML, 8). Colour: Yellow with greenish gray spots. Cara- Pablo Goloboff). pace with median dark stripe, chelicerae with anteri- or and lateral spots. Sternum with sinuous dark bands on sides, legs with diffuse irregular dark spots. Ab- RESULTS domen with dark greenish spots on yellow back- Naevius calilegua spec. nov. ground, white guanine patches (Fig. 5) on dorsum, venter paler, with two lateral lines of white guanine (Figs. 1-13) patches and median, diffuse, greenish gray strip. Types Female paratype: Total length 3.48. Carapace ( holotype and 1 & paratype from Argentina, Pro- length 1.58, width 1.26, height 0.72. Sternum length vincia de Jujuy, Parque Nacional Calilegua, 19.7 km 0.80, width 0.76. Abdomen length 1.92, width 1.36. up from park entrance, 1600m, 23-24.IX.1995, M. Epigynum (Fig. 12) slightly sclerotized, median area Ramírez, P. Goloboff, C. Szumik (MACN 9651). posteriorly notched, membranous, folded. Lateral lobes posteriorly projected in sclerotized mounds, at Diagnosis sides of posterior atrium. Copulatory ducts long, Males are very similar to those of N. manu, but differ passing through posterior margin of lateral lobes, by the less globose palpal tibia, the distally wider copulatory openings hidden inside atrium (Fig. 13). ventral apophysis, and the basally wider and distally Spines: Femora I-IV, d 0-1-1; p d1a; r d1a. Patellae bifid dorsal apophysis. I-IV, d1-0-1. 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- Description p1-2;p d1-1;r d1-d1;d r1-1. IV, v 0-p1-2; p d1-d1; r Male holotype: Total lenth 3.52. Carapace length d1-d1; d r1-1. Metatarsi: I, v 2-2-1; p d1-d1-2; r d1- 1.46, width 1.12, height 0.82, posteriorly elevated d1-2; d 0-p1-0. II, v 2-2-1; p d1-d1-2; r d1-d1-2; d 1- (Fig. 6), thoracic fovea on posterior slope, very long, 1-0. III, v 2-2-1; p d1-d1-1; r d1-d1-1; d p1-p1-0. IV, slightly sunken. Anterior median eyes smaller than v 2-2-1; p d1-d1-1; r d1-d1-1; d p1-p1-p1. Colour as others (Fig. 7). Sternum length 0.76, width 0.70. Ab- in female. Tracheal system as in N. zongo (Brescovit domen length 1.66, width 1.20. Chelicerae with 3 & Bonaldo, 1996: Fig. 6). teeth on promargin, 5 on retromargin: apical tooth isolated, smaller, 4 basals contiguous, in crescent se- Variability ries to apex. Palp: Patella lacking apophysis. Tibia The number of retromarginal cheliceral teeth varies globose, ventral apophysis bifid, apex as wide as between 5 and 6, even in the same specimen. The base. Dorsal apophysis very long, curved, articulated size of the two apical teeth are quite homogenous at base, as long as cymbium; base wide, bilobate (Fig. among specimens. 2), tip bifid (Fig. 4). Tibial gland voluminous, duct passing into dorsal apophysis, discharging through Natural history oblique opening (Fig. 11). Cymbium with apical, ret- All specimens were collected beating epiphytic moss- rolateral longitudinal groove where apex of dorsal es and lichens in moderately moist yungas. apophysis rests. Embolus arising from prolateral side of tegulum, not articulated. Only one tegular articu- Etymology late sclerite, probably a conductor, with furrow where The specific name is a noun in apposition, taken from embolus fits (Figs. 3, 10). Spines: Femora I-IV: d 0- the type locality. 1-p1; p 0-0-d1; r 0-0d1. Patellae I-IV: d 1-0-1. Tib- iae I: v p1-2-2 or 2-2-0, p 1-d1-1, r 0-d1-1, d r1-0-1- Other material examined 0. II: v 0-2-0, p 0-d1-1, r 0-d1-1, d r1-0-1-0. III: v Same data as types, 10 ((, 1 &&, 28 immatures 0-p1-2, p 0-d1-d1, r 0-d1-1, d r1-0-1-0. IV: v 0-p1-2, (MACN), 5 (( (IML); same, 11.3 km up from Park p 0-d1-1, r 0-d1-1, d r1-0-1-0. Metatarsi: I: v 2-2-1; entrance, 1 ( (MACN); same, 20.7 km up from Park p d1-d1-1; r d1-d1-1; d 0-p1-2. II: v 2-2-1; p d1-d1-1; entrance, 7 ((, 2 &&, 4 immatures (MACN); same, r d1-d1-1; d 1-p1-2. III: v 2-2-1; p d1-d1-1; r d1-d1- near Monolito, 1700m, 9.VIII.1997, M. Ramírez, L. 1; d 1-p1-2. IV: v 2-2-1; p d1-d1-1; r d1-d1-1; d 1- Compagnucci, 1 (, 4 immatures (MACN). 206 L.A. COMPAGNUCCI AND M.J. RAMêREZ

Figs. 5-13. 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. NEW NAEVIUS FROM ARGENTINA 207

ACKNOWLEDGEMENTS Lehtinen PT (1967): Classification of the cribellate spiders and some allied families, with notes on the evolution of Helpful comments on the manuscript were provided by the suborder . Ann zool fennici 4: 199- María Elena Galiano, Valerie Todd Davies, Charles Gris- 468. wold, Norman Platnick and an anonymous reviewer. Ger- Ramírez MJ (1995): A phylogenetic analysis of the sub- ardo Lamas and Janine Duarez (Museo de Historia Natural families of Anyphaenidae (Arachnida, Araneae). Ent de la Universidad Nacional Mayor de San Marcos, Lima) scandinavica 26: 361-384. sent the type of N. manu. The Administración de Parques Ramírez MJ (1996): On the genus Callevopsis (Araneae, Nacionales helped during field work. Patricia Sarmiento Dictynoidea). Revta Mus Arg Cien Nat (NS) 144: 1-8. (Servicio de Microscopía Electrónica, Museo de la Plata) Ramírez MJ, Grismado CJ (1997): A review of the spider made the scanning micrographs. Lucrecia Nieto and Pablo family Filistatidae in Argentina (Arachnida, Araneae), Goloboff collaborated with the morphological studies of with a cladistic reanalysis of filistatid genera. Ent scan- tibial glands. Specimens were reared in laboratory by cour- dinavica 28: 319-349. tesy of María Elena Galiano and Susana Ledesma Roth VD (1967): A review of the South American spiders (MACN). Support for this project was provided by a grad- of the family Agelenidae (Arachnida, Araneae). Bull uate fellowship and EXO085 fund from the Universidad de Amer Mus nat Hist 134: 299-345. Buenos Aires to MJR. Wanless FR (1979): A revision of the spider genus Brettus (Araneae: Salticidae). Bull British Mus Nat Hist (Zool) 35: 127-200. Wanless FR (1984): A review of the spider subfamily Spar- REFERENCES taeinae nom. n. (Araneae: Salticidae), with descriptions of six new genera. Bull British Mus Nat Hist (Zool) 46: Brescovit AD, Bonaldo AB (1996): Two new species of 135-205. the genus Naevius Roth (Arachnida, Araneae, Desidae). Wanless FR (1987): Notes on spiders of the family Saltici- Spixiana 19: 233-238. dae, 1: The genera Spartaeus, Mintonia and Taraxella. Forster RR (1970): The spiders of New Zealand, Part III: Bull British Mus Nat Hist (Zool) 52: 107-137. Desidae, Dictynidae, Hahniidae, Amaurobioididae, Ni- codamidae. Otago Mus Zool Bull 3: 1-184. Griswold CE, Coddington JA, Platnick NI, Forster RR (1999): Towards a phylogeny of entelegyne spiders Received: 2 November 1999 (Araneae, Araneomorphae, Entelegynae). J Arachnol Accepted: 29 May 2000 27: 53-63.