Zootaxa 1470: 4757 (2007) www.mapress.

com / zootaxa/ Copyright 2007 Magnolia Press

ISSN 1175-5326 (print edition)

ZOOTAXA
ISSN 1175-5334 (online edition)

A new species of Leiosaurus (Iguania: Leiosauridae) from central-western Argentina

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ALEJANDRO LASPIUR, 2JUAN CARLOS ACOSTA & 3CRISTIAN S. ABDALA

Departamento de Biologa e Instituto y Museo de Ciencias Naturales, Facultad de Ciencias Exactas, Fsicas y Naturales, Universidad Nacional de San Juan. Av. Espaa 400 (N), CP: 5400, San Juan, Argentina 3 Instituto de Herpetologa, Fundacin Miguel Lillo CONICET y Facultad de Ciencias Naturales e IML, UNT. Miguel Lillo 205. Tucumn. Argentina. E-mail: 1laspiursaurus@gmail.com; 2jcacosta@sinectis.com.ar; 3popper@tucbbs.com.ar

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Abstract
In this paper we describe a new species of the Leiosaurus genus from central-western Argentina. This new taxon presents remarkable differences regarding the lepidosis and coloration pattern compared to the other species of the genus: L. catamarcensis, L. paronae and L. bellii. The dorsal coloration pattern is unique and is characterized by dorsal markings similar to the colour design of some felines like the jaguar. This new species inhabits the highlands of central-western Argentina where steppe bunch grasses with low plant formation and low spiny shrubs prevail. However, little is known of its biology as with the other species of the genus Leiosaurus. The discovery of this new taxon is significant, because it has been one hundred years since the last description of a new species of these taxa. Key words: Leiosauridae, Leiosaurus sp. nov., La Rioja, San Juan, Argentina

Introduction The genus Leiosaurus was described by Dumril and Bibron (1837) to include a new species which was assigned Mexico as type locality by mistake. The species described was Leiosaurus bellii, widely distributed among southern Argentina. Thereinafter, new species were described also in Argentina, Leiosaurus darwini Bell, Leiosaurus fasciatus dOrbigny (Koslowsky, 1898) and Leiosaurus bardensis (Gallardo, 1968). All these species were included into the Leiosaurus genus and removed from it due to their synonymy with Diplolaemus according to the proposal of Donoso-Barros (1965). At present the genus Leiosaurus is represented according to Frost et al., (2001) by three species: L. catamarcensis Koslowsky 1898, L. paronae Peracca, 1897 (Cei, 1986) and L. bellii Dumril and Bibron, 1837, all of them distributed in Argentina. The genus Leiosaurus is composed by species which have stout body, wide head caused by the presence of large jaw muscles. However, the characters unique to the genus are: surface of the subdigital lamellae keeled; smooth tail scales; without caudal autotomy; tail slightly longer than body; no contact between orbital semicircles; dorsal coloration pattern with defined vertebral spots, shark teeth or fleur-de-lis shaped (Cei, 1986; Frost et al., 2001). There is little information about the geographic distribution of Leiosaurus and little knowledge of the species included in the group. Nonetheless, Cei (1973) presents a general map with Leiosaurus together to Diplolaemus, Aperopristis and Cupriguanus geographic distribution. The San Juan and La Rioja Provinces represent a complex system where high geological formations alternate with intermountained tectonic valleys (Suvires et al., 1999). These geomorphologic regional features represent geographical barriers and can generate morphological differences in the species (Irschick et al., 1997; McCranie et al., 2001) considering in
Accepted by S. Carranza: 13 Feb. 2007; published: 10 May 2007

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this case the geographical isolation as the possible cause of speciation. During the period comprehended between 2000 and 2004, in San Juan and La Rioja Provinces some specimens of the genus Leiosaurus were collected in high Pre-Andean foothills areas (between 2.400 and 2.800 mts.) being this a new entity related to the group. In this paper a new species is described based in characteristics regarding the coloration pattern and some morphological variations.

Materials and methods The new taxon was compared to Leiosaurus paronae, L. bellii and L. catamarcensis specimens (syntypes included). We also obtained data from the bibliography (Dumril & Bibron, 1837; Koslowsky, 1898). The quantification and characterization of scales were examined using a binocular microscope (1040 x) following Abdala (2002; 2003). The body sizes were obtained using a precision caliper to the nearest 0.01 mm. The specimens collected were fixed in 10% formaldehyde and then preserved in 70% ethanol. The collected material was deposited in the Coleccin Herpetolgica de la Fundacin Miguel Lillo (FML) and in the Coleccin Herpetolgica del Instituto y Museo de Ciencias Naturales de la Universidad Nacional de San Juan (IMCNUNSJ). The list of the examined material is presented in Appendix 1.

Results Leiosaurus jaguaris sp. nov. Holotype. FML 17584. Adult male. Gualcamayo (29 49 48.5 S; 68 4545.9 W) at 2.440 mts., Jchal Department, San Juan Province. Col. J. Marinero, R. Buff y J. Villavicencio, September 31st, 2000. Paratypes. IMCN-UNSJ 3002. Adult male. Riverbed of the Guandacol River (28 57 S; 68 46W) at 2.762 mts., Coronel Felipe Varela Department. La Rioja Province. Col. M. Jordn and J. Mrquez, February 6th, 2005. FML 17585. Adult male. Between Entre Rios School and Villa Mercedes Town, Jchal Department, San Juan Province. Col. C. Abdala, S. Barrionuevo y M.J. Tulli, November 5th, 2004. FML 7484-95. 12 specimens. Between Punta de Agua and las Chacaritas, 33 km to the West of Alto Jage, General Lamadrid Department, La Rioja Province. Col. S. Torres, S. Kretzschmar, J.C. Moreta and C. Salvatierra, February, 1998. Etymology. It makes reference to the aspect of the coloration pattern which is similar to the american feline Panthera onca. Diagnosis. Leiosaurus jaguaris (Fig. 1) belongs to the Leiosauridae family according to Frost et al., (2001). Together with Diplolaemus and Pristidactylus genera, Leiosaurus belongs to the Leiosaurinae subfamily (Frost et al., 2001). It clearly differs from all other members of genus due to the dorsal coloration pattern, the lepidosis and morphometric characters. The dorsal coloration pattern of L. jaguaris presents dorsal markings on the vertebral line in form of circles or semicircles with diffuse borders, absent or faded dorsolateral markings, rounded marked tail or with irregular rings and ventral body with or without little markings or dark scales irregularly disposed while L. paronae presents big and defined lily shaped spots on the vertebral line, big dorsolateral and circular spots or transverse to the axis body, tail with alternate dark and light rings and ventrally body with clearly defined markings (Fig. 2). Leiosaurus jaguaris presents dorsal and dorsolateral uniform and granular squamation which do not form a protruding crest over the vertebral line or groupings of dorsolateral scales in contrast to L. paronae which presents in its body (mainly over the vertebral line) protruding conical scales which are bigger than the rest and which are keeled forming a vertebral crest which

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is unique among the species of the group. Moreover, L. jaguaris present uniform head scales without two protruding scales placed to both sides and behind the interparietal scale, characteristic in L. paronae. Leiosaurus jaguaris presents gular and ventral rounded scales which are smooth and subjuxtaposed or juxtaposed while L. paronae has keeled conic gular scales that ending in nib and triangular ventrals, monocarinate and imbricate. Leiosaurus jaguaris presents dorsal markings on the vertebral line, but they are never shark tooth shaped like in L. bellii (Fig. 3). Leiosaurus jaguaris has round ventral scales or little markings irregularly distributed while L. bellii has a darker abdomen with longitudinal thin markings and a white line on the central part of the abdomen; this character is absent in the species hereby described. The rostral scale of the L. jaguaris can not be distinguished form the supralabial ones; temporal region with a line of big scales which continues together with the scales of the subocular region, in contrast to the L. bellii, whose rostral scale is well differentiated and protruding with a rectangular form and whose temporal scales are more consistent with no big differenced scales. Moreover, L. jaguaris presents subdigital lamellae with mucrones in all the toes of feet and hands, while L. bellii presents subdigital lamellae without mucrones. Leiosaurus jaguaris presents a coloration pattern which is very different from the one of L. catamarcensis (Fig. 4), which has diffuse spots on the vertebral line in form of circles or semicircles and a lot of circular little spots irregularly distributed and with different colours to give a tiger striped look, while in L. catamarcensis the dorsal spots are bigger and more defined with no little spots on the back of the body. Besides, L. jaguaris has a continuous or discontinuous dark line which follows the supralabial scales up to the auditory meatus and, as was stated before, present smooth ventral scales on the pectoral region, while in L. catamarcensis, in some specimens, presents some dark or orange supralabial scales and the ventral scales are lightly careened. Also ventrally, in the abdominal region, the scales of the L. jaguaris are juxtaposed while in L. catamarcensis are imbricate. In L. jaguaris there are not conic shaped scales protruding from the rest of the vertebral line in contrast to L. catamarcensis which presents some protruding scales, mainly on the front part of the body.

FIGURE 1. Leiosaurus sp. nov. (Paratype, FML 17585). Photo: Cristian S. Abdala. A NEW SPECIES OF LEIOSAURUS Zootaxa 1470 2007 Magnolia Press

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FIGURE 2. Leiosaurus paronae Adult male, from Reserva Bosques Telteca. Mendoza. Photo: Leonardo Muoz.

Description of Holotype. Adult male, snout-vent length (92.7 mm.). Tail length (114.5 mm.) measured from the base of the vent up to the distal end; length of the torso (59.2 mm.) measured from the base of the neck up to the vent; width of the torso (23.9 mm.). Big and distinctive triangular head 1.32 times longer (27.5 mm.) than wide (21 mm.), head height (15.5 mm.), eye diameter (5.7mm.), eye-upper lip distance (3.5 mm.), eye-ear length (11.2 mm.). Very narrow auditory meatus, height of (2.9 mm.), and width (1.4 mm.). Distance between nares (5.8 mm.). Snout length (10.2 mm.) measured from the end of snout to the orbit. Size of the thigh (20.7 mm.), tibia-fibula (18.4 mm.) and foot (23 mm.). Distance between anterior and posterior limbs (39.5 mm). Humerus length (17.1 mm.), forearm (14.4 mm.) and hand (13.8 mm.). Vent width (8.5 mm.). Tail thin and depressed, 1.9 times longer than torso. Gular fold complete. Rough dorsal surface of the head, cephalic irregular and sub-conic scales. Supraorbital semicircles not very differentiated, separated by two rows of cuspidate scales indistinct from 1619 marginal scales also cuspidate which skirt the semicircles. These semicircles are occupied by tiny granular and juxtaposed scales indistinct from 17 irregular and non prominent superciliars. Frontal and front parietal scales not regionalized and subdivided, parietal scales softly cuspidate, always smaller than the interparietal, elongated and surrounded by 9 irregular scales, small pineal organ but recognizable. Temporal scales with small granular scales in the centre, surrounded to the anterior area by irregular scales which are strongly cuspidate and limited by an upper furrow of prominent variable in size and form scales. Granular tympanic scales not prominent. Rounded and prominent nasal scale, pentagonal canthal scale separated from the nasal scale by 2 scales. Irregular and small internasal scales. Rostral undistinguishable from postrostrals, flat prominent pentagonal mental scale, wider than high, in contact with 8 rounded postmentals, 18 pentagonal supralabials not very variable separated from the nasal scale by four rows of flat small scales, 17 pentagonal infralabials. No flattened snout, slightly prominent loreolabial area, irregular supranasal scales undistinguishable from loreolabials. One hundred seventy eight scales around the body. Two

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hundred eight round, smooth and imbricate ventral scales. Tail with cuspidate and imbricate dorsal and lateral scales. Smooth, rounded and imbricate infracauldal scales. Distal end of tail with regularly rectangular scales, strongly imbricate. Sub-rounded antehumeral scales, the same as antebrachial scales which are enlarged in the near end of the manus. Granular back antehumeral and antebranchial scales. Prominent, sub-conical fore tibial scales, plain, rounded back tibials. Fourth finger with 20 tricarenate subdigital lamellae and fourth foot finger with 34 tricarenate subdigital lamellae. Irregular post auricular fold and antehumeral fold well marked. Infracarpals and infratarsals imbricate bicarenate or tricarenate lamellae. Monocuspidate teeth.

FIGURE 3. Leiosaurus bellii Adult male, from El Cortaderal, Malarge Department, Mendoza Province. Photo: Cristian S. Abdala.

Coloration in ethanol. Dorsally, head with numerous dark brown spots distributed irregularly and with a drawing with a light gray whale tail form surrounded by a black rim. Head with two parallel stripes of dark brown or black colour laterally. The wider stripe goes from the back of the eye up to the upper front area of the auditory meatus. The other one has the same colour, but it is thinner and broken; it goes over the supralabial scale to the lower front area of the auditory meatus. Ventrally the head has some diffused brown spots. Dorsal body of light gray colour with numerous spots and irregular dots of dark brown colour distributed all over the body and limbs in an irregular way. Vertebral stripe not very much distinguishable and three vertebral spots which are light gray in the centre and black in the rim with a reinforce form. The back irregular vertebral spots are not distinguishable. The sides of the body have numerous light brown spots of different sizes which are irregularly distributed. In ventral pattern, it has light gray body with few diffused dots. In a dorsal and ventral pattern, tail has same colour and pattern as the body. Variation in paratypes. Based on fourteen adult individuals, seven males and seven females. Morphometry and squamation. The head is longer (23.2 27.8 mm. x= 24.9 mm.) than wider (21.3 25.8 mm. x= 22.3 mm.) (Fig. 5). Height of the head (14.4 18.5 mm. x = 14.7 mm.), neck is narrower than the head and trunk. Robust body, the length of the trunk is 1.3 times bigger in relation to the body and the snoutvent length is 2.8 times bigger in relation to the body. Snout-vent length in males and females jointly: (80.9

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97.5 mm. x= 89.6 mm.). Tail length (88.2 115.9 mm. x= 98, 7 mm.). The tail length is 1.2 times longer than snout-vent length. Dorsal head surface is rough, formed by conical scales. Rostral divided in 2 or 4 scales. Among 35 rows of loreolabials of minor size than the supralabials. Fourteen 22 supralabials. Seventeen 20 infralabials. Distinguishable interparietal of bigger size than parietals and surrounded by 8 10 scales. Six 8 scales in contact with the mental scale. Smaller granular auricular scales in the lower front margin of the meatus. Scales around the body (both sex overall) 189 212, x= 205, 2. Without precloacal pores in males and females. Very marked gular fold. Dorsal scales, granular, conical, which finish in end, juxtaposed without keels. Ventral scales are bigger than dorsal scales. Hands and feet subdigital tricarenate scales. Laminar infracarpal and infratarsal scales, imbricate and triffids.

FIGURE 4. Leiosaurus catamarcensis Adult male, from Baldecitos, San Juan Province. Photo: Cristian S. Abdala.

Coloration. Dorsal head (Fig. 5) with numerous dark brown spots and dots distributed irregularly and with a white or light gray whale tail drawing surrounded by a black or dark brown rim. This drawing is found in the other species in the genus. Two parallel dark brown or black stripes are distinguished in the head laterally. The widest stripe is found from the back part of the eye to the upper front area of the auditory meatus. The other stripe, same colour but thinner than the first one and broken in various specimens, is found from the supralabials scales to the lower front area of the auditory meatus. Ventrally, the head with numerous light and dark brown spots or stripes longitudinally to the body, in most cases with cascade appearance. Light brown or light grey dorsal body with numerous irregular spots or dark brown and light brown dots irregularly distributed all over the body and limbs. With marked vertebral line from the occiput to the upper limbs, then it mixes with big vertebral marks in reniforme form or in complete circle or broken in the middle. Such spots of light grey colour in the center and with various black or dark brown dots in the exterior rim. Five-7 vertebral spots. Some L. jaguaris specimens exhibit variation to this coloration pattern. The vertebral spots can be diffused or absent, or totally distinguishable and prominent. At the sides of the body it can be dis-

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tinguished numerous spots of different size, dark brown or light brown distributed irregularly. Without paravertebral spots. Ventrally white or light gray body with some dots of the same colour as the back. Dorsal tail with the same pattern as the body; in some specimens the vertebral spots form incomplete rings. Ventral tail of the same colour as the abdomen, with some diffuse or immaculate spots.

FIGURE 5. Head of Leiosaurus sp. nov. (Paratype, FML 17585). Photo: Cristian S. Abdala.

Distribution. (Fig.6). L. jaguaris is only known in Gualcamayo (type locality), San Juan Province, and near regions, such as Guandacol River to the north of type locality and to the west of Alto Jage, La Rioja Province. Natural History and Discussion. Leiosaurus jaguaris lives in a high-altitude area where steppe grazing land with low vegetal coverage and prickly low height bushes predominate (Fig. 7). Among the most common species are: Gochnatia glutinous, Heliotropium sp., Larrea cuneifolia, Plectocarpa tetracantha and P. ruogesii. The type specimen was associated to a bush of Larrea cuneifolia, in a stony substratum, other specimens under Gochnatia glutinous. It seems to be a marked association among the representatives of this kind to this type of biotopes in accordance with Cei (1986) in the description of the kind of habits of Leiosaurus. There is little information about the geographic distribution of Leiosaurus and little knowledge of the species included in the group. For Van Devender (1977), L. bellii highly depend, for protection, of his cryptical coloration. Also, a mighty bite allows this species to have a specialized diet based on Tenebrionidae beetles. Cei (1986) confirm this diet for L. catamarcensis and L. paronae. For reproduction, there is data about clutch size of a female of L. catamarcensis (Blanco & Acosta, 2003), which, in captivity, placed 10 eggs. About thermal-

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physiology in L. catamarcensis, Villavicencio et al., (2006), established the selected temperature (27.8C), and get a thin range (7.9C) of max and min voluntaries temperatures or set points. However, the range of thermal tolerance is amazing wide (37.87 C), exhibiting eurithermy, a condition which relate this species with the thermal characteristics of its habitat, a best aptitude to resist low temperature. Probably, the thermalphysiology of L. catamarcensis, which lives on a warm and desert environment, with thermal variation, like the Argentinean Monte, strongly restrict the activity time (in the dawn or in the crepuscule), and the use of microhabitat (caves or tree branches). The thermal characteristics, mimicry and low mobility, are maybe the cause at the low numbers of register and specimens in collections.

FIGURE 6. Distribution of the Leiosaurus species of Argentina based in collections specimen (see Appendix 1).

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FIGURE 7. View of habitat of Leiosaurus sp. nov. Type Locality. San Juan Province, Argentina. Photo: Alejandro Laspiur.

Among the Leiosaurus species, there are variation in both, coloration patterns and lepidosis characters. Being these differences obvious when we differentiate L. jaguaris at L. paronae and L. bellii. However, the differences between these species, there are important similitude. The presence of smooth ventral scales in the pectoral zone is character states which allow us to differentiate L. jaguaris at L. catamarcensis, who exhibit slightly keeled ventral scales (Koslowsky, 1898). In Frost et al., (2001), there is no mention about the slightly keeled state in this character. So, with this work, we contributed to a more exhaustive study of the known characters and adding new morphological characters, present in the taxon described here, for phylogenetic analysis. A general morphological study could establish the relationships between the new species and the others members of the genus. The sexual dichromatism present in the three species of the genus (Frost et al., 2001) is not present in L. jaguaris. In the same work, this author indicate that the adult females of L. bellii; L. paronae y L. catamarcensis are larger than adult males; this condition neither is present in L. jaguaris.

Acknowledgments To Ricardo Buff, Jos Villavicencio and Jos Marinero for their collaboration during the development of the description and for the capture of one of the individuals. To Pablo Gmez, Pablo Meglioli, Eduardo Sanabria and Lorena Quiroga for their constant support. To Gustavo Scrocchi for giving L. jaguaris specimens unselfishly for its description. To Daniel Perez for giving Leiosaurus bellii specimens. To Jorge Williams and DaiA NEW SPECIES OF LEIOSAURUS Zootaxa 1470 2007 Magnolia Press

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ana Ferraro for the loan of L. catamarcensis type material. To Sebastin Barrionuevo and Maria Jos Tulli for their help in the field. To Marcelo Jordn and Justo Mrquez for giving generously the paratype specimen. To Carlos Borghi and Hctor Mendoza for their collaboration.We thank to Salvador Carranza and an anonymous reviewer for their opportune suggestions and comments on the manuscript. To Direccin de Conservacin y reas Protegidas. To Subsecretara de Medio Ambiente of San Juan Province and to Direccin General de Medio Ambiente y Desarrollo Sustentable, La Rioja Province for the permission for capture.

Literature cited
Abdala, C.S. (2002) Nuevo Liolaemus (Iguania: Liolemidae) perteneciente al grupo boulengeri de la provincia de Neuqun, Argentina. Cuadernos de Herpetologa, 16 (1), 313. Abdala, C.S. (2003) Cuatro nuevas especies del gnero Liolaemus (Iguania: Liolaemidae), pertenecientes al grupo boulengeri, en la Patagonia, Argentina. Cuadernos de Herpetologa, 17 (12), 332. Blanco, G.M. & Acosta, J.C. (2003) Leiosaurus catamarcensis (NCN) Clutch Size. Herpetological Review, 34 (2), 145 pp. Cei J.M. (1973) Comentarios Sobre algunos gneros de Igunidos: Diplolaemus, Leiosaurus, Aperopristis y Cupriguanus. Physis, 32 (85), 269276. Cei, J.M. (1986). Reptiles del Centro, Centro-Oeste y Sur de la Argentina. Herpetofauna de Zonas ridas y Semiridas. Museo Regionale di Scienze Naturali Torino. Monografie IV, 163168 pp. Donoso-Barros, R. (1965) El gnero Diplolaemus Bell en Sudamrica. II Congreso. Latinoamericano de Zoologa, Sao Pablo, 219226 pp. Dumril, A.M.C. & Bibron, G. (1837). Erpetologie gnrale ou historie naturelle complete des reptiles. Paris: Libraire Encyclopedique de Loret, Vol. IV:ii, 554 pp. Frost, D.R., Etheridge, R., Janies, D. & Titus, T.A. (2001) Total evidence, sequence alignment, evolution of polychrotid lizards, and a reclassification of the Iguania (Squamata: Iguania). American Museum Novitates, 3343, 138. Gallardo, J.M. (1961) Estudio Zoogeogrfico del Gnero Leiosaurus (Reptilia: Sauria). Physis, 22 (63), 113118. Gallardo, J.M. (1968) Dos Nuevas Especies de Iguanidae (Sauria) de la Argentina. Neotrpica, 14 (43), 18. Irschick D. J., Vitt, L.J., Zani, P.A. & Losos, J.B. (1997) A Comparison of evolutionary radiations in mainland and Caribbean Anolis lizard. Ecology, 78 (7), 21912203. Koslowsky, J. (1898) Enumeracin Sistemtica y distribucin geogrfica de los reptiles argentinos. Revista del Museo de La Plata, Tomo VIII, 161 pp. Mc Cranie, J.R., Nicholson, K.E. & Khler, G. (2001) A new species of Norops (Squamata: Polychrotidae) from northwestern Honduras. Amphibia-Reptilia, 22 (4), 465 473. Suvires, G., Pereyra, B., Zambrano, J. & Oviedo, M. (1999) Rasgos geomorfolgicos regionales de la provincia de San Juan. CD Sntesis del cuaternario de la Provincia de San Juan INGEO, UNSJ, San Juan, Argentina. Van Devender, T.R. (1977) Observations on the Argentine Iguanid Lizard Leiosaurus bellii Dumril and Bibron (Reptilia, Lacertia; Iguanidae). Journal of Herpetology, 11 (2), 238241 pp. Villavicencio, H.J., Laspiur, A. & Acosta, J.C. (2006) Aportes al Conocimiento de la termofisiologa de Leiosaurus cataamarcensis (IGUANIA: LEIOSAURIDAE), en San Juan, Argentina. Actas VII Congreso Argentino de Herpetologa, Corrientes, 15 pp.

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Appendix 1
Leiosaurus jaguaris La Rioja: IMCN-UNSJ 3002. Coronel Felipe Varela Department. FML 7484-95. 12 specimens. Between Punta de Agua and las Chacaritas, 33 Km. West of Alto Jage, General Lamadrid Department. San Juan: FML 17584. Gualcamayo. Jchal Department. FML 17585. Between Entre Ros School and Villa Mercedes town, Jchal Department. Leiosaurus catamarcensisCatamarca: MLP S. 905-906, 2 specimens, (syntypes). FML 63. Famabalasto, Sierra del Cajn. FML 670. Tampa-Tampa y Agua Tapada 15 Km. North from Faralln Negro, Beln Department. FML 1221. Valle Carrizalito, between Bajo la Lumbrera y Bajo el Durazno Agua de Dionisio, Beln Department. FML 1319. Campo del Pucar, to southeast from Agua de las Palomas, Andalgal Department. FML 1707. Km. 110, Cuesta de la Chilca, Beln Department. FML 1849. Campo Pozuelo, Beln Department. FML 1850. San Fernando Locality, Beln Department. FML 3616. Quebrada de los Viscos, Central Camp of the Faralln Negro Mine, Departamento Beln. FML 16665. Between El Ingenio y Puesto El Arenal, Andalgal Department. FML 17086-7. 2 specimens Cazadero Zone, Andalgal Department. FML 17088. Campo El Arenal, Andalgal Department. La Rioja: FML 2066. Campo de Loma Larga, Antinaco, Famatina Department. FML 2693. Between Punta del Agua y Las Chacritas, Gral. Sarmiento Department. FML 9452-5. 6 Km. To East from Anillaco, Castro Barros Deparment. FML 9456. Talampaya National Park, Provincial Route 26, Km. 139. Mendoza: FML 1066. Las Crucecitas Locality, Lujn de Cuyo. San Juan: IMCN-UNSJ 3003-05. 3 specimens. Baos del Salado, Albardn Department. IMCN-UNSJ 3006 Baldecitos Locality, Valle Frtil Department. IMCN-UNSJ 3007 Way to Iglesia, Ullm Departament. IMCN-UNSJ 870 Aguada El Molle Co. Pie de Palo, Caucete Departament. Leiosaurus paronae Crdoba: IMCN-UNSJ 3009-3152 Chancan. IMCN-UNSJ 3010-2716 San Marcos Sierra, Cruz del Eje Departament. La Pampa: IMCN-UNSJ 3008-2020 Parque Luro, Santa Rosa Departament. Mendoza: FML 2039. 25 de Mayo, San Rafael Deparment. Santiago del Estero: FML 35. Puesto Belgrano, Guayasan Deparment. FML 276. Santiago del Estero, Capital. FML 877. Toro Negro. FML 3718. Estancia Toro Negro, Pozo Hondo, Jimnez Deparment. Leiosaurus bellii Chubut: FML 1600. Paso de los Indios, FML 1934. Puerto Madryn, FML 2688. Punta Pirmide, PennsulaValds, FML 2742. Laguna los Indios Establishment, National Route N 1. FML 9463-64. 2 specimens. Baha Escondida, Rawson Deparment. Mendoza: FML 2689. 7 Km. To West of Cortadera, Malarge Deparment. Neuqun: IMCN-UNSJ 3011. El Mangrullo, Picn Leuf Department. FML 2160. Piedra del Aguila. Colln-Cur Deparment. Ro Negro: FML 8322. Ing. Jacobacci, 25 de Mayo Deparment. FML 8324. 35 Km. To North West of Chinchinales, Gral. Roca Deparment. Santa Cruz: FML 2127. Punta Maqueda, 35 Km. To South of Comodoro Rivadavia, Deseado Deparment.

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