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Accepted by S. Carranza: 13 Feb. 2007; published: 10 May 2007 47 ZOOTAXA ISSN 1175-5326 (print edition) ISSN 1175-5334 (online edition) Copyright © 2007 · Magnolia Press Zootaxa 1470: 4757 (2007) www.mapress.com/ zootaxa/ A new species of Leiosaurus (Iguania: Leiosauridae) from central-western Argentina 1 ALEJANDRO LASPIUR, 2 JUAN CARLOS ACOSTA & 3 CRISTIAN S. ABDALA 1-2 Departamento de Biología e Instituto y Museo de Ciencias Naturales, Facultad de Ciencias Exactas, Físicas y Naturales, Univer- sidad Nacional de San Juan. Av. España 400 (N), CP: 5400, San Juan, Argentina 3 Instituto de Herpetología, Fundación Miguel Lillo – CONICET y Facultad de Ciencias Naturales e IML, UNT. Miguel Lillo 205. Tucumán. Argentina. E-mail: 1 [email protected]; 2 [email protected]; 3 [email protected] 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 Duméril 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 d´Orbigny (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 repre- sented according to Frost et al., (2001) by three species: L. catamarcensis Koslowsky 1898, L. paronae Peracca, 1897 (Cei, 1986) and L. bellii Duméril 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 semi- circles; 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 spe- cies 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

Laspiur Et Al 2007-A New Species of Leiosaurus (Iguania Leiosauridae) From Central-western Argentina-Zootaxa

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Accepted by S. Carranza: 13 Feb. 2007; published: 10 May 2007 47

ZOOTAXAISSN 1175-5326 (print edition)

ISSN 1175-5334 (online edition)Copyright © 2007 · Magnolia Press

Zootaxa 1470: 47–57 (2007) www.mapress.com/zootaxa/

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

1ALEJANDRO LASPIUR, 2JUAN CARLOS ACOSTA & 3CRISTIAN S. ABDALA 1-2Departamento de Biología e Instituto y Museo de Ciencias Naturales, Facultad de Ciencias Exactas, Físicas y Naturales, Univer-sidad Nacional de San Juan. Av. España 400 (N), CP: 5400, San Juan, Argentina3Instituto de Herpetología, Fundación Miguel Lillo – CONICET y Facultad de Ciencias Naturales e IML, UNT. Miguel Lillo 205. Tucumán. Argentina. E-mail: [email protected]; [email protected]; [email protected]

Abstract

In this paper we describe a new species of the Leiosaurus genus from central-western Argentina. This new taxon presentsremarkable 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 markingssimilar to the colour design of some felines like the jaguar. This new species inhabits the highlands of central-westernArgentina where steppe bunch grasses with low plant formation and low spiny shrubs prevail. However, little is knownof its biology as with the other species of the genus Leiosaurus. The discovery of this new taxon is significant, because ithas 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 Duméril and Bibron (1837) to include a new species which wasassigned Mexico as type locality by mistake. The species described was Leiosaurus bellii, widely distributedamong southern Argentina. Thereinafter, new species were described also in Argentina, Leiosaurus darwiniBell, Leiosaurus fasciatus d´Orbigny (Koslowsky, 1898) and Leiosaurus bardensis (Gallardo, 1968). Allthese species were included into the Leiosaurus genus and removed from it due to their synonymy withDiplolaemus according to the proposal of Donoso-Barros (1965). At present the genus Leiosaurus is repre-sented according to Frost et al., (2001) by three species: L. catamarcensis Koslowsky 1898, L. paronaePeracca, 1897 (Cei, 1986) and L. bellii Duméril and Bibron, 1837, all of them distributed in Argentina. Thegenus Leiosaurus is composed by species which have stout body, wide head caused by the presence of largejaw 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 semi-circles; 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 spe-cies included in the group. Nonetheless, Cei (1973) presents a general map with Leiosaurus together toDiplolaemus, Aperopristis and Cupriguanus geographic distribution. The San Juan and La Rioja Provincesrepresent a complex system where high geological formations alternate with intermountained tectonic valleys(Suvires et al., 1999). These geomorphologic regional features represent geographical barriers and cangenerate morphological differences in the species (Irschick et al., 1997; McCranie et al., 2001) considering in

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

Materials and methods

The new taxon was compared to Leiosaurus paronae, L. bellii and L. catamarcensis specimens (syntypesincluded). We also obtained data from the bibliography (Duméril & Bibron, 1837; Koslowsky, 1898). Thequantification and characterization of scales were examined using a binocular microscope (10–40 x) follow-ing Abdala (2002; 2003). The body sizes were obtained using a precision caliper to the nearest 0.01 mm. Thespecimens collected were fixed in 10% formaldehyde and then preserved in 70% ethanol. The collected mate-rial was deposited in the Colección Herpetológica de la Fundación Miguel Lillo (FML) and in the ColecciónHerpetológica del Instituto y Museo de Ciencias Naturales de la Universidad Nacional de San Juan (IMCN-UNSJ). 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° 45´45.9” W) at 2.440 mts., Jáchal

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° 46´W) at

2.762 mts., Coronel Felipe Varela Department. La Rioja Province. Col. M. Jordán and J. Márquez, February

6th, 2005.FML 17585. Adult male. Between Entre Rios School and Villa Mercedes Town, Jáchal 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

Jagüe, 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 americanfeline 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 sub-family (Frost et al., 2001). It clearly differs from all other members of genus due to the dorsal coloration pat-tern, the lepidosis and morphometric characters. The dorsal coloration pattern of L. jaguaris presents dorsalmarkings on the vertebral line in form of circles or semicircles with diffuse borders, absent or faded dorsolat-eral markings, rounded marked tail or with irregular rings and ventral body with or without little markings ordark scales irregularly disposed while L. paronae presents big and defined “lily” shaped spots on the vertebralline, big dorsolateral and circular spots or transverse to the axis body, tail with alternate dark and light ringsand ventrally body with clearly defined markings (Fig. 2). Leiosaurus jaguaris presents dorsal and dorsolat-eral uniform and granular squamation which do not form a protruding crest over the vertebral line or group-ings 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 pro-truding scales placed to both sides and behind the interparietal scale, characteristic in L. paronae. Leiosaurusjaguaris presents gular and ventral rounded scales which are smooth and subjuxtaposed or juxtaposed whileL. paronae has keeled conic gular scales that ending in nib and triangular ventrals, monocarinate and imbri-cate. 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 dis-tributed while L. bellii has a darker abdomen with longitudinal thin markings and a white line on the centralpart of the abdomen; this character is absent in the species hereby described. The rostral scale of theL. jaguaris can not be distinguished form the supralabial ones; temporal region with a line of big scales whichcontinues together with the scales of the subocular region, in contrast to the L. bellii, whose rostral scale iswell differentiated and protruding with a rectangular form and whose temporal scales are more consistent withno big differenced scales. Moreover, L. jaguaris presents subdigital lamellae with mucrones in all the toes offeet and hands, while L. bellii presents subdigital lamellae without mucrones. Leiosaurus jaguaris presents acoloration pattern which is very different from the one of L. catamarcensis (Fig. 4), which has diffuse spots onthe vertebral line in form of circles or semicircles and a lot of circular little spots irregularly distributed andwith different colours to give a tiger “striped” look, while in L. catamarcensis the dorsal spots are bigger andmore defined with no little spots on the back of the body. Besides, L. jaguaris has a continuous ordiscontinuous dark line which follows the supralabial scales up to the auditory meatus and, as was statedbefore, 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, inthe abdominal region, the scales of the L. jaguaris are juxtaposed while in L. catamarcensis are imbricate. InL. jaguaris there are not conic shaped scales protruding from the rest of the vertebral line in contrast to L. cat-amarcensis 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.

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

Description of Holotype. Adult male, snout-vent length (92.7 mm.). Tail length (114.5 mm.) measuredfrom the base of the vent up to the distal end; length of the torso (59.2 mm.) measured from the base of theneck up to the vent; width of the torso (23.9 mm.). Big and distinctive triangular head 1.32 times longer (27.5mm.) 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.). Distancebetween nares (5.8 mm.). Snout length (10.2 mm.) measured from the end of snout to the orbit. Size of thethigh (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.). Tailthin and depressed, 1.9 times longer than torso. Gular fold complete. Rough dorsal surface of the head, cepha-lic irregular and sub-conic scales. Supraorbital semicircles not very differentiated, separated by two rows ofcuspidate scales indistinct from 16–19 marginal scales also cuspidate which skirt the semicircles. These semi-circles are occupied by tiny granular and juxtaposed scales indistinct from 17 irregular and non prominentsuperciliars. 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 butrecognizable. Temporal scales with small granular scales in the centre, surrounded to the anterior area byirregular scales which are strongly cuspidate and limited by an upper furrow of prominent variable in size andform scales. Granular tympanic scales not prominent. Rounded and prominent nasal scale, pentagonal canthalscale separated from the nasal scale by 2 scales. Irregular and small internasal scales. Rostral undistinguish-able from postrostrals, flat prominent pentagonal mental scale, wider than high, in contact with 8 roundedpostmentals, 18 pentagonal supralabials not very variable separated from the nasal scale by four rows of flatsmall scales, 17 pentagonal infralabials. No flattened snout, slightly prominent loreolabial area, irregularsupranasal 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 lateralscales. 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 thenear end of the manus. Granular back antehumeral and antebranchial scales. Prominent, sub-conical fore tibialscales, plain, rounded back tibials. Fourth finger with 20 tricarenate subdigital lamellae and fourth foot fingerwith 34 tricarenate subdigital lamellae. Irregular post auricular fold and antehumeral fold well marked. Infra-carpals and infratarsals imbricate bicarenate or tricarenate lamellae. Monocuspidate teeth.

FIGURE 3. Leiosaurus bellii Adult male, from El Cortaderal, Malargüe Department, Mendoza Province. Photo: CristianS. Abdala.

Coloration in ethanol. Dorsally, head with numerous dark brown spots distributed irregularly and with adrawing with a light gray “whale tail” form surrounded by a black rim. Head with two parallel stripes of darkbrown or black colour laterally. The wider stripe goes from the back of the eye up to the upper front area of theauditory meatus. The other one has the same colour, but it is thinner and broken; it goes over the supralabialscale to the lower front area of the auditory meatus. Ventrally the head has some diffused brown spots. Dorsalbody of light gray colour with numerous spots and irregular dots of dark brown colour distributed all over thebody and limbs in an irregular way. Vertebral stripe not very much distinguishable and three vertebral spotswhich are light gray in the centre and black in the rim with a reinforce form. The back irregular vertebral spotsare not distinguishable. The sides of the body have numerous light brown spots of different sizes which areirregularly distributed. In ventral pattern, it has light gray body with few diffused dots. In a dorsal and ventralpattern, 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 thehead and trunk. Robust body, the length of the trunk is 1.3 times bigger in relation to the body and the snout-vent 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 thansnout-vent length. Dorsal head surface is rough, formed by conical scales. Rostral divided in 2 or 4 scales.Among 3–5 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 ofthe meatus. Scales around the body (both sex overall) 189 – 212, x= 205, 2. Without precloacal pores in malesand females. Very marked gular fold. Dorsal scales, granular, conical, which finish in end, juxtaposed withoutkeels. Ventral scales are bigger than dorsal scales. Hands and feet subdigital tricarenate scales. Laminar infra-carpal 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 andwith a white or light gray “whale tail” drawing surrounded by a black or dark brown rim. This drawing isfound in the other species in the genus. Two parallel dark brown or black stripes are distinguished in the headlaterally. The widest stripe is found from the back part of the eye to the upper front area of the auditorymeatus. The other stripe, same colour but thinner than the first one and broken in various specimens, is foundfrom the supralabials scales to the lower front area of the auditory meatus. Ventrally, the head with numerouslight 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 dotsirregularly distributed all over the body and limbs. With marked vertebral line from the occiput to the upperlimbs, 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 vertebralspots 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. Withoutparavertebral spots. Ventrally white or light gray body with some dots of the same colour as the back. Dorsaltail with the same pattern as the body; in some specimens the vertebral spots form incomplete rings. Ventraltail 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, andnear regions, such as Guandacol River to the north of type locality and to the west of Alto Jagüe, La RiojaProvince.

Natural History and Discussion. Leiosaurus jaguaris lives in a high-altitude area where steppe grazingland with low vegetal coverage and prickly low height bushes predominate (Fig. 7). Among the most commonspecies are: Gochnatia glutinous, Heliotropium sp., Larrea cuneifolia, Plectocarpa tetracantha and P. ruoge-sii. The type specimen was associated to a bush of Larrea cuneifolia, in a stony substratum, other specimensunder Gochnatia glutinous. It seems to be a marked association among the representatives of this kind to thistype of biotopes in accordance with Cei (1986) in the description of the kind of habits of Leiosaurus. There islittle information about the geographic distribution of Leiosaurus and little knowledge of the species includedin 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 afemale 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 (27.8ºC),and get a thin range (7.9ºC) of max and min voluntaries temperatures or “set points”. However, the range ofthermal tolerance is amazing wide (37.87 ºC), exhibiting eurithermy, a condition which relate this specieswith the thermal characteristics of its habitat, a best aptitude to resist low temperature. Probably, the thermal-physiology of L. catamarcensis, which lives on a warm and desert environment, with thermal variation, likethe Argentinean “Monte”, strongly restrict the activity time (in the dawn or in the crepuscule), and the use ofmicrohabitat (caves or tree branches). The thermal characteristics, mimicry and low mobility, are maybe thecause 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: AlejandroLaspiur.

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, thedifferences between these species, there are important similitude. The presence of smooth ventral scales in thepectoral zone is character states which allow us to differentiate L. jaguaris at L. catamarcensis, who exhibitslightly keeled ventral scales (Koslowsky, 1898). In Frost et al., (2001), there is no mention about the “slightlykeeled” state in this character. So, with this work, we contributed to a more exhaustive study of the knowncharacters and adding new morphological characters, present in the taxon described here, for phylogeneticanalysis. A general morphological study could establish the relationships between the new species and theothers 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, José Villavicencio and José Marinero for their collaboration during the development of thedescription and for the capture of one of the individuals. To Pablo Gómez, Pablo Meglioli, Eduardo Sanabriaand Lorena Quiroga for their constant support. To Gustavo Scrocchi for giving L. jaguaris specimens unself-ishly for its description. To Daniel Perez for giving Leiosaurus bellii specimens. To Jorge Williams and Dai-

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ana Ferraro for the loan of L. catamarcensis type material. To Sebastián Barrionuevo and Maria José Tulli fortheir help in the field. To Marcelo Jordán and Justo Márquez for giving generously the paratype specimen. ToCarlos Borghi and Héctor Mendoza for their collaboration.We thank to Salvador Carranza and an anonymousreviewer for their opportune suggestions and comments on the manuscript. To Dirección de Conservación yÁreas Protegidas. To Subsecretaría de Medio Ambiente of San Juan Province and to Dirección General deMedio Ambiente y Desarrollo Sustentable, La Rioja Province for the permission for capture.

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Abdala, C.S. (2003) Cuatro nuevas especies del género Liolaemus (Iguania: Liolaemidae), pertenecientes al grupo bou-lengeri, en la Patagonia, Argentina. Cuadernos de Herpetología, 17 (1–2), 3–32.

Blanco, G.M. & Acosta, J.C. (2003) Leiosaurus catamarcensis (NCN) Clutch Size. Herpetological Review, 34 (2), 145pp.

Cei J.M. (1973) Comentarios Sobre algunos géneros de Iguánidos: Diplolaemus, Leiosaurus, Aperopristis y Cuprigua-nus. Physis, 32 (85), 269–276.

Cei, J.M. (1986). Reptiles del Centro, Centro-Oeste y Sur de la Argentina. Herpetofauna de Zonas Áridas y Semiáridas.Museo Regionale di Scienze Naturali Torino. Monografie IV, 163–168 pp.

Donoso-Barros, R. (1965) El género Diplolaemus Bell en Sudamérica. II Congreso. Latinoamericano de Zoología, SaoPablo, 219–226 pp.

Duméril, A.M.C. & Bibron, G. (1837). Erpetologie générale ou historie naturelle complete des reptiles. Paris: LibraireEncyclopedique de Loret, Vol. IV:ii, 554 pp.

Frost, D.R., Etheridge, R., Janies, D. & Titus, T.A. (2001) Total evidence, sequence alignment, evolution of polychrotidlizards, and a reclassification of the Iguania (Squamata: Iguania). American Museum Novitates, 3343, 1–38.

Gallardo, J.M. (1961) Estudio Zoogeográfico del Género Leiosaurus (Reptilia: Sauria). Physis, 22 (63), 113–118.Gallardo, J.M. (1968) Dos Nuevas Especies de Iguanidae (Sauria) de la Argentina. Neotrópica, 14 (43), 1–8. 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), 2191–2203.Koslowsky, J. (1898) Enumeración Sistemática y distribución geográfica de los reptiles argentinos. Revista del Museo de

La Plata, Tomo VIII, 161 pp.Mc Cranie, J.R., Nicholson, K.E. & Köhler, G. (2001) A new species of Norops (Squamata: Polychrotidae) from north-

western Honduras. Amphibia-Reptilia, 22 (4), 465 – 473. Suvires, G., Pereyra, B., Zambrano, J. & Oviedo, M. (1999) Rasgos geomorfológicos regionales de la provincia de San

Juan. CD Síntesis 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 Duméril and Bibron (Rep-

tilia, Lacertia; Iguanidae). Journal of Herpetology, 11 (2), 238–241 pp.Villavicencio, H.J., Laspiur, A. & Acosta, J.C. (2006) Aportes al Conocimiento de la termofisiología de Leiosaurus cata-

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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 Jagüe, General Lamadrid Department. San Juan:FML 17584. Gualcamayo. Jáchal Department. FML 17585. Between Entre Ríos School and Villa Mercedes town,Jáchal Department.

Leiosaurus catamarcensis—Catamarca: MLP S. 905-906, 2 specimens, (syntypes). FML 63. Famabalasto, Sierra delCajón. FML 670. Tampa-Tampa y Agua Tapada 15 Km. North from Farallón Negro, Belén Department. FML 1221.Valle Carrizalito, between Bajo la Lumbrera y Bajo el Durazno Agua de Dionisio, Belén Department. FML 1319.Campo del Pucará, to southeast from Agua de las Palomas, Andalgalá Department. FML 1707. Km. 110, Cuesta dela Chilca, Belén Department. FML 1849. Campo Pozuelo, Belén Department. FML 1850. San Fernando Locality,Belén Department. FML 3616. Quebrada de los Viscos, Central Camp of the Farallón Negro Mine, DepartamentoBelén. FML 16665. Between El Ingenio y Puesto El Arenal, Andalgalá Department. FML 17086-7. 2 specimensCazadero Zone, Andalgalá Department. FML 17088. Campo El Arenal, Andalgalá Department. La Rioja: FML2066. Campo de Loma Larga, Antinaco, Famatina Department. FML 2693. Between Punta del Agua y Las Chacri-tas, 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, Luján deCuyo. San Juan: IMCN-UNSJ 3003-05. 3 specimens. Baños del Salado, Albardón Department. IMCN-UNSJ 3006Baldecitos Locality, Valle Fértil Department. IMCN-UNSJ 3007 Way to Iglesia, Ullúm Departament. IMCN-UNSJ870 Aguada El Molle Co. Pie de Palo, Caucete Departament.

Leiosaurus paronae— Córdoba: IMCN-UNSJ 3009-3152 Chancaní. IMCN-UNSJ 3010-2716 San Marcos Sierra, Cruzdel Eje Departament. La Pampa: IMCN-UNSJ 3008-2020 Parque Luro, Santa Rosa Departament. Mendoza: FML2039. 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,Jiménez Deparment.

Leiosaurus bellii— Chubut: FML 1600. Paso de los Indios, FML 1934. Puerto Madryn, FML 2688. Punta Pirámide,PenínsulaValdés, FML 2742. Laguna los Indios Establishment, National Route N° 1. FML 9463-64. 2 specimens.Bahía Escondida, Rawson Deparment. Mendoza: FML 2689. 7 Km. To West of Cortadera, Malargüe Deparment.Neuquén: IMCN-UNSJ 3011. El Mangrullo, Picún Leufú Department. FML 2160. Piedra del Aguila. Collón-CuráDeparment. Río Negro: FML 8322. Ing. Jacobacci, 25 de Mayo Deparment. FML 8324. 35 Km. To North West ofChinchinales, Gral. Roca Deparment. Santa Cruz: FML 2127. Punta Maqueda, 35 Km. To South of ComodoroRivadavia, Deseado Deparment.

58 · Zootaxa 1470 © 2007 Magnolia Press