Zootaxa, Redescription of Liolaemus dorbignyi ...

Zootaxa 1717: 51–67 (2008) www.mapress.com / zootaxa/

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Redescription of Liolaemus dorbignyi Koslowsky, 1898 and description of a new species of Liolaemus (Iguania: Liolaemidae) ANDRÉS SEBASTIÁN QUINTEROS 1, CRISTIAN SIMÓN ABDALA 2 & FERNANDO JOSÉ LOBO 3 1, 3

Cátedra de Anatomía Comparada. Facultad de Ciencias Naturales. Universidad Nacional de Salta. Av. Bolivia 5150. 4400. Salta. Argentina. 54-387-4255591. E-mail: [email protected]; [email protected] 2 Instituto de herpetología, Funcación Miguel Lillo y Facultad de Ciencias Naturales e IML. Miguel Lillo 251. 4000. Tucumán. Argentina. 54-381-4230056. E-mail: [email protected].

Abstract Liolaemus dorbignyi is re-described, and its type locality restricted to the Sierra de Fiambalá, Catamarca Province, Argentina. Also, Liolaemus scrocchii sp. nov., a new species of Liolaemus belonging to the L. montanus series, previously confused with L. dorbignyi , is described. This new species inhabits the phytogeographic region of Puna in Argentina at 3500 m. and is distributed in Jujuy and northwestern Salta provinces. Diagnostic characters include the color pattern, (composed of transverse paravertebral spots, that are band shaped and never reach the lateral region, and never contact the vertebral zone), and the number of scales around midbody. Several populations from different localities were considered L. dorbignyi because of their color pattern and squamation. Even though part of this confusion is clarified in this contribution, some populations still require further study. Key words: Lizards, Liolaemus, montanus group, Koslowsky, Northwestern Argentina, Puna highlands

Resumen Se re-describe a Liolaemus dorbignyi, y su localidad tipo es restringida a la Sierra de Fiambal, en la Provincia de Catamarca, Argentina. También se describe una nueva especie de Liolaemus, Liolaemus scrocchii sp. nov., previamente confundida con L. dorbignyi, perteneciente a la serie de L. montanus. Esta especie nueva habita en la región fitogeográfica de la puna en Argentina, a 3500 m. Esta distribuida en la provincia de Jujuy y en el noroeste de la provincia de Salta. Sus caracteres diagnsticos incluyen el patrón de coloración, (compuesto por manchas paravertebrales transversales, con forma de bandas y que nunca llegan a los laterales del cuerpo ni contactan la zona vertebral), y el número de escamas alrededor del cuerpo. Diversas poblaciones de diferentes localidades fueron consideradas bajo el nombre de L. dorbignyi, debido a su patrón de coloración y escamación. Aunque parte de esta confusión es resuelta en este trabajo, algunas poblaciones aún requieren estudio.

Introduction The genus Liolaemus is composed of more than 200 species distributed from Tierra del Fuego, in southern Argentina to Peru, occupying diverse regions in Argentina, Bolivia, Chile, Paraguay and the coasts of Brazil and Uruguay. In the last years many works brought knowledge about the taxonomy and phylogenetic relationships within the genus. Laurent (1983) divided the genus in two major groups, Liolaemus sensu stricto, or the Chilean group (chiliensis), and Eulaemus, or the Argentinean group. Halloy et al. (1998) presented a phylogenetic analysis of the L. wiegmannii group using behavioral characters; Etheridge (2000) analyzed the same group based on morphological characters; Lobo (2001, 2005) studied the relationships within the chiliensis Accepted by S. Carranza: 18 Jan. 2008; published: 3 Mar. 2008

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group; Schulte et al. (2000) studied the phylogenetic relationships of 59 species in the genus, based on mitochondrial DNA, and determined sections within the two major groups of Laurent (1983). Espinoza et al., (2004) studied the relationships of the genus based on 12s, NADH dehydrogenase subunit I and NADH dehydrogenase subunit II sequences. In these two works the major groups of Laurent (1983) were recovered. Cruz et al. performed a molecular-based phylogenetic analysis to study the Bergmann’s and Rapoport’s rules. Avila et al. (2006) studied the relationships of the species of the Liolaemus boulengeri group. Also, Abdala (2007) perform a phylogeny of the L. boulengeri group based on both, molecular and morphological characters. Given the similarity among species and the lack of study of some groups, several problems appear in the identification of diverse species. Etheridge (1993) revised the L. darwinii complex, describing four new species previously confused with L. darwinii Bell. Abdala (2005) redescribed L. boulengeri Koslowsky, and described two new species, and suggested that at least four additional species may be confused with L. boulengeri. Koslowsky (1898) described Liolaemus dorbignyi in detail, without designating a holotype. The type locality was imprecisely stated as “Catamarca Province.” In subsequent publications, many populations were identified as L. dorbignyi, extending its distribution to the Puna of Chile (Núñez, 1992) and Bolivia (Laurent, 1995). Many Liolaemus species from northwestern Argentina have been described in recent years (Lobo and Espinoza, 1999, 2004; Martinez Oliver and Lobo, 2002; Abdala, 2006, and Abdala and Lobo, 2006); however, most of them are species belonging to the L. darwinii group or L. alticolor group. The study of specimens from several populations from Catamarca, Salta and Jujuy identified in collections as L. dorbignyi indicated clear differences in color pattern and squamation characteristics. This situation indicated the need to clearly establish the origin of the material used to describe L. dorbignyi by Koslowsky. Goals of this study include the redescription of L. dorbignyi, including providing data about its distribution and biology, and the description of a new species that has been confused with L. dorbignyi in the past.

Materials and methods Morphological characters studied were those commonly used for Liolaemus taxonomy, described by Laurent (1985), Cei (1986), Etheridge (1993, 1995, 2000), Lobo (2001), and Abdala (2002, 2003, 2005, 2007). Terminology for the description of squamation is that of Smith (1946), and neck-fold terminology follows Frost (1992). The description of colors in life is based on observations made in the field and /or pictures taken at the moment of capture. Terminology for body color patterns follows Lobo and Espinoza (1999). Scale observation and measurements were taken using digital a caliper to the nearest 0.02 mm, under a stereoscope (10– 40x). All measurements are in mm. Studied specimens are deposited in FML (Fundación Miguel Lillo, Tucumán), MCN (Museo de Ciencas Naturales, Salta), and MLP.S (Museo de la Plata).

Results Liolaemus dorbignyi Koslowsky 1898 (Fig. 1–4) Designation of a lectotype— (Fig. 1) Among the specimens of Liolaemus dorbignyi collected by Koslowsky (in his original description there is no mention of the number of specimens collected) which he used to write his description of this species, five adult specimens (three males and two females) are deposited in the Museo de La Plata (MLP). Koslowsky (1898) included a plate with three photographs of adult specimens (his Lam.

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VIII). Both ventral and dorsal pictures corresponded to two male specimens, but none seem to correspond to the preserved males extant in the MLP collection. There is a picture of a lateral view of a head, but this picture does not allow us to compare with the specimens preserved. The description also does not provide enough information to allow a determination of which specimens were used for the type description. Ferraro and Williams (2006) studied the type material of amphibians and reptiles deposited in the Museo de La Plata, which includes the species described by Koslowsky. In relation to L. dorbignyi, they identified a syntype (MLP.S 844), and argued that it corresponded to the ventral picture of a specimen in Koslowsky’s plate. According to Art 73.2 of the ICZN, all specimens belonging to the type series are automatically considered syntypes, when no holotype or lectotype was designed, so the five specimens of the Koslowsky’s series are syntypes. We don’t agree with Ferraro and Williams (2006), and we consider that none of the specimens of Koslowsky’s series match the specimens illustrated, and therefore we designate as lectotype the individual catalogued as MLP.S 057 (in compliance with art 74, of ICZN), a specimen which better represent the species. Also, the specimens catalogued as MLP.S 057; 844–45; 990–91 are designated paralectotypes.

FIGURE 1. Dorsal view of the Lectotype of L. dorbignyi (top) and a specimen collected in Sierra de Fiambalá. The arrows indicate the similar bands and spot that the two specimens exhibit.

Designtion of the type locality—Since Koslowsky (1898) described the type locality of Liolaemus dorbignyi as “Catamarca province,” the identity of this species has been problematic. Unfortunately, Koslowsky’s field notes, which would be very helpful, are lost (Jorge Williams pers. comm.). During the revision of materials deposited at the FML (Fundación Miguel Lillo, Tucumán) and MCN (Museo de Ciencias Naturales, Salta), we found specimens corresponding to different populations assigned to L. dorbignyi. Among these samples, only one individual (FML 1855) is similar to those collected by Koslowsky. However, these specimens had an incorrect label about their provenance (Saujil, National Road 60, 1550 m. Department Tinogasta, Catamarca Province). This incorrect label was confirmed during field trips, and O. Pagaburo (collector of FML 1855), told us that the specimens were collected in the Sierra de Fiambalá, over 3000 m. Later, from those mountains we collected specimens, which are similar to those collected by Pagaburo and Koslowsky.

REDESCRIPTION OF L. DORBIGNYI AND A NEW SPECIES

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Koslowsky (1898), in the original description of L. dorbignyi, designated as type locality “Catamarca Province.” To determine where Koslowsky could have collected the specimens, preserved specimens were compared to specimens of different populations collected in several localities of Catamarca province, Argentina. Also we analyzed the pictures from Koslowsky’s publication; because of the five preserved specimens only two retain the original color pattern, while the others have this pattern obscure and inconspicuous owing to their preservation conditions. From the populations studied, those from Sierra de Fiambalá, Catamarca Province exhibit the same color pattern (with dark bands transverse to the body axis), flat and keeled dorsal scales, and better match with the preserved specimens, the pictures and the original description. There are no known records from Koslowsky’s journeys in Catamarca. However, the comparisons between the different populations of Catamarca with the preserved material (Fig. 1), allows us to restrict the type locality of L. dorbignyi to Sierra de Fiambalá, Catamarca province. Diagnosis—Liolaemus dorbignyi belongs the L. montanus group (Etheridge, 1995), because it has a blade-like process on posterior distal tibia, associated with a greatly hypertrophied M. tibialis anticus (Abdala, et al. 2006). Within this group, the presence of scales of equal size on the posterior face of the femur differentiates it from the L. boulengeri group (Abdala, 2007). Among the species that have not a patch of enlarged scales on the posterior face of the femur, L. dorbignyi is a robust and large sized Liolaemus (max SVL 98.3 mm) which distinguishes it from L. andinus Koslowsky, L. audituvelatus Núñez and Yáñez, L. disjunctus Laurent, L. eleodori Cei et al., L. erroneus Núñez and Yáñez, L. etheridgei Laurent, L. fabiani Yáñez and Núñez, L. famatinae Cei, L. fittkaui Laurent, L. griseus Laurent, L. huacahuasicus Laurent, L. insolitus Cei and Péfaur, L. islugensis Ortiz and Marquet, L. molinai Valladares et al., L. montanus Koslowsky, L. multicolor Koslowsky, L. ortizi Laurent, L. pantherinus Pellegrin, L. poconchilensis Valladares, L. poecilochromus Laurent, L. pulcherrimus Laurent, L. reichei Werner, L. rosenmanni Núñez and Navarro, L. ruibali DonosoBarros, L. signifier Dumerill and Bibrón, and L. vallecurensis Pereyra, (max SVL 50–70 mm). Liolaemus dorbignyi has flat dorsal scales, imbricate, and slightly keeled, character states that distinguish it from L. andinus, L. audituvelatus, L. erguetae Laurent, L. erroneus Núñez and Yáñez, L. forsteri Laurent, L. insolitus, L. islugensis, L. molinai, L. patriciaiturrae Navarro and Núñez, L. poecilochromus, L. robertoi Pincheira-Donoso and Núñez, L. rosenmanni, and L. schmidti Marx (smooth and juxtaposed dorsal scales). It may also be distinguished from L. aymararum Veloso et al., L. disjunctus, L. etheridgei, L. fittkaui, L. huacahuasicus, L. montanus, L. ortizi, L. polystictus Laurent, L. stolzmanni Steindachner, L. thomasi Laurent, and L. williamsi Laurent, which have dorsal scales imbricate but strongly keeled. Liolaemus chlorostictus Laurent, L. jamesi Boulenger, L. orientalis Müller, L. poconchilensis, L. robustus Laurent, L. signifier, and L. stolzmanni, have weakly keeled but juxtaposed dorsal scales, characteristic that distinguishes it from L. dorbignyi. The number of scales around midbody in L. dorbignyi is 48 to 59, fewer than in L. andinus, L. disjunctus, L. duellmani Cei, L. eleodori, L. erguetae, L. foxi Núñez et al., L. islugensis, L. molinai, L. multicolor, L. nigriceps Philippi, L. patriciaiturrae, L. pleopholis Laurent, L. poecilochromus, L. pulcherrimus, L. robertoi, L. rosenmanni, L. ruibali, L. signifer, and L. vallecurensis, which have 64 to 110. Females of L. dorbignyi have precloacal pores, differing from following species in which females lack precloacal pores, L. andinus (Koslowsky, 1895), L. duellmani (Cei, 1986), L. fabiani (Pincheira-Donoso and Núñez, 2005), L. islugensis (Ortiz & Marquet, 1987), L. jamesi (Pincheira-Donoso and Núñez, 2005), L. melanogaster (Lurent, 1998), L. pantherinus ((Pincheira-Donoso and Núñez, 2005), L. puritamensis Núñez and Fox, L. robertoi, L. rosenmanni, L. signifer and L. stolzmanni (Pincheira-Donoso & Núñez, 2005), and L vallecurensis. The presence of precloacal pores in females could be problematic for the diagnosis, because some species are known for few specimens, but we used this character because the new species show a high percentage of females with precloacal pores and in all the samples (from different localities), at least one female exhibit pores. The dorsal color pattern is formed by dark bands transverse to the longitudinal body axis, and which searate the laterals and the ventral color, distinguishing it from the others species of the L. montanus group.


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Description of the Lectotype — MLP.S 057—Male, SVL 95.0. Trunk length 45.8. Head length 20.7. Head width 20.3. Head height (at parietal) 15.2. Eye diameter 3.2. Interorbital distance 9.7. Orbit-auditory meatus distance 8.9. Auditory meatus height 3.7, width 1.9. Orbit-commissure of mouth distance 2.7, distance between nares 3.2. Subocular scale 4.9. Femur 14.2, tibia 17.8, foot 25.0 and humerus 9.1.

FIGURE 2. Dorsal view of a male specimen of L. dorbignyi from the Sierra de Fiambalá, Catamarca, Argentina. Photo: C. S. Abdala.

FIGURE 3. Ventral view of a male specimen of L. dorbignyi from the Sierra de Fiambalá, Catamarca, Argentina. Photo: C. S. Abdala.



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Dorsal surface of the head smooth, with sixteen scales (From occiput to rostral sclae). Rostral scale wider than higher, bordered by six scales. Mental scale larger than rostral, trapezoidal, bordered by four scales. Rostral scale not in contact with nasal. Two internasals. Nasal scale surround by seven scales, separated from canthal by two scales. Five scales between rostral and supercilliaries. Six scales between rostral and frontal. Frontal horizontally divided. Two postrostrals. Interparietal smaller than or equal to parietals, surrounded by six scales. Twelve upper ciliaries. Supraorbital semicircles complete. Five supraoculars. Preocular scale separated from lorilabials row by one scale. Two projecting scales on anterior and superior margin of auditory meatus. Eight smooth horizontal temporals and six verticals. Nine lorilabials, five in contact with subocular scale. Seven supralabials, fourth larger than the others, without sense organs, not in contact with subocular scale. Four infralabials, second ventrally in contact with two scales. Four chinshields; second imbricate separated by two scales. Forty eight scales around midbody. Forty nine dorsal scales, from occiput to forelimbs, with rounded anterior margins, and slightly keeled. Twenty seven rows of scales on dorsum. Ventrals larger than dorsals, flat, juxtaposed in abdominal region, and subjuxtaposed on chest and laterals regions. Thirty five subjuxtaposed and smooth gulars, between auditory meatus. Seven precloacal pores. Antehumeral scales flat and larger than dorsals. Postauricular, rictal and longitudinal nuchal folds present. Scales on the longitudinal fold smooth. Seventeen subdigital lamellae on IV finger, and twenty four on toe IV. Third and fourth left toes absent. Flanks scales triangulars, smooth with several small granular intercalated scales. Femoral patch absent. Infracarpals scales flat, imbricate and trifid. Infratarsals scales flat, imbricate and keeled. Color in ethanol—Head gray dorsally, with dark brown tonalities, darker than the body. Trunk, lead-gray with seven to nine dark transverse bands, faded in vertebral and paravertebral regions. Fore and hind limbs with same pattern and color as trunk. Vertebral line absent, scapular spots and dorsolateral stripes absent. Lateral region dark gray, like the dorsum, showing projections of the dorsal transverse bands. Scales iridescent or with bright color, absent. Tail autotomyzed near the base. Ventrally, from mental to cloacal region light gray, variegated, with dark and light spots. In belly region predominate dark brown spots.

FIGURE 4. Dorsal view of a female specimen of L. dorbignyi from the Sierra de Fiambalá, Catamarca, Argentina. Photo: C. S. Abdala.


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FIGURE 5. Map of the Northwestern Argentina, showing known localities of L. dorbignyi (diamonds) and L. scrocchii (Black circles). The arrows indicate the restricted type locality of L. dorbignyi and the type locality of L. scrocchii. The white dots indicate the populated areas.

Variation—Based on 4 paralectotypes and 17 additional specimens. Head longer (16.4–29.3; x = 19.8 than wider (14.7–20.3; x = 17.6). Head height 8.9–15.2, (x = 11.0). Neck wider than the head. Snout Vent Length 2.2 times longer than trunk length. Snout Vent Length 73.5–98.3, (x = 89.5). Tail length 78.0–117.4, (x = 96.6). Width of tail base 8.9–14.5, (x = 11.6). Humerus length 7.3–14.1, (x = 10.4). Tibial length 11.0–18.7, (x = 15.5). Foot length 21–27.2, (x = 24.4). Fourth finger length 8.3–11.4, (x =9.6) and IV toe 11.4–16.0, (x = 13.2). Dorsal surface of the head slightly rugose, with 14–19 (x = 16.5) scales. Six to 8 supercilliaries. Twelve



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to 14 upper cilliaries. Eight to 10 lorilabials in a single row equal in size to supralabials. Six to 8 supralabials scales. Frontal scale azygous or horizontally divided in 2. Interparietal scale smaller, equal to, or larger than parietals, surrounded by 5–9 scales. Nasal scale surrounded by 6–8 scales. Four to 6 infralabials. Four scales in contact with the mental scale. Five to 9 smooth horizontal temporals and 4–7 verticals. Up to 4 auriculars scales on the anterior margin of the auditory meatus. Horizontal fold “Y” shaped, between shoulder and auditory meatus. Scales around midbody 48–59 (x = 54.7). Thirty two to 52 gulars scales and 35-45 on neck (from posterior margin of auditory meatus to shoulder, along horizontal fold). Dorsal scales between occiput and hindlimbs 45–56 (x = 50.6). Seventy seven to 94 (x = 85.9) ventrals scales. Precloacal pores in males 5–8 (x = 6.5). Three of six females with 5 to 6 precloacals pores. Gulars scales 32–52 (x = 40.3). Dorsal scales flat, imbricated and slightly keeled or unkeeled. Ventral scales same size as dorsal scales. Seventeen to 22 subdigital lamellae on the fourth finger and 21–26 on the fourth toe. Without femoral patch.

FIGURE. 6. Habitat of L. dorbignyi from Sierra de Fiambalá, Catamarca, Argentina. Photo: S. Quinteros.

Color in life—Sexual dichromatism present (Fig. 2–4). Dorsal head in males, black or dark brown; gray or brown in females, but never darker than males. Dorsum and sides of the body intense yellow or dark orange, with large paravertebral spots transverse to the body axis, divided on the sides of the body (Fig. 2). Paravertebral spots black or dark brown, band-shaped. In females the dorsal background is gray or reddishbrown. Without vertebral line, dorsolateral stripes nor scapulars spots. In males the tail is dorsally light gray or greenish-gray. Ventrally yellow or intense orange from the mental scale to the cloacal region, the tail ventrally exhibit same color of dorsum (Fig. 3). Females exhibit the same pattern but lighter (Fig. 4). Distribution—The main problem with the identification of Liolaemus dorbignyi, was the lack, in the original description, of the designation of a precise type locality. According to the results obtained in this work, the distribution of L. dorbignyi is restricted to Sierra de Fiambal, Catamarca Province, Argentina. Specifically we found populations in the localities known as Puesto la Lagunita, between 35 and 38 Km NW of Medanitos, at 4350 m, and in La Cienaga, Campo El Potrerito, 3900 m. However it is probable that L. dorbignyi is distributed along the entire Sierra de Fiambalá, over the 3500 m. (Fig.5.)


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Natural History—Liolaemus dorbignyi, inhabits large rocky areas of the Sierra de Fiambalá (Fig. 6). It is a viviparous species. No more data about its biology are known. It lives in syntopy with Phymaturus cf. antofagastensis, L. cf. andinus and L. cf. puna.

Liolaemus scrocchii sp. nov. (Fig. 7–8) Liolaemus dorbignyi (non Koslowsky, 1898): Laurent (1991), Núñez (1992), Núñez and Jacksic (1992), Laurent (1995), Etheridge (1995), Laurent (1998), Schulte et al. (2000), Valladares et al. (2002), Espinoza et al. (2004), PincheiraDonoso and Núñez (2005), Cruz et al. (2005), and Abdala (2007).

Holotype—FML 17228. Male. Two Km SE of Susques on National road 52. Dto. Susques, Jujuy, Argentina. Abdala, C., Barrionuevo, S., Quinteros, S., and G., Scrocchi, Cols. 25-Feb-05. Paratypes—FML 17229. Male, MCN 2136–2137, male and female; same data as the holotype. FML 01763, 5 females, 4 males; Santa Rosa de los Pastos Grandes. Dto. Los Andes, Salta. O. Pagaburo col. 06May-86. FML 01759 male and female; Agua de los Sapos. (Puesto de Cruz Soriano), 20 Km from Mina Tincalayo. Dto. Los Andes, Salta; O.Pagaburo, col. 06-May-86. FML 01817. Male, El Quemado, Abra de Pives. Dto. Tumbaya, Jujuy, 4000 m. E. Lavilla, A. Marcus, and R. Montero cols. 20-Feb-87. FML 06481. Male. La Mesada, 64 km from La Poma to Abra del Acay. Dto. La Poma, Salta. 4665 m. S. Torres col. 21-Jul-97. FML 00992. male and 2 females. Around base of the Paño mountain. Serranía de Chañi. 4200–4300 m. Dto. Belgrano, Jujuy. G. Scrocchi col. MCN 714; MCN 898; MCN 961. Females. Casa Mocha, climbing from NW of the Nevado del Chañi. 4500–4700 m. C. Bracamonte col. 29-Mar-2002. MCN 899. Male. Abra del Cerro Toro. Sierra de Aguilar. R. Seggiaro col. 10-Oct-2002. MCN 964. Male. Road to Abra del Acay from national road 51. Near 4900 m. I. Martínez Oliver, S. Lobo and F. Lobo cols. 19-Jan-03. MCN 1922–1925. Female and 2 males. Km 210, Nacional road 51. 0.6 Km from nacional road 51. 6.4 Km to S of Olacapato. 24.2411475 S; 66.69711 W. 4070 m. B. Casimiro, Espinoza, R., Lobo, F. and S. Quinteros cols. 17-Jan-06. Diagnosis—Liolaemus scrochii belongs to the L. montanus group (Etheridge, 1995), diagnosed by a blade-like process on posterior distal tibia, associated with a greatly hypertrophied M. tibialis anticus (Abdala, et al. 2006). It has scales of equal size on the posterior face of the femur, distinguishing it from the L. boulengeri group (Abdala, 2007). Within the L. montanus group, L. scrocchii is a robust lizard, with a large size (max. SVL 95 mm) which distinguishes it from L. andinus, L. audituvelatus, L. disjunctus, L. eleodori, L. erroneus, L. etheridgei, L. fabiani, L. famatinae, L. fittkaui, L. griseus, L. huacahuasicus, L. insolitus, L. islugensis, L. molinai, L. montanus, L. multicolor, L. ortizi, L. pantherinus, L. poconchilensis, L. poecilochromus, L. pulcherrimus, L. rechei, L. rosenmanni, L. ruibali, L. signifier, and L. vallecurensis, which are smaller (SVL 50– 76 mm). Liolaemus scrocchii has flat, imbricated and slightly keeled dorsal scales which distinguish it form L. andinus, L. audituvelatus, L. erguetae, L. erroneus, L. forsteri, L. insolitus, L. islugensis, L.molinai, L. patriciaiturrae, L. poecilochromus L. robertoi, L. rosenmanni, and L. schmidti, (smooth and juxtaposed dorsal scales) and from L. aymararum, L. disjunctus, L. etheridgei, L. fittkaui, L. huacahuasicus, L. montanus, L. ortizi, L. polystictus, L. stolzmanni, L. thomasi, and L. williamsi (imbricate and strongly keeled scales). Liolaemus chlorostictus, L. jamesi, L. orientalis, L. poconchilensis, L. robustus, L. signifer and L. stolzmanni have slightly keeled but juxtaposed scales. The number of scales around midbody in L. scrocchii is 55–68, distinguishing it from L. andinus, L. disjunctus, L. duellmani, L. eleodori, L. erguetae, L. foxi, L. islugensis, L. molinai, L. multicolor, L. nigriceps, L. patriciaiturrae L. pleopholis,, L. poecilochromus, L. pulcherrimus, L. robertoi, L. rosenmanni, L. ruibali, L. signifer, and L. vallecurensis (between 64 and 110). The females of L. scrocchii have precloacal pores 54.14% (eight of fourteen females), differing from L. andinus, L. duellmani, L. fabiani, L. islugensis, L. jamesi, L. melanogaster, L. pantherinus, L. puritamensis, L. robertoi, L. rosenmanni, L. signifer, L. stolzmanni, and L vallecurensis in which females lack precloacal pores. REDESCRIPTION OF L. DORBIGNYI AND A NEW SPECIES


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The dorsal color pattern is composed of transverse paravertebral spots. These spots are band-shaped, never reach the lateral region, and are never in contact in the vertebral zone (Fig. 7). This is the main difference between L. dorbignyi, L. griseus and other members of the L. montanus group. Also, the number of scales around midbody in L. scrocchi is generally higher 55–68 (x = 61.8) than L. dorbignyi 48–59 (x = 54.5). The presence of juxtaposed or subjuxtaposed dorsal scales and the absence of keeled or slightly keeled dorsal scales further differentiate L. scrocchii from L. dorbignyi. Description of the Holotype—Snout-vent-length 87.3 mm. Trunk length 41.7. Head larger (19.2 mm) than wider (17.2 mm). Head height 11.1 mm. Eye diameter 7.1 mm. Interorbital distance 11 mm. Orbit-auditory meatus distance 7.1 mm. Auditory meatus height 2.8 mm, wide 0.9 mm. Orbit-comisure of mouth distance 3.1 mm. Internares 3 mm. Sub-ocular scale 4.3 mm long. Femur length 14.6, tibia 17.3, and foot 25.8 mm. Humerus length 11.8 mm. Dorsal surface of the head smooth, with 14 dorsal scales. Rostral wider than high, bordered by six scales. Mental larger than rostral, bordered by four scales, trapezoidal. Nasals do not contact rostral. Two internasal scales. Seven scales surrounding nasal separated from canthal by two scales. Five scales between rostral and supercilliaries, six between rostral and frontal. Frontal scale divided. Two postrostrals. Interparietal smaller than parietals, in contact with five scales. Fourteen upper cilliaries. Supraorbital semicircles incomplete. Preocular scale separated from lorilabials by one scale. Two scales on anterior margin of auditory meatus and three on upper margin. Six smooth temporals (from upper margin of the auditory meatus to postocular scale) Three lorilabials contact subocular. Seven supralabials, the fourth of equal in size to others, without scales organs, and not in contact with the subocular . Six supraoculars. Eight lorilabials. Six infralabials, the second ventrally in contact with two scales. Five chinshields, the second separated by two scales. Fifty seven scales around midbody. Forty eight, rounded, sub-juxtaposed, smooth dorsals, from occiput to hindlimbs. Twenty nine scale rows on dorsum. Ventrals scales same size as dorsals, flat and juxtaposed. Forty eight sub-juxtaposed and smooth gulars. Five precloacal pores. Antehumeral scales flat, larger or equal to dorsals. Postauricular, rictal and longitudinal folds present. Scales on the longitudinal fold, granular and smooth. Fourth finger with 17 subdigital lamellae and 26 on fourth toe. Flank scales, rounded, smooth with some small granular intercalated scales. Without femoral patch. Flat, imbricate and triphid infracarpal scales. imbricated and keeled infratarsal scales. Color in ethanol—Head dorsally dark brown, with scattered dark gray scales. Head darker than rest of body. Trunk reddish-brown, with seven conspicuous, rhomboidal, black paravertebral spots, arranged transverse to body axis; posteriorly spots in contact in vertebral zone. A black spot present on posterior region of trunk, owing, probably, to scale damage. Fore and hind limbs same color as trunk, but without dark spots. Vertebral line, scapular spots and dorsolateral stripes absent. Lateral region of body with same color trunk, but light gray in ventrolateral zone. Iridescent or brightly colored scales, absents. Dorsally tail with same color as trunk. Cloacal region gray. Ventrally, from mental to cloacal region light gray, variegated with dark and light spots. On thighs a light yellow color. Tail exhibits a light gray, ringed pattern. Variation—Based on 27 paratypes. Head larger (15.1–20.6 mm; x = 17,9 mm, than wider (13.0–19.4 mm; x = 16.4 mm). Head height 9.1–12.3 mm (x = 10.8 mm). Neck wider than head. Snout-vent length, 2.1 times longer than trunk. Snout-vent length 72.8–94.5 mm (x = 83.9 mm). Tail length 85.1–101.3 mm, (x = 92.3 mm). Tail base width 8.9–14.1 mm, (x = 11.3 mm). Humerus length 5.5–12.7 mm, (x = 9.3 mm). Tibia length 12.2–18.5 mm, (x = 16.3 mm). Foot length 21.8–27.9 mm, (x = 24.0 mm). Fourth finger length 7.1– 10.1 mm, (x =9.0 mm) and fourth toe 11.1–17.3 mm, (x = 14.1 mm). Dorsal surface of head, slightly rugose, with 14–19 scales. Six to 7 supercilliaries. Twelve to 16 upper cilliaries. One row of lorilabials same size as supralabials. Seven to 10 lorilabials. Six to 8 supralabials. Frontal scale divided in two, three or four. Interparietal scale larger, smaller or equal to parietals, and surrounded by 5–8 scales. Nasal scale surrounded by 5–8 scales. Four to 6 infraliabals. Mental in contact with 4 scales. Four to 8 smooth horizontal temporals. Up to 4 auriculars scales on anterior margin of auditory meatus. Neck folds present; horizontal fold


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is “Y” shaped between shoulder and auditory meatus. Scales around midbody 55–68 (x = 61.8). Gulars 34–46 and 36–48 scales on neck (from posterior margin of auditory meatus to shoulder, along horizontal fold). Dorsal scales between occiput and hind limbs 42–53 (x = 49.6). Ventrals 72–98 (x = 86.4). Males with 2–10 (x = 5.6) precloacal pores. Among fourteen females, eight exhibits 3–6 precloacal pores. Flat, juxtaposed or subjuxtaposed, smooth dorsal scales. Sixteen to 21 infradigital lamellae on fourth finger and 23–28 in fourth toe. Without femoral patch.

FIGURE 7. Dorsal view of a male specimen of L. scrocchii from the type locality, near Susques, Jujuy, Argentina. Photo: G. Scrocchi.

FIGURE 8. Ventral view of a male specimen of L. scrocchii from the type locality, near Susques, Jujuy, Argentina. Photo: G. Scrocchi. REDESCRIPTION OF L. DORBIGNYI AND A NEW SPECIES


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FIGURE 9. Habitat of L. scrocchii from type locality, near Susques, Jujuy, Argentina. Photo: G. Scrocchi.

Color in life—(Fig. 7–8) Without sexual dichromatism. Dorsal head design uniform, light brown or reddish-brown, with yellowish shades and dark or black scales. Body background light brown or reddish-brown, in some specimens orange tones are present. Large dark brown or black rhomboidal or rectangular paravertebral spots, always transverse to body axis (Fig. 7). These spots never fused in vertebral zone of trunk, but are fused at base of tail. Lateral fields yellow, orange or reddish-brown, darker than dorsum. Light blue or white scales irregularly spread on laterals fields and on fore and hind limbs. Without vertebral line, dorsolateral bands, or scapulars spots. Tail light brown, white or yellow shaded. Paravertebral spots disappear toward end of tail. Ventrally intense yellow from mental to cloacal region, variegated with dark and light brown scales (Fig. 8). Some specimens with gular region orange or reddish. Tail pattern same as that of body. Distribution—Liolaemus scrocchii occurs in the Puna region of Jujuy and in the northwest of Salta province (Fig. 5). In Jujuy province it is known from Susques, Abra de Pives, Cerranías de Chañi, Chorcan, Abdón Castro Tolay, Sey and Sierra de Aguilar, and in Salta province it is found at Abra del Acay, Olacapato, Santa Rosa de los Pastos Grandes, Huaytiquina and Mina Tincalayo (Fig 3). Natural History—Liolaemus scrocchii, is known from localities in the phytogeographic region of the Puna (Cabrera and Willink, 1980), always above the 3000 m. It is commonly found basking on rocks and on the sand banks nearby (Fig. 9). It is viviparous, omnivorous with a high degree of plant material in its diet (Espinoza et al. 2004). It lives in sympatry with L. multicolor, L. ornatus Koslowsky and L. puna Lobo and Espinoza. Etymology—We name this new taxon scrocchii to acknowledge our colleague and friend Gustavo Scrocchi for his extensive and dedicated work on the systematics and biology of Argentine reptiles, and in particular for his invaluable kindness and companionship. Remarks—The species described here belongs to the L. montanus group (Etheridge, 1995) within the subgenus Eulaemus (Laurent, 1983). The squamation and color pattern distinguish it from L. dorbignyi and the other species of the genus. The species here described inhabits many localities of the Puna in Jujuy Province and in northwestern Salta Province, between 23º–25º S and 65º–67º W. This species is viviparous, saxi-


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colous and lives on rocky mountain slopes. Individuals of L. scrocchii from the Nevado del Chañi (Jujuy province) and from Abra del Acay (Salta, Province), in captivity, exhibit a particular behavior to stay, part of the day, in water.

Discussion All previous published studies in which the name Liolaemus dorbignyi was used were based on specimens, described here as L. scrocchi. Núñez and Jaksic (1992), synonymyzed L. puritamensis (Núñez and Fox, 1989) with L. dorbignyi, without studying the type material of Koslowsky (1898), but instead based their decisions on specimens of L. scrochii deposited in the IBA (Instituto de Biología Animal, Mendoza). However, Peters and Donoso-Barros (1970) provided a schematic drawing of the dorsal pattern of L. dorbignyi (their Fig 35, page 177), which totally differs from that of L. puritamensis. Years later, Pincheira-Donoso and Núñez (2005) revalidate L. puritamensis, to specific status, but in this book they present a photograph of L. puritamensis, comparing it with L. scrocchii (their Fig 15, page 119). The presence of L. dorbignyi in Chile is uncertain. Donoso-Barros (1960, 1966) and Donoso-Barros and Codoceo (1962) cite L. dorbignyi from Laguna Amarga in southern Chile near 52º S latitude (far away from the type locality of L. dorbignyi, approximately 27º S). Donoso-Barros (1966) provided a picture (ventral view) of a specimen identified as L. dorbignyi from Laguna Amarga, but this picture corresponds to L. sarmientoi Donoso Barros (Pincheira-Donoso and Núñez, 2005). In this last work L. dorbignyi is not listed as a member of the Chilean herpetofauna. Both L. dorbignyi and L. scrochii belongs to the montanus group of Etheridge (1995). According to Cei (1993) L. dorbignyi belongs to the signifer group with L. nigriceps and L. orientalis while the montanus group is formed by L. griseus, L. huacahusicus and L. montanus. Few studies have addressed the relationships of the montanus group. In a phylogenetic analysis of Liolaemus using the COI, NDI, NDII and different regions of tRNA (tRNAGln, tRNAIle, tRNAMet, tRNATrp, tRNAAla, tRNAAsn, tRNACys, tRNATyr), Schulte et al. (2000) found the same major groups that Laurent (1983) referred to as the subgenera Liolaemus and Eulaemus. Inside the Eulaemus group there are two sections, one of which is the montanus section. This section is divided into two series, the monatnus series and the boulengeri series. The montanus series represents the montanus group of Etheridge (1995) with the exception of the boulengeri subgroup. Etheridge (1995) considered the boulengeri group (diagnosed by the presence of a femoral patch) to be a subset of the montanus group because he could find no morphological synapomorphies that would define the species if the montanus group was not included within the boulengeri subgroup. For Schulte et al. (2000) the monatnus series is a monophyletic group formed by L. ruibali, L. andinus-La Rioja, L. famatinae, L. orinetalis, L. dorbignyi, L. poecilochromus, L. multicolor and L. andinus-Jujuy. Espinoza et al. (2004) found a monophyletic montanus group formed by L. orientalis chlorostictus, L. ruibali, L. famatinae, L. andinus andinus, L. dorbignyi, L. andinus poecilochromus, L. cf. andinus, and L. multicolor. Although several phylogenetic analyses have been made, the phylogenetic position of L. dorbignyi remains unknown, because in all the analyses the researchers had identified specimens of L. scrochii as L. dorbignyi (Schulte et al., 2000, Valladares, et al., 2002, Espinoza et al., 2004, Cruz et al., 2005). In Schulte et al. (2000) the specimen identified as L. dorbignyi (SDSU 3443, from Olacapato, Los Andes in Salta province) is actually L. scrocchii. In this analysis L. scrocchii is the sister taxon to a clade formed by (L. poecilocrhomus (L. multicolor - L. andinus-Jujuy)) within the montanus series. The analysis of Valladares et al.'s (2002) use the same genes as Schulte et al. (2000). This hypothesis shows a polytomy formed by L. scrocchii, L. molinai, L. aymararum and a clade formed by (L. poecilochromus (L. multicolor – L. andinus)). In Espinoza et al. (2004) the results are identical to those of Schulte et al. (2000). For Cruz et al. (2005) L. scrocchii is the sister taxon of L. poecilochromus.



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The behavior to stay in water is similar to that described by Labra et al. (2002) for L. fabiani in the Atacama Desert. According Labra et al. (2002), L. fabiani use the water, part of the day, as a shelter and avoids overheating. Both L. fabiani Yañez and Nuñez, and L. scrocchii belong to the montanus series in the phylogenetic analysis performed by Valladares et al. (2002), which could explain why these two species share this behavior.

Material examined Species names are arranged alphabetically by species, then by country, province, museum acronym, number and details of the localities. Liolaemus andinus: Argentina, Catamarca: FML 1409, Aguas calientes, Dto. Tinogasta. FML 1913; 2025, Las Grutas, to 20 Km of Paso San Francisco, Dto. Tinogasta. FML 2545, Near Rio Chaschuil, 42 Km SE of Paso San Francisco, Dto. Tinogasta. FML 2546, 68 Km NW of San Francisco river, Dto. Tinogasta. MCN 1648–53, Cazadero Grande, Dto. Tinogasta. Chile, III Región: FML 2553, Salar de Maricunga. Liolaemus chlorostictus: Argentina, Jujuy: FML 1515, Cuesta de Fundiciones, Mina Pirquita. Dto. Rinconada. Liolaemus dorbignyi: Argentina, Catamarca: MLPS 057; MLP S 844–45; MLPS991–92, Cordillera de Catamarca. MCN 2119–21, Puesto la Lagunita, 35–38 km NE of Medanitos, climbing from Medanitos. Sierra de Fiambalá, Dto. Tinogasta. MCN 2132–33, Los Aujeritos, 34–36 km NE de Medanitos, 2 Km SW of Puesto la Lagunita, climbing from Medanitos. Sierra de Fiambalá, Dto. Tinogasta. FML 1855. Saujil, Road 60, Dto. Tinogasta. FML 17230–236, Puesto la Lagunita, 35–38 km NE of Medanitos, climbing from Medanitos. Sierra de Fiambalá, Dto. Tinogasta. Liolaemus eleodori: Argentina, San Juan: FML 1703, Llano de los Hoyos, San Guillermo river. Dto. Iglesia, San Juan. Liolaemus famatinae: Argentina, La Rioja: FML 232. Cerro de Famatina, Dto. Famatina. FML 1720, Cueva de Perez, Nevados de Famatina, Dto. Famatina. Liolaemus fitkaui: Bolivia, Cochabamba: FML 1612, Tiraque, Dto. Cochabamba. Liolaemus forsteri: Bolivia, La Paz: FML 2211, Chacaltay, Dto. La Paz. Liolaemus griseus: Argentina, Tucumán: FML 1354, Vega Mataderos, Vacahuasi, Tafi del Valle. FML 1582, SW of Cerro Lomo Ballena, Cumbres Calchaquíes, Tafi del Valle. FML 1586, Cerro Negrito, Cumbres Calchaquíes, Tafi del Valle. Liolaemus huacahuasicus: Argentina, Catamarca: FML 486–7, El Cerrillo. FML 665, 1 Km S of Cerro El Overo, Nevados del Aconquija, Dto. Santa Maria. FML 674–5, Filo Colorado, 2–3 Km W Cerro el Overo, Nevados del Aconquija, Dto. Santa María. FML 677, 5 Km W of Nevado del Candado, Dto. Santa María. FML 885, Nevado de las Animas, Dto. Santa Maria.1224, Filo los Heladitos, climbing from Capillitas, Dto. Andalgala. Tucumán: FML 0066, Laguna de El Negrito, Dto. Tafi del Valle. FML 00203; FML 0469–70, Cerro El Negrito, Dto. Tafi del Valle. Liolaemus jamesi: Chile, I Region Tarapacá: FML 1193, Tarapacá, Punta Arica. FML 1775, Parinacot, Arica. Liolaemus montanus: Argentina, Catamarca: FML 279; 289, El Rodeo, Dto. Ambato. FML 908–10; 981–2; 1723; 2141, Cerro El Manchao, Dto. Ambato. Liolaemus multicolor: Argentina, Jujuy: FML 00259; 0272; 1873; 2065; 2098; 2349; 2592, Abrapampa, Dto. Cochinoca. FML 1464; 1539, 3 Km NW of Abrapampa, Dto. Cochinoca. Liolaemus nigriceps: Argentina, Salta: FML 1633, South of Socompa Volcano, Dto. Los Andes. FML 1635, South of Socompa Volcano, Dto. Los Andes. Liolaemus orientalis: Argentina, Jujuy: FML 928; 949, Road to Laguna Blanca, Dto. Humahuaca. FML 930; 938–39; 944, Chorcan, Dto. Humahuaca.


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Liolaemus poecilocromus: Argentina, Catamarca: FML 00878, Salar de Antofalla, Dto. Antofagasta de la Sierra. FML 01171, Los Nacimientos, Dto. Antofagasta de la Sierra. FML 01173, Volcanes of Los Nacimientos, Dto. Antofagasta de la Sierra. FML 3068, Ojo de Calalaste, Dto. Antofagasta de la Sierra. FML 0369–71, Las Quinuas, Salar de Antofalla, Dto. Antofagasta de la Sierra. MCN 249–51; 288–91, 4–5 Km N of Antofagasta de la Sierra, Dto. Antofagasta de la Sierra. MCN 278–79; 298–301, Paycuqui, Dto. Antofagasta de la Sierra. MCN 280–82, 3 Km N of Antofagasta de la Sierra, Dto. Antofagasta de la Sierra. MCN 285–87, Cuesta de Randolfo, Dto. Antofagasta de la Sierra. Liolaemus pulcherrimus: Argentina, Jujuy: FML 1961, Mudana, Dto. Humahuaca. FML 2184, Mudana, Dto. Humahuaca. FML 18238–249, Road to Mudana, 23º 20´30´´ S 65º 13´27.5´´ W, Dto. Humahuaca. Liolaemus puritamensis: Argentina, Jujuy: MCN 2498–99, West Coast of Vilama Lake, 4495 msnm, 22º33'38''S; 66º54'55,6''W, Dto. Rinconada . MCN 2509–13, Road to Isla Grande lake, 4533 msnm, 22º35'46,3''S; 66º42'35''W, Dto. Rinconada. FML 985, Vega Afluente Vilama Lake, Dto. Rinconada. Liolaemus ruibali: Argentina, La Rioja: FML 2975–76, Agua Quemada, 47 Km E from Alto Jagüe, Dto. Vinchina. Mendoza: FML 1076; 1104, Paramillo de Uspallata, Dto. Las Heras. FML 2549, National Road 7, Paramillo, Uspallata, Dto. Las Heras. FML 3462–63, Twenty four Km SE of Uspallata, Dto. Las Heras. Liolaemus schmidti: Chile, II Region Antofagasta: FML1192, Antofagasta, Atacama. FMl 1197, Volcán Tatio, Antofagasta. Liolaemus scrocchii: Argentina, Jujuy: FML 17228–29, 2 Km SE of Susques over National road 52. Dto. Susques. FML 01817–1, El Quemado, Abra de Pives. Dto. Tumbaya. FML 00992, Around base of the Cerro Paño. Serranía de Chañi, Dto. Belgrano. MCN 714; MCN 898; MCN 961, Casa Mocha, climbing from NW of the Nevado del Chañi. MCN 899, Abra del Cerro Toro. Sierra de Aguilar. Salta: FML 01763, Santa Rosa de los Pastos Grandes, Dto. Los Andes. FML 01759, Agua de los Sapos. (Puesto de Cruz Soriano), 20 Km from Mina Tincalcayo, Dto. Los Andes. FML 06481, La Mesada, 64 Km from La Poma to Abra del Acay, Dto. La Poma. MCN 964, Road to Abra del Acay from national road 51, Dto. Los Andes. MCN 1922– 25, Km 210, Nacional road 51, 0.6 Km from national road 51, 6.4 Km to S of Olacapato. 24.2411475 S; 66.69711 W, Dto. Los Andes. Liolaemus signifer: Perú: FML 1434, Titicaca Lake, 3840 msnm. FML 1557, Titicaca Lake, road to Puno. Liolaemus vallecurensis: Argentina, San Juan: FML 2404, Valle de Cura, Dto. Iglesia.

Acknowledgments We thank to J. Faivovich and J. M. Díaz Gómez for valuable comments of a first draft. We would like to thank E. Lavilla, S. Kretzschmar (FML) and J. Williams (MLP) for allowing us access to their collections. We thank to F. Cruz, R. Montero, R. E. Espinoza, J. M. Díaz Gómez, S. Barrionuevo, G. Scrocchii, F. Arias, S. Valdecantos and O. Pagaburo, for helping us in field, lab or discussing ideas related to this work. We acknowledge the provincial departments of Fauna of Catamarca, Jujuy and Salta for providing permits for the collection of specimens. This study was supported by grants (FL) from Consejo de Investigaciones de la Universidad Nacional de Salta (CIUNSa, 1236), and from Consejo Nacional de Investigaciones Científicas y Técnicas (PIP 5986), a grant from Consejo Nacional de Investigaciones Científicas y Técnicas (PIP, 6287) to F. Cruz, (PIP, 5780) to E. Lavilla, and Consejo de Investigaciones de la Universidad Nacional de Tucumán (CIUNT, G315) to Ricardo Montero.



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