Article LUCIANO JAVIER AVILA 1,4, MARIANA MORANDO 1, DANIEL ROBERTO PEREZ 2 & JACK W. SITES, JR 3 1. Abstract. Resúmen.

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1 Zootaxa 2667: (2010) Copyright 2010 Magnolia Press Article ISSN (print edition) ZOOTAXA ISSN (online edition) A new species of the Liolaemus elongatus clade (Reptilia: Iguania: Liolaemini) from Cordillera del Viento, northwestern Patagonia, Neuquén, Argentina LUCIANO JAVIER AVILA 1,4, MARIANA MORANDO 1, DANIEL ROBERTO PEREZ 2 & JACK W. SITES, JR 3 1 CENPAT-CONICET. Boulevard Almirante Brown 2915, U9120ACD, Puerto Madryn, Chubut, Argentina. avila@cenpat.edu.ar; morando@cenpat.edu.ar 2 Escuela Superior de Salud y Ambiente, Universidad Nacional del Comahue. Buenos Aires Neuquén. Argentina. ddeneuquen@yahoo.com 3 Department of Biology and M. L. Bean Life Science Museum, Brigham Young University, 401 WIDB, 84602, Provo, Utah, United States of America. jack_sites@byu.edu 4 Corresponding author: avila@cenpat.edu.ar Abstract A new species of lizard of the genus Liolaemus from Cordillera del Viento, northwestern Neuquén Province, Argentina is described. The new species is a member of the Andean-Patagonian Liolaemus elongatus clade and is known only from a single locality above 3000 m in the eastern slopes of Domuyo Volcano. Liolaemus antumalguen sp. nov. is a stout species, saxicolous, probably viviparous and omnivorous, and seems to have low population density. It is easily differentiated from all other species of the elongatus complex for a highly variable but characteristic dorsal pattern and a completely black ventral coloration. Key words: Sauria, Iguanidae, Liolaemus antumalguen sp. nov., new species Resúmen Una nueva especie de lagartija del genero Liolaemus de la Cordillera del Viento, noroeste de la provincia de Neuquén, Argentina, es descrita. La nueva especie es miembro del cclado andino-patagonico Liolaemus elongatus y es conocida solo para una unica localidad sobre los 3000 m de altura en el faldeo oriental del volcán Domuyo. Liolaemus antumalguen sp. nov. es una especie robusta, rupícola, probablemente vivípara y omnívora, y parece tener baja densidad poblacional. Se diferencia fácilmente de todas las otras especies del grupo por un patrón de coloración dorsal variable pero característico y por poseer la zona ventral completamente negra. Palabras claves: Sauria, Iguanidae, Liolaemus antumalguen sp. nov., nueva especie Introduction Lizards of the genus Liolaemus are a common vertebrate component of the Southern Andean and Patagonian biotas. With a number of described species reaching 225, the genus Liolaemus constitutes the most speciesrich squamate genus of southern South America. During the last 15 years more than 50 species were described mainly in Argentina and Chile, and one of the main causes of this dramatic recent increase in the number of known species is that new collections are being made in previously unexplored areas throughout the Andes and Patagonia. Several new species limited to small basins, high-elevation valleys or single mountaintops, have been discovered in recent years in northwestern Patagonia and the Southern Andes (e.g. Abdala 2002, 2003, 2005; Avila et al. 2003, 2004, 2006, 2007a; Cei and Videla 2003; Etheridge and Christie 2003, 28 Accepted by S. Carranza: 7 Oct. 2010; published: 4 Nov. 2010

2 Pincheira-Donoso and Scolaro 2007, Pincheira-Donoso et al. 2007; Videla and Cei, 1996), and new survey efforts carried out in several isolated areas of Patagonia, as well as molecular studies of several lizards clades, demostrate that species diversity in this area is greater than presently known (Morando 2004, Morando et al. 2003, 2007). Laurent (1983, 1985) split the genus Liolaemus in two subgenera: Liolaemus sensu stricto and Eulaemus, criteria followed by Etheridge (1995) and supported in a molecular study by Schulte et al. (2000). Within the subgenus Liolaemus, several subgroups can be recognized on the basis of morphological traits (Lobo 2001, 2005), and in a molecular study, Morando et al. (2003) defined a well-supported clade of saxicolous lizards distributed across mountain landscapes of Andes and volcanics plateaus of Patagonia. This was recognized as the Liolaemus elongatus-kriegi complex (Cei 1979, Morando et al. 2003), and now includes three sub-clades: petrophilus, kriegi, and elongatus. As currently known, the elongatus clade includes L. elongatus Koslowsky 1896, L. chillanensis Müller and Hellmich 1932, L. thermarum Cei and Videla 1996 (Morando unpublished data, Torres-Perez et al. 2009, but see Lobo et al and discussion below), and at least twelve still undescribed species. These species are distributed in the western slope of the Andes, restricted to Andean and Patagonian steppe environments of southwestern Mendoza and western areas of Neuquén, Rio Negro, Chubut, and probably Santa Cruz provinces, in Argentinean Patagonia (Morando et al. 2003; Morando 2004). Other authors have found or suggested different arrangements for these lizards (Lobo 2001, 2005; Pincheira- Donoso and Nuñez 2005, Pincheira-Donoso et al. 2008, but see Lobo et al. 2010). Resolution of alternative hypotheses of relationships must await more inclusive geographic, taxonomic, and character sampling. During a recent field exploration, a sample of Liolaemus was collected on a large volcanic outcrops and lava flows on the eastern slopes of the Domuyo Volcano, the highest peak of southern Andes in northwestern Neuquén province. A morphological and molecular analysis of these lizards show that they represent a new taxon in the elongatus species clade, which we describe here as a new species. Material and methods We examined sample series of the species determined by Morando et al. (2003) and Morando (2004) as members of the elongatus clade (Table 1, Appendix I), and samples of Liolaemus buergeri, L. ceii, L. flavipiceus, L. kriegi, and L. punmahuida, because they are distributed in the closest mountain range region and at similar altitudes, also because of their body sizes or general morphology they can be confused with the new species. All are deposited in the herpetological collections of Fundación Miguel Lillo (FML), Argentina; Monte L. Bean Museum, Brigham Young University (BYU), Provo, USA; Museo de La Plata, Universidad Nacional de La Plata (MLP.S), Argentina; Museum of Vertebrate Zoology, UC Berkeley (MVZ), Berkeley, USA; Museo Argentino de Ciencias Naturales (MACN), Argentina, and the collection LJAMM-CNP of the Centro Nacional Patagonico (CENPAT-CONICET), Puerto Madryn, Argentina. Ten specimens of the new species were examined. Specimens were collected by hand, sacrificed by a pericardiac injection of sodium pentothal Abbot, fixed in 10 20% formalin and later transferred to 70% ethanol. Measurements were taken with a Mitutoyo digital caliper to the nearest 0.1 mm. Some character states were observed with the aid of a binocular stereomicroscope. Scale definitions, measurements, and chromatic characters follow Smith (1946), and recent treatments of the genus Liolaemus by Etheridge and Christie (2003) and Espinoza et al. (2000); neck-fold terminology follows Frost (1992). Descriptions of color in life are based on notes taken in the field and color photographs of recently captured animals. Data for some species used for comparison were obtained from the original description, Cei (1986) and Pincheira-Donoso and Ñuñez (2005). Protocols for DNA extraction, mtdna primer descriptions, PCR, and sequencing procedures for the cytochrome b (811 bp) region, as well as analytical methodology follow Morando et al. (2003) and Morando (2004). A NEW SPECIES OF LIOLAEMUS FROM PATAGONIA Zootaxa Magnolia Press 29

3 TABLE 1. Morphometric, meristic, and chromatic characteristics from species phylogenetically related to the Liolaemus elongatus clade. Data for Liolaemus chillanensis were taken from Pincheira-Donoso and Nuñez (2005). Character TERMS OF USE Liolaemus antumalguen (n = 9) Liolaemus chillanensis Liolaemus elongatus (n = 128) Liolaemus thermarum (n = 27) Maximum SVL (mm) Scales around midbody 78.5±3.5(72 82) ±4.0(68 87) 73.37±3.5(66 79) Scales between occiput 73.6±2.6(70 78) ±5.2(60 87) 71.3±4.3(64 83) and rump Ventral scales 109.2±4.9( ) ±5.6(92 121) 103.4±4.9(94 112) Dorsal pattern Variable, from patternless to two dorsolateral series of black ocelli sometimes fused longitudinally Variable but usually a wide vertebral band, with longitudinal lines of black dots in longitudinal lines A vertebral band formed by transversal black lines irregularly fused, flanked by two more clear dorsal longitudinal bands A vertebral band formed by small and irregular black dots, sometimes fused forming a black vertebral solid band to almost disappearing Ventral melanism Present Absent Absent Absent Precloacal pores Body color Light gray to ochre Ochre to light gray Ochre to almost black Light tan to ochre Tail rings absent absent Present in some populations absent Results Liolaemus antumalguen sp. nov. (Figure 1, 2) Type material. Holotype. MACN 38985, an adult male from rocky boulders from the eastern piedmont of Domuyo volcano, around Chadileu Creek (36º 39 S, 70º 20 W), Chos Malal Department, Neuquén Province, Argentina; collected by D. R. Perez; 23 Enero Paratypes. MACN 38987, MLP.S , (males); BYU 12592, MACN 38986, MLP.S , LJAMM (females); same data as holotype. Diagnosis. Liolaemus antumalguen is a member of the elongatus clade that currently includes L. elongatus, L. thermarum (Morando et al., 2003; Morando, 2004), L. chillanensis (Torrez-Perez et al. 2009), and many candidate species represented by well-supported mtdna haploclades (Morando et al. unpublished). Differs from all other members of the clade due to a unique and very variable dorsal coloration pattern, ranging from large irregular black dots distributed along the dorsolateral areas on ochre background coloration, to an almost plain pattern without any dorsal mark. Liolaemus antumalguen has a complete ventral melanism in adult specimens, characteristic not observed in any other species within the L. elongatus clade, with the exception of populations of L. elongatus from western Rio Negro province, Argentina (Morando et al. 2003, Avila et al. unpublished data). It is the largest species in the clade (maximum SVL mm vs 94 mm in L. elongatus, vs 70.3 mm in L. chillanensis, and 85.0 mm in L. thermarum), and at difference of all other species has a robust body vs a more slender shape, with well developed and evident neck folds. Liolaemus elongatus is characterized by a pattern of longitudinal bands, a vertebral band defined by irregular transversal black lines, frequently fused, with a dark background, two more clearly-defined dorsal longitudinal bands, and dorsolateral bands with black, gray, and a few white scales, usually the tail is ringed; all of these characteristics are absent in L. antumalguen. Liolaemus antumalguen differs from L. thermarum in that males of this species lack precloacal pores (according to original description) or have less (2 3 vs 4, according to our data), and have a dorsal coloration characterized by two wide and dark lateral bands; a pattern never present in L. antumalguen. Liolaemus chillanensis have slender and smaller body, with higher midbody scale count (81 95 vs 72 82), lack of conspicuous neck pouches, have a white or gray venter, and in 30 Zootaxa Magnolia Press AVILA ET AL.

4 life a conspicuous lateral and posterior body cyan coloration never found in L. antumalguen. From L. buergeri, L. ceii, and L. kriegi, L. antumalguen can be distinguished because this species have smaller dorsal scales, and a higher number of scales around midbody (94 98, , vs 72 82), lack of ventral melanic coloration (usually is white or gray, with small grey dots or uniformily slight grey), and have darker body coloration. Liolaemus curis is a smaller species (maximum SVL vs mm), have completely black body coloration or with a dorsal pattern of longitudinal bars not observed in L. antumalguen. Liolaemus flavipiceus have smaller size (SVL 90.3 vs mm) and a very particular dorsal coloration with widely distributed yellow and/or orange scales never found in L. altumalguen. Liolaemus leopardinus, L. ramonensis and L. valdesianus are smaller species (87.5, 86.6, 90.4 mm vs mm), with different dorsal coloration patterns, and higher number of midbody scales in L. ramonensis (87 96 vs 72 82). Liolaemus antumalguen can be easily distinguished from L. punmahuida by the lack of the conspicuous black line between orbit and nasal scales, white cream head coloration, and absence of any extendend brightly red or yellow ventral coloration. Liolaemus tregenzai, L. coeruleus, and L. neuquensis are smaller species (80.2, and 61.1 mm vs mm), have a conspicuous cyan ventral coloration and lack of precloacal pores. Liolaemus cristiani have a very conspicuous cyan coloration, with a dorsal pattern very different from L. antumalguen, and lack of extended ventral melanism and precloacal pores. Description of holotype. Adult male (Fig. 1, 2) mm snout-vent length (SVL); tail length mm, regenerated 70.4 mm. Axilla-groin distance 46.0 mm. Head length 22.5 mm (from anterior border of tympanum to tip of snout), 19.5 mm wide (at anterior border of tympanum), 11.5 mm high (at anterior border of tympanum). Snout length 7.3 mm (posterior margin of canthal to tip of snout). Interorbital distance 4.2 mm. Eye-nostril distance 6.4 mm. Orbit-auditory meatus distance 8.9 mm. Orbit-tip of snout distance 9.9 mm. Forelimb length 29.6 mm. Tibial length 21.1 mm. Foot length 30.8 mm (ankle to tip of claw on fourth toe). Dorsal head scales bulged, rough, 14 between occiput at level of anterior border of tympanum to rostral, pitted with numerous scale organs. Rostral scale wider (4.2 mm) than high (2.0 mm). Two postrostrals, together with anterior lorilabial, separate nasal scales from rostral. Nasal scales longer than wide, irregularly triangular; nostril over one-half length of nasal, posterior in position. Scales surrounding nasals 8 8. Four internasals, central scales four times bigger than laterals, right subdivided, with small supernumeraries behind. Frontonasal 8, irregular in size and position, with several supernumeraries as small granular scales; prefrontals 5, almost equal in size, but irregular in shape, with scar damage. A subtriangular scale on each side overlapping the superior border of posterior canthal. Frontal scale with scar damage and small supernumeraries. Frontoparietals in two rows, two anterior and three posterior scales. Interparietal depressed, with scar damage, subpentagonal, surrounded by six scales; four smaller in front and sides, two larger in back. Parietal eye not evident. Parietals bulged, oval to irregularly shaped, two similar in size to interparietal, other smaller. Circumorbitals: Transversally expanded supraoculars 7 7. Smaller lateral supraoculars: Anterior canthal wider than long, separate from nasal by two postnasals. Posterior canthal longer than wide; a well marked canthal ridge. Loreal scales, concave (4 3). Lorilabials small, reduce (2 4). Superciliaries 8 7, flattened and elongated, anterior four broadly overlapping dorsally. Orbit with upper and lower ciliaries. Orbit diameter 2.5 x 4.5 mm. Preocular small, unfragmented, longer than wide. Subocular scale elongated, six times longer than wide (6.5 x 1 mm). A well marked longitudinal ridge along upper margin of preocular and subocular scales. Postocular small, slightly bulged, without ridge. Palpebral scales small, irregular, flat, a few anterior granular and bulged. Supralabials 7 8, convex. Fifth supralabial curved upward posteriorly slightly overlapping sixth scale. Temporals smooth, bulged, subimbricate. Anterior auriculars slightly smaller than adjacent posterior temporals, projecting outward (2 3). Posterior auriculars small, granular, outprojecting. External auditory meatus inconspicuous, higher (4.5 mm) than wide (1.1 mm). Lateral scales of neck granular with inflated skin. Mental scale wider (4.9 mm) than high (2.6 mm), in contact with four scales. Mental followed posteriorly by two rows of five chinshields. Five infralabials on each side, first on each side quadrangular, two times wider than supralabials; all others stretched, slightly smaller than supralabials. Gular scales smooth, flat, imbricate, with rounded posterior margins. Scales of throat between chinshields slightly juxtaposed, becoming slightly imbricate toward auditory meatus. Forty-two gulars between tympanum openings. Infralabials separated from chinshields by one to three rows of smaller scales. A NEW SPECIES OF LIOLAEMUS FROM PATAGONIA Zootaxa Magnolia Press 31

5 Antehumeral, gular, longitudinal neck, rictal, dorsolateral, and postauricular well developed, large, very distinctive; oblique unconspicuous. A distinct lateral body fold in the first quarter between axilla and groin. Scales of dorsal neck region granular, projecting outward. Seventy-eight dorsal scales between occiput and anterior surface of thighs. Dorsal body scales rhomboidal to lanceolate, imbricate, with a keel well marked on dorsal region. At midbody, eighteen longitudinal rows of keeled scales. Dorsal scales grade laterally into slightly smaller, smooth scales at midbody. Scales immediately anterior to, above, and posterior to forelimb and hindlimb insertion small, smooth, granular, and non-overlapping, with conspicuous 1 2 organ scales. Body lateral scales grading from rhomboidal to quadrangular at midbody. Ventral body scales quadrangular, smooth, flat, imbricate, same size or a little smaller than dorsal scales. Eigthty two scales around midbody; scales between mental and precloacal pores 107. Scales of cloacal region about equal in size to ventral body scales. Four insconspicuous precloacal pores. FIGURE 1. Liolaemus antumalguen, holotype adult male (MACN 38985), from Chadileu Creek, eastern piedmont of Domuyo Volcano, Chos Malal Department, Neuquén Province, Argentina. Anterior suprabrachials rhomboidal, some faintly keeled, equal in size to dorsal body scales, grading into rounded and smooth scales posteriorly. Posterior suprabrachials smaller, smooth, becoming granular near axilla. Anterior antebrachials similar to suprabrachial. Posterior antebrachials smaller, smooth, rounded. Supracarpals imbricated, rhomboidal, smooth. Infracarpals strongly imbricate, rhomboidal, keeled. Pre and postdigital scales of manus smooth. Subdigital lamellae with three blunt keels, each terminating in a short mucron, numbering: I: 15, II: 24, II: 22, IV: 16, V: 11. Claws robust, moderately curved, black. Anterior suprafemorals as large as dorsal body scales, rhomboidal to lanceolate, slighltly keeled or smooth. Posterior suprafemorals small, granular shape. Supratibial rhomboidal to lanceolate, keeled, grading into rounded, smooth, posterior supratibials, same size as ventral body scales. Supratarsal and first supradigital keeled, middle and distal supradigital smooth. Infratarsal small, rhomboidal, imbricate, keeled, some with a small mucron. Subdigital scales keeled, most with two or three keels, mucronate, numbering: I: 12, II: 18, III: 23, IV: 26, V: 19. Claws robust, moderately curved, opaque brown or black. Dorsal and lateral caudals on non-regenerated tail, slightly keeled, ventral smooth. Caudal scales on regenerated tail strongly keeled and mucronated, ventrally becaming less keeled and non-mucronated. Color in life. Ground color dorsal and lateral scales in body, limbs, and non-regenerated portion of tail ochre to light brown, with lighter margins (Fig. 1). Dorsal head scales black. Two conspicuous lateral rows of large, irregular black spots between limbs. Lateral areas of neck, insertion zones of limbs, posterior and anterior surface of limbs, dully grey. Ventral areas of head, body, limbs and tail (including regenerated portion) brilliant black, a few scales of femoral and interfemoral areas brightly yellow. Same color in preservative than live animals after two years but less vibrant. Yellow coloration disappears. 32 Zootaxa Magnolia Press AVILA ET AL.

6 FIGURE 2. Dorsal and ventral view of the preserved holotype (MACN 38985). Variation in paratypes. Based on the paratypes (Table 2, Fig. 3): In three males: SVL (mean ± SD (range)): 95.5 ± 10.9 ( ). Axilla groin distance: 40.6 ± 4.3 ( ). Head length: 20.2 ± 2.04 ( ). Head width: 17.2 ± 1.95 ( ). Foot length: 30.0 ± 0.8 ( ). Tibial length: 19.9 ± 1.0 ( ). Hand length: 28.8 ± 0.7 ( ). Midbody scales: 80.3 ± 2.0 (78 82). Dorsal scales: 75.0 ± 3.0 (72 78). Ventral scales: 111 ± 6.0 ( ). Precloacal pores: 3 4. Scales around interparietal: Supraoculars scales: 6 5. Scales around nasal 5 7. Supralabial scales: 7 8. Infralabial scales: 6. Fourth toe lamellae: Third toe lamellae: In six females: SVL (mean ± SD (range)): 95.9 ± 6.9 ( ). Axilla groin distance: 44.7 ± 4.5 ( ). Head length: 19.5 ± 1.0 ( ). Head width: 16.7 ± 0.8 ( ). Foot length: 27.7 ± 0.2 ( ). Tibial length: 18.5 ± 0.7 ( ). Hand length: 28.3 ± 0.9 ( ). Midbody scales: 77.4 ± 3.8 (72 82). Dorsal scales: 72.8 ± 2.1 (70 76). Ventral scales: ± 4.5 ( ). Precloacal pores not present in females. Interparietal usually irregularly shaped. Scales around interparietal: 7 8. Supraoculars scales: 6 5. Supralabial scales: 7 8. Infralabial scales: 5 6. Fourth toe lamellae: Third toe lamellae: As in other members of this complex, no strong body-size morphometric dimorphism or squamation differences were observed. The tail base of males is expanded laterally and cloacal opening is slightly quadrangular; at difference of other species of the group precloacal A NEW SPECIES OF LIOLAEMUS FROM PATAGONIA Zootaxa Magnolia Press 33

7 pores of males were small and inconspicuos. Dorsal coloration exhibits a strong variation between individuals. In some lizards head is completely black, meanwhile others exhibits almost any black scale and head color is light-tan or ochre. Some lizards exhibit only parts of the head black. Dorsal body coloration show similar variation, some lizards are almost completely dark brown or ochre with a pattern of large and rounded dorsolateral spots with different grades of fusion. In some individuals, this fussion is almost complete showing an irregular dorsolateral black band. MLP.S 2594 show complete light tan or ochre body coloration with small and irregular black spots irregularly distributed on dorsal areas, with some dark areas on head scales. Some lizards have gray lateral coloration in head, body, limbs, and tail. Ventral coloration varies from completely black to dark gray dotted with irregular rounded black spots. Tail coloration is plain, without rings, but varies from complete black, ochre, or gray, sometimes with small irregularly shaped and distributed black dots. A faded yellow coloration is restricted to the femoral and precloacal areas in a few individuals. TABLE 2. Morphometric and meristic variation in Liolaemus antumalguen type series. MACN MLPS 2594 MLPS 2595 LJAMM 6173 LJAMM 6172 BYU MACN MLPS 2592 sex F F F F F F M M M SVL Axilla-groin distance Head length Head width Head high Foot length Tibial length Arm length Midbody scales Dorsal scales Ventral scales Fourth toe lamellae Third toe lamellae Supralabial scales Infralabial scales Precloacal pores MLPS 2593 Etymology. The species name is in reference to a mithological fairy, Antú Malguén, wife of the sun, which according to a legend of the Mapuche people, inhabits the summit of the Domuyo Volcano. Geographic distribution. Liolaemus antumalguen is known only from its type locality in the eastern slope of Domuyo Volcano along Chadileu Creek, Chos Malal Department, Neuquén Province (Fig. 4, 5). It may be expected to occur in other localities around Domuyo Volcano and Cordillera del Viento mountains where habitats similar to the type locality exist but was not found in several field surveys carried out in surrounding lowlands and adjacent mountains. Cordillera del Viento is a higher mountainous range than the southern Andes in the region. The distributional area belong phytogeographycally to the Andean Dominio, High Andean District (Cabrera, 1976). The substrate is composed of between % rocky outcrops or rocky fields, with some soil in areas with some kind of protection from wind or flat areas. Liolaemus antumalguen appears to be restricted to this rocky zone that extends ~ 200 m on each side of the Chadileu Creek. Vegetation is dominated by bushes Mulinum spinosum, Chuquiraga oppositifolia, Anarthrophyllum rigidum, Adesmia rigida, Fabiana imbricata, intermixed with the grasses Festuca spp. and Stipa speciosa. 34 Zootaxa Magnolia Press AVILA ET AL.

8 FIGURE 3. Dorsal and ventral variation in color patterns of individuals of the type series. From upper to below: males, dorsal and ventral view, females, dorsal and ventral view. A NEW SPECIES OF LIOLAEMUS FROM PATAGONIA Zootaxa Magnolia Press 35

9 FIGURE 4. Type locality of Liolaemus antumalguen (red dot). Open circles identify main towns in the region. Yellow doted area: approximate extension of the Cordillera del Viento massif. Red dot-line mark boundaries between Mendoza and Neuquén province. Insert: the area in South America. Natural history. Liolaemus antumalguen was observed basking on rocks along a narrow zone on each side of Chadileu Creek. A few lizards were observed running or foraging between small plants along stream shores, but they usually retreated to seek refuge under bushes, rocks or crevices if a capture attempt was made. Liolaemus antumalguen is sympatric with other lizards of the genera Liolaemus and Phymaturus, but the majority of these species are still undescribed. Liolaemus punmahuida, a recently described lizard initially 36 Zootaxa Magnolia Press AVILA ET AL.

10 known only from the nearby Tromen Volcano, is syntopic in same areas (Avila and Perez, 2006), as well as Phymaturus verdugo (Avila et al., 2007b). At least two small species related to the elongatus and rothi clades of Liolaemus are also found in syntopy. With the exception of this undescribed species of the elongatus clade, no other species related with this clade are found sintopically with L. antumalguen, all other species of the clade are in lower altitudes or in separated mountains ranges. As was observed in other saxicolous Liolaemus, this new species seems to be territorial; males were observed defending areas in the rocks, making body displays, and fighting with neighbors (Perez, personal observation). No conclusive evidence of viviparity can be offered, but all related species have this reproductive mode. Difficulty of access to the type locality did not allow additional trips to this area to obtain more data about L. antumalguen. Remarks. A mtdna gene tree analysis, including the new described species as well as other related species and candidate species of the clade is depicted on Fig. 6. This tree is based on the mitochondrial gene fragment cyt-b (805 bp). We used JModeltest v0.1.1 (Guindon and Gascuel, 2003; Posada, 2008) to select the appropriate model of evolution (GTR+I+G). Two independent MrBayes analyses were run for 10 million generations, with Markov chains sampled at intervals of 4,000 generations. The equilibrium samples (after 10% of burn-in) were used to generate a 50% majority rule consensus tree, and posterior probabilities (Pp) were considered significant when 0.95 (Huelsenbeck and Ronquist, 2001). Liolaemus antumalguen is recovered within the elongatus clade that also includes L. chillanensis, L. thermarum and four candidate species (PP: 100%). The petrophilus and buergeri clades also include candidate species and are recovered with strong support (94% and 99% respectively); here we renamed the clade ceii-kriegi used by (Morando et al. 2003) as buergeri. The objective of this tree is to show the position of this new species in relation with these three related clades. Phylogenetic relationships for these clades are under study by our research group and a detailed analysis will be published elsewhere. Discussion The Liolaemus elongatus kriegi complex as defined by Cei (1979) on the basis of several diagnostic morphological characters includes widespread polytypic species for which species recognition rests on an inadequate taxonomy (Morando et al. 2003). Different taxonomic groupings for the species included in this complex have been proposed by Cei (1974, 1979, but also see Cei 1975, 1986, 1993), Ortiz (1981), Etheridge (1995), Espinoza et al. (2000), Schulte et al. (2000), Lobo (2001, 2005), Espinoza and Lobo (2003), Pincheira Donoso and Nuñez (2005), and Scolaro and Pincheira Donoso (2007). Morando et al. (2003) used the name Liolaemus elongatus-kriegi complex to be consistent with the original hypothesis of group content by Cei (1979), but resolved three well-supported species groups on the basis of mtdna sequences. The Liolaemus elongatus clade includes L. elongatus Koslowsky 1896, L. antumalguen described here, L. chillanensis (Torres-Perez et al. 2009) and L. thermarum Videla and Cei, 1996 (L. thermarum was originally described as a member of the neuquensis clade by these authors; but see Espinoza et al., 2000). We analyzed lizards from the type series of L. thermarum deposited in the MACN but they are in poor condition (completely black and deformed) as result of improper preservation techniques. Thus, we were not able to compare these specimens with other taxa, and no tissues were preserved for genetic analysis. We collected lizards in the type locality a few years later, and they match the photographs of the original description of Videla and Cei (1996), and those shown by Espinoza et al. (2000), allowing us to infer that this population corresponds to L. thermarum. As was suggested by Espinoza et al. (2000), we recovered this species as a member of the elongatus clade, and not closely related to L. neuquensis (in disagreement with Cei and Videla [2002, 2003] and Pincheira-Donoso and Scolaro [2007]). Also, in Morando (2004) in a phylogenetic tree including many species from the chiliensis group and several genes, L. neuquensis was not recovered as part of the elongatus clade. Other species such as L. leopardinus, L. ramonensis, L. valdesianus, L. curis, L.cristiani, as well as some subspecies of the L. monticola clade, may also be related to the Liolaemus elongatus clade (Lobo, 2001, 2005, Torres Perez et al. 2009), here we find L. chillanensis recovered within the elongatus clade, but a test of this hypothesis must await a more extensive study. In a recent publication, A NEW SPECIES OF LIOLAEMUS FROM PATAGONIA Zootaxa Magnolia Press 37

11 FIGURE 5. Upper: general view of the type locality of Liolaemus antumalguen from a southern perspective. Below: detail of the type locality rocky environment. Lizard activity was more intense around the rocks surrounding the Chadileu Creek. 38 Zootaxa Magnolia Press AVILA ET AL.

12 FIGURE 6. Phylogenetic relationships of Liolaemus antumalguen with other members of the elongatus-kriegi complex, based on Bayesian analyses; numbers above the nodes are posterior probability values. Lobo et al. (2010) includes as part of the elongatus clade, the following species: austromendocinus, elongatus, flavipiceus, gununakuna, parvus, petrophilus, punmahuida, thermarum, tregenzai, and all other species previously suggested as related with the elongatus clade are included in new created groups (leopardinus, chillanensis, monticola and kriegi) of the subgenus Liolaemus. Strong phylogenetic hypothesis for all these related groups are still missing and more geographic, taxonomic, and character sampling is still needed. Acknowledgments We thank Consejo Federal de Inversiones (CFI) for providing funds for field trips. Studies on the Liolaemus elongatus species are supported by fellowships to L. J. Avila and M. Morando from Consejo Nacional de Investigaciones Científicas y Técnicas (CONICET), and Fondo Nacional de Ciencia y Tecnica (FONCYT); and funds from the Biology Department, M. L. Bean Museum Life Science, and Kennedy Center for International Studies of BYU issued to J. W. Sites, Jr. We acknowledge the NSF Partnership for International Research and Education award (OISE ) for support of collaborative research on Patagonian biodiversity, granted to the following institutions (listed alphabetically): BYU, CENPAT, Dalhousie University, Darwinion Botanical Institute, Universidad Austral de Chile, Universidad Nacional de Cordoba, Universidad Nacional de Comahue, Universidad de Concepcion, and University of Nebraska. M. Archangelsky, J. C. Acosta, L. Belver, M. I. Christie, K. Delhey, K. Dittmar, N. Frutos, R. Kiesling, C. Perez, P. Petracci, and Y. Vilina helped on field trips and/or provided samples of the Liolaemus elongatus species. A NEW SPECIES OF LIOLAEMUS FROM PATAGONIA Zootaxa Magnolia Press 39

13 We thank Fauna authorities of Neuquén Province for collection permits. We thank scout H. Castillo and provincial park ranger D. Castillo for help in field work. References Abdala, C.S. (2002) Nuevo Liolaemus (Iguania: Liolaemidae) perteneciente al grupo boulengeri de la provincia de Neuquén, Argentina. Cuadernos de Herpetología, 16, Abdala, C.S. (2003) Cuatro nuevas especies del género Liolaemus (Iguania: Liolaemidae), pertenecientes al grupo boulengeri, de la Patagonia Argentina. Cuadernos de Herpetología, 17, Abdala, C.S. (2005) Dos nuevas especies del genero Liolaemus (Iguania: Liolaemidae) y redescripción de Liolaemus boulengeri (Koslowsky, 1898). Cuadernos de Herpetología, 19, Avila, L.J., Perez, C.H.F. & Morando, M. (2003) A new species of Liolaemus (Squamata: Iguania: Liolaemidae) from northwestern Patagonia (Neuquén, Argentina). Herpetológica, 59, Avila, L.J., Morando, M., Perez, C.H.F. & Sites, J.W., Jr. (2004) Phylogenetic relationships of lizards of the Liolaemus petrophilus group (Squamata, Liolaemidae), with description of two new species from western Argentina. Herpetológica, 60, Avila, L.J. & Perez, D.R. (2006) Reptilia, Iguania, Liolaemini, Liolaemus punmahuida: new geographic record. Checklist, 2, Avila, L.J., Morando, M. & Sites, J.W., Jr. (2006) Inferring evolutionary processes in inertial species an example in Patagonian lizards of the Liolaemus fitzingerii complex (Squamata: Liolaemidae). Biological Journal of Linnean Society, 89, Avila, L.J., Morando, M., Perez, C.H.F. & Sites, J.W., Jr. (2007) A new species of Liolaemus (Reptilia: Squamata: Liolaeminii) from southern Mendoza province, Argentina. Zootaxa, 1452, Avila, L.J., Perez, C.H.F., Perez, D. & Morando, M. (2007) Reptilia, Liolaemidae, Phymaturus verdugo: Distribution extension, new provincial records, filling gaps, and geographic distribution map. Checklist, 3, Cabrera, A.L. (1976) Regiones Fitogeográficas Argentinas. Enciclopedia de Agricultura y Jardinería (2ª ed.). Tomo II. Fase I. Acme. Buenos Aires. Cei, J.M. (1974) Revision of the Patagonian iguanids of the Liolaemus elongatus Complex. Journal of Herpetology, 8, Cei, J.M. (1979) The Patagonian Herpetofauna. In: Duellman, W.E. (Ed.), The South American Herpetofauna: Its origin, Evolution, and Dispersal. Museum of Natural History, University of Kansas, Lawrence, Kansas, pp Cei, J.M. & Videla, F. (2003) A new species of Liolaemus lacking precloacal pores in males from the Andean southeastern mountains of Mendoza Province, Argentina (Liolaemidae, Iguania, Lacertilia, Reptilia). Bolletino del Museo Regionale di Scienze Naturali di Torino, 20, Cei, J.M. & Videla, F. (2002) Singulares hallazgos evolutivos y taxonomicos en generos de iguanidos relevantes de la herpetofauna andina y de zonas limitrofes. Multequina, Latin American Journal of Natural Resources, 11, Cei, J.M. & Videla, F. (2003) A new Phymaturus species from Volcanic Cordilleran Mountains of the South-Western Mendoza province, Argentina (Liolaemidae, Iguania, Reptilia). Bolletino Museo Regionale di Scienze Naturali, Torino, 20, Espinoza, R.E., Lobo, F. & Cruz, F.B. (2000) Liolaemus heliodermis, a new lizard from northwestern Argentina with remarks on the content of the elongatus group (Iguania: Liolaemidae). Herpetologica, 59, Etheridge, R. & Christie, M.I. (2003) Two new species of the lizard genus Liolaemus (Squamata: Liolaemidae) from northern Patagonia, with comments on Liolaemus rothi. Journal of Herpetology, 37, Frost, D. (1992) Phylogenetic analysis and taxonomy of the Tropidurus group of lizards (Iguania, Tropiduridae). American Museum Novitates, 3033, Guindon, S. & Gascuel, O. (2003) A simple, fast and accurate method to estimate large phylogenies by maximumlikelihood. Systematic Biology, 52, Huelsenbeck, J.P. & Ronquist F. (2001) MrBayes: Bayesian inference of phylogeny. Bioinformatics, 17, Lobo, F. (2001) A phylogenetic analysis of lizards of the Liolaemus chiliensis group (Iguania: Tropiduridae). Herpetological Journal, 11, Lobo, F. (2005) Las relaciones filogeneticas dentro del grupo chiliensis (Iguania, Liolaemidae, Liolaemus): sumando nuevos caracteres y taxones. Acta Zoologica Lilloana, 49, Lobo, F., Espinoza, R.E. & Quinteros, S. (2010) A critical review and systematic discusión of recent classification proposals for liolaemid lizards. Zootaxa, 2549, Morando, M. (2004) Sistemática y filogenia de grupos de especies de los géneros Phymaturus y Liolaemus (Squamata: Tropiduridae: Liolaeminae) del oeste y sur de Argentina. Ph.D. Dissertation, Universidad Nacional de Tucumán, San Miguel de Tucumán, Argentina. 40 Zootaxa Magnolia Press AVILA ET AL.

14 Morando, M., Avila, L.J., & Sites, J.W., Jr. (2003) Sampling strategies for delimiting species: genes, individuals, and populations in the Liolaemus elongatus kriegi complex (Squamata: Liolaemidae) in Andean Patagonian South America. Systematic Biology, 52, Morando, M., Avila, L.J., Turner, C. & Sites, J.W. Jr. (2007) Molecular evidence for species complex in the Patagonian lizard Liolaemus bibronii and phylogeography of the closely related Liolaemus gracilis (Squamata: Liolaemini). Molecular Phylogenetics and Evolution, 43, Roig, F.A. (1998) La vegetación de la Patagonia. In: Correa, M.N. (Ed.). Flora Patagonica. INTA 7 1, Buenos Aires, Argentina, pp Pincheira-Donoso, D. & Nuñez, H. (2005) Las especies chilenas del genero Liolaemus Wiegmann, 1834 (Iguania: Tropiduridae: Liolaeminae). Taxonomia, sistematica y evolucion. Publicación Ocasional del Museo Nacional de Historia Natural 59, Chile, 486 pp. Pincheira-Donoso, D., Scolaro, J.A. & Schulte II, J.A. (2007) The limits of polymorphism in Liolaemus rothi: molecular and phenotypic evidence for a new species of the Liolaemus boulengeri clade (Iguanidae, Liolaemini) from boreal Patagonia of Chile. Zootaxa, 1452, Pincheira-Donoso, D. & Scolaro, J.A. (2007) Iguanian species-richness in the Andes of boreal Patagonia: Evidence for an additional new Liolaemus lizard from Argentina lacking precloacal glands (Iguania, Liolaeminae). Zootaxa, 1452, Pincheira-Donoso, D., Scolaro, J.A. & Sura, P. (2008) A monographic catalogue on the systematics and phylogeny of the South American iguanian lizard family Liolaemidae (Squamata, Iguania). Zootaxa, 1800, Schulte, J.A., II, Macey, J., R., Espinoza, R.E. & Larson, A. (2000) Phylogenetic relationships in the iguanid lizard genus Liolaemus: multiple origins of viviparous reproduction and evidence for recurring Andean vicariance and dispersal. Biological Journal of Linnean Society, 69, Smith, H.M. (1946) Handbook of lizards, Comstock Publishing Company. Ithaca, New York, U.S.A, 557 pp. Torres-Pérez, F., Mendez, M.A., Benavides, E., Moreno, R.A., Lamborot, M., Palma, R.E. & Ortiz, J.C. (2009) Systematics and evolutionary relationships of the mountain lizard Liolaemus monticola (Liolaemini): how morphological and molecular evidence contributes to reveal hidden species diversity. Biological Journal of the Linnean Society, 96, Videla, F. & Cei, J.M. (1996) A new peculiar Liolaemus species of the chiliensis phyletic group from the volcanic Cordilleran landscapes of southern Mendoza Province, Argentina (Iguania, Lacertilia, Reptilia). Bolletino Museo Regionale di Scienze Naturali di Torino, 14, Appendix I Specimens examined Liolaemus antumalguen (10). ARGENTINA: NEUQUEN: Chos Malal Department, eastern piedmont of Domuyo volcano, around Chadileu Creek: MACN to 87, MLP.S 2592 to 5, LJAMM-CNP , BYU Liolaemus buergeri (12). ARGENTINA: MENDOZA: Malargue Department, 16 km W Las Leñas: LJAMM-CNP 2682, Mallines Colgados: LJAMM-CNP , NEUQUEN: Ñorquin, Cascada del Rio Agrio: LJAMM-CNP Liolaemus ceii (14). ARGENTINA: NEUQUEN: Alumine Department, Provincial Road 13, Pampa de Lonco Luan: LJAMM-CNP , 1198, Liolaemus elongatus (128). ARGENTINA: CHUBUT: Cushamen Department: Ruta Provincial 12, 9.1 km E La Cancha Railroad Post, road to Gualjaina: LJAMM-CNP Futaleufu Department: Ruta Nacional 40, Km 1530, 17 km S Esquel, 5 km S junction Ruta Nacional 40 and Ruta Nacional 259: LJAMM-CNP 2128, 2156/7, Ruta Nacional 40, Km 1589, 18 km N Tecka: LJAMM-CNP Languineo Department: Ruta Nacional 25, 5 km W Colan Conhue, Cuesta del Paisano: LJAMM-CNP 6177 to 80. Rio Senguer Department: Ruta Provincial 20, 23 km W Los Manantiales: FML13070, LJAMM-CNP 3046/7. Tehuelches Department: Ruta Nacional 40, 22 km S Gobernador Costa: FML 13071, LJAMM-CNP Ruta Provincial 53, 40 km S junction Ruta Nacional 25: LJAMM-CNP 4681 to 83. Near Gobernador Costa: LJAMM-CNP 6145/6. NEUQUEN: Alumine Department. Portal La Atravesada, 3 km S, 7 km W Primeros Pinos: MVZ /5. 3 km ENE Lago Ruca Choroi, 8 km E, 9 km N Cerro Ruca Choroi: MVZ Along Arroyo Rucaco, SE end of Lago Ruca Choroi, 3.5 km E and 6.5 km N Cerro Ruca Choroi: MVZ /3. Catan Lil Departament. Campo de la Pistola, 4 km W and 2 km N Las Coloradas: MVZ /67, /70, /3/4. On E side of Rio Alumine, 31 km S Rahue via Ruta Provincial 23: MVZ Lacar Department. Pampa de Alicura on Ruta 40 and 237, 5 km W and 6 km N Paso Flores: MVZ Los Lagos Department, on W side of Rio Limay, 2 km E and 16 km S Confluencia: MVZ Sandy flat along highway, Estancia Tehuel Malal, ca. 6 km WNW Nahuel Huapi: MVZ km S, 3 km W Cerro de las A NEW SPECIES OF LIOLAEMUS FROM PATAGONIA Zootaxa Magnolia Press 41

15 Ardillas: MVZ Zapala Department. Ruta Provincial 13, 12 km W, 1 km N Estacion Zapala: MVZ /4/ 7/8. SW end of Laguna Blanca, 8.5 km W and 1 km N Cerro Mellizo Sud: MVZ /8. W end of Laguna Blanca: MVZ Ruta Provincial 46, 0.5 km N limite PN Laguna Blanca: MVZ RIO NEGRO: Bariloche Deparment. Ridge above Refugio Neumeyer, 15 km S Bariloche: MVZ /90/81/82. Chalhuaco Valley: FML 13072/3, LJAMM-CNP 2811, 3051/2, 2055 to 57, 3051/2, 3055 to 7. El Cuy Department: Ruta Provincial 67, 19.2 km NE Mencue: LJAMM-CNP 5532 to 57. Ruta Provincial 67, 20 km S Mencue: LJAMM 5559/60. Ñorquinco Department. Along Rimrock, 4 km S and 1 km E Alto del Escorial: MVZ , , Along Rio Chenqueniyen, 10 km E and 3 km S Cerro Pico Quemado: MVZ to 34, to 41. Laguna de Los Juncos, Escorial de Chenqueniyen, 5 km N Alto del Escorial: LJAMM-CNP /9. Ruta Provincial 6, 1 km NW Ojo de Agua: LJAMM-CNP Pilcaniyeu Department. Rocky knoll, Cañadon Bonito, 23 km NE Pilcaniyeu: MVZ /8. Ruta Provincial 23, 2.3 km SE Comallo: LJAMM-CNP Ruta Provincial 23, 4.8 km SE Comallo: LJAMM 5449/50. Ruta Nacional 40, 2.7 km S Estancia San Pedro: LJAMM- CNP 5428 to Ruta Provincial 67, 2 km N Cañadon Chileno, 37 km NE Comallo: LJAMM-CNP Ruta Provincial 67, 3.5 km 3.5 km N Cañadon Chileno: 5621 to 26. Veinticinco de Mayo Department: Ruta Provincial 8, 17 km S San Antonio del Cuy: FML 13060/1, LJAMM-CNP 1519/20, 1615, 1636/7, 1639 to 1641, 1818 to 22, 1915, 2433 to 36, 2721, Ingeniero Jacobacci: LJAMM-CNP 263/4, 269/70, 278, Liolaemus flavipiceus (4). ARGENTINA: MENDOZA: Malargüe Department: Provincial Road 145, Paso Pehuenche: LJAMM-CNP Liolaemus kriegi (22). ARGENTINA: RIO NEGRO: Ñorquinco Department: Provincial Road 6, 1 km NW Ojo de Agua: LJAMM-CNP , , National Road 1s40, 2.5 km N Chenqueniyen: LJAMM-CNP , 3501, de Mayo Department: Provincial Road 76, 57 km S Ingeniero Jacobacci: LJAMM-CNP 3044, 3071, , Provincial Road 5, 40 km SE Maquinchao. LJAMM-CNP 3045, , Liolaemus thermarum (27). ARGENTINA: MENDOZA: Malargüe Department: Provincial Road 226, 1 km N Baños del Azufre: LJAMM-CNP , 5786 to Provincial Road 226, 11.4 km S Baños del Azufre: LJAMM-CNP 5799 to Provincial Road 226, Paraje Mallines Colgados: LJAMM-CNP 2748 to San Carlos Department: Along Rio Diamante, 3 km S Lago Diamante: MVZ ; S side Laguna Diamante: MVZ to Specimens used for phylogenetic analyses Liolaemus elongatus-kriegi complex: -buergeri clade: L. aff. austromendocinus: 2244; L. buergeri NQN: 5796; L. sp. nov. A: 2533; L. kriegi: SDSU 3695; L. aff. buergeri 5297; L. kriegi: fn elongatus clade: L. antumalguen: 6155 ; L. sp. nov. 7: 2602; L. sp. nov. 6: 2454; L. chillanensis: 3434; L. sp. nov. 15: 5268; L. elongatus: 2128; L. sp. nov. 16: 5238 ; L. thermarum: 5792; L. sp. nov. 19: petrophilus clade : L. austromendocinus MZ: 5147; L. sp. nov. 48: 540; L. parvus: 2711; L. gununakuna: 2690; L. sp. nov. 12: 5375; L. sp. nov. 13: 5356; L. petrophilus CH: 2125; L. petrophilus RN: 1914; L. capillitas: 2786; L. umbrifer: 5029; L. tulkas: 4227; L. dicktracy: 5750; L. talampaya: punmahuida clade: Liolaemus flavipiceus: 7906; L. punmahuida: Outgroup: L. kingii : All LJAMM-CNP collection. 42 Zootaxa Magnolia Press AVILA ET AL.