Liolaemus puelche, Avila, Luciano Javier, Morando, Mariana, Perez, Cristian Hernan Fulvio & Sites, Jack W., 2007

Avila, Luciano Javier, Morando, Mariana, Perez, Cristian Hernan Fulvio & Sites, Jack W., 2007, A new species of Liolaemus (Reptilia: Squamata: Liolaemini) from southern Mendoza province, Argentina, Zootaxa 1452, pp. 43-54 : 45-52

Liolaemus puelche sp. nov.

( Figure 1 )

Type Material. Holotype: MACN 38992, adult male collected along the south side of Ruta Nacional 40, 3 km N Ranquil Norte (36º 38’ S, 69º 49’ W, 1600 m), Malargüe Department, Mendoza province, Argentina, 9 March 2000, by L. J. Avila, and M. Morando.

Paratypes: MLP.R 5063, adult male; MLP.R 5064, subadult male; MACN 38991, subadult female. All collected on marginal slopes on southern edge of Ruta Nacional 40, 3 km N Ranquil Norte (36º 38’ S, 69º 49’ W, 1600 m), Malargüe Department, Mendoza Province, Argentina, 7 February 2003, by L. J. Avila, K. Dittmar, M. Morando, and C. H. F. Perez.

Diagnosis. Liolaemus puelche is a robust and medium size member of the clade of Liolaemus lizards referred to as the boulengeri group by Etheridge (1995) and is a member of the Liolaemus donosobarrosi group, that includes L. cuyanus , L. donosobarrosi , L. josei , and several other potential species still undescribed ( Avila et al, 2006; Morando et al, 2004). Liolaemus cuyanus has a light brown dorsal coloration crossed by well defined transversal brown bands with posterior white-borders, a conspicuous brown to black antehumeral band from throat to shoulder, and white ventral areas; these characteristics are not present in L. puelche . Liolaemus mapuche has a very different dorsal coloration characterized by a light blue head and scattered blue scales on a light green-blue background, a black antehumeral arch, four series of gray paravertebral and lateral spots, and scattered yellow and blue scales in the lateral area of the body ( Abdala, 2002); these characteristics are never present in L. puelche . Liolaemus donosobarrosi is a smaller species (maximum SVL = 64 vs 89 mm in L. puelche ), and has a dorsal pattern consisting of transverse series of well-marked four half-moon spots surrounding white spots, with a background coloration orange brown or orange tan, sometimes very bright, more scales around midbody (71–87 vs 67–76 in L. puelche ), and more scales between the occiput and rump (82–101 vs 74–80 in L. puelche ). Liolaemus josei is a smaller species (maximum SVL = 73.1 vs 89 mm in L. puelche ), with a marked sexual dichromatism not present in L. puelche ; L. josei males have gular melanism, two series of well marked paravertebral black spots, and a dorsal background of white/ light blue speckles and a light-blue sheen, characters never present in males of L. puelche .

Description of the holotype. Adult male 85.5 mm (SVL), regenerated tail 62.0 mm (46.0 mm regenerated portion). Axilla-groin distance 37.4 mm. Head length 17.1 mm; head width 14.9 mm; head depth 10.6 mm; snout length 6.2 mm, horizontal diameter of the orbit 5.0 mm. Arm length 23.2 mm; tibial length 15.9 mm; foot length 23.5 mm.

Upper head scales smooth, convex, bulged, pitted with scale organs, in postrostral, internasals, frontonasals, and prefrontals. Rostral pentagonal, twice as wide as high (2.9 x 1.3 mm). Two postrostrals, wider than high, with four conspicuous scale organs each; together with anterior supralabials separate nasal scales from rostral. Nasal scales subpentagonal in shape (1.6 x 1.6 mm). Nostril roughly rhomboidal in shape, occupying almost two thirds of the nasal scale. Nasal scales in contact with seven scales on each side. Internasal scales convex. Six internasal, two median, almost quadrangular; four lateral in tandem, a smaller one in front and a long and narrow scale in back. Two frontonasals rows in front, two paired scales in wider contact in medial position, two smaller lateral scales in contact with nasals. Along the back, a row of five scales, three about equal in size and subpentagonal in medial position; lateral frontonasals rhomboidal equal in size each other, and in wide contact with second canthal. Nine prefrontals, anterior three regularly arranged, with a small medial and two lateral more that three times larger; a second row of three scales with a medial scale larger than laterals; a third row with two scales in tandem in the left side, and a single scale in the right side. Frontal scale irregular, slightly larger that last prefrontals. Seven frontoparietals irregular in shape, smaller that frontal and parietal. Interparietal pentagonal with a large and conspicuous white cream “eye” in the middle.

A lateral parietal on each side, irregular, equal or slightly smaller than interparietal. Four larger parietals in back, very irregular. Circumorbitals: 11–11. Supraoculars 8–7, laterally expanded. Three rows of irregular, small scales between supraoculars and supercilliaries, 26 on each side. A small scale between nasal and first canthal. First canthal higher than wide. Posterior canthal longer than wide, with a well marked but blunt ridge. Posterior canthal overlap only a small part of first supercilliary. Supercilliaries 8–7 (left-right), strongly overlapped. Loreal irregular in size and shape, 9-5; together with preocular and anterior subocular forming a slight concavity. Upper ciliary scales in two rows, those of inner rows flat and quadrangular, those of outer row rectangular, compressed, and moderately projecting. Lower and upper ciliaries similar in size and shape. Palpebral scales small, irregulars, flat. One preocular, longer than wide; one elongate subocular (5.6 x 0.9 mm), one small postocular; a very evident keel in subocular and postocular, less marked in preocular. Lorilabials convex, 7-7, except the first, roughly quadrangular, higher than supralabials pitted with conspicuous and numerous scale organs. Supralabials 9-8, flat. Temporal scales smooth, swollen, juxtaposed, with a scale organ in the tip. Auditory meatus higher than wide (3.2 x 1.0 mm) surrounded by granular scales. Mental pentagonal, wider than high (3.3 x 1.9); in contact with anterior infralabial and postmental but not in contact with anterior sublabials. Infralabials 6-6. Chinshields 7-7, transversally expanded, separated from infralabials by series of 2–3 elongated but smaller sublabials. Larger sublabials with several conspicuous scales organs each, less numerous to absent in smaller scales. Only a few scale organs present in supralabials and infralabials. Gular scales smooth, flat, imbricate, rhomboidal, immediately after postmental, gradually becoming rounded posteriorly and with a distinct posterior apical notch. Lateral neck folds (longitudinal, oblique, antegular, gular, antehumeral and post auricular) distinct to well developed.

Dorsal body scales subtriangular to obovate, moderately imbricated. Twenty three to 25 longitudinal rows with scales with distinct but blunt keels, a few with an apical scale organ. At midbody, dorsal scales grade laterally into smaller scales, smooth, imbricate, many with an apical notch. Scales anterior to, above, and posterior to forelimb and hind limb insertions, small, smooth, non-overlapping, becoming granular close to the insertions. Ventral body scales smooth, flat, imbricate, with an apical notch; subtriangular to obovate, same size or smaller than dorsal body scales. Scales around midbody 71; scales between occiput an anterior margin of hind-limb articulations 80. Scales of cloacal apron slightly smaller in size than ventral body scales. Precloacal pores 8.

Suprabrachials smooth, rhomboidal, imbricated, with a small notch in the tip. Infrabrachials small, convex, and not overlapping. Supra-antebrachials smooth, imbricate, rhomboidal to obovate, some with a small notch in the tip, others larger than suprabrachials. Infra-antebrachials smooth, granular to progressively rhomboidal distally, becoming with one to three small but distinct keel and spines near the hand insertion. Supracarpals smooth, strongly imbricate. Infracarpals strongly keeled, imbricate. Supradigital lamellae smooth, imbricate. Subdigital lamellae tricarinate, imbricate, numbering: I: 14, II: 22, III: 22, IV: 16, V; 10.

Suprafemorals smooth, imbricate, rhomboidal to rounded, few with an apical scale organ or a blunt keel. Infrafemorals smooth, imbricate, rhomboidal. Supratibials smooth to slightly keeled, imbricate, becoming rounded distally. Twenty to 25 spinose scales in a postfemoral patch. Infratibials smooth, imbricate. Supratarsals, smooth, imbricate, a few with a apical notch. Infratarsals strongly imbricate, with a conspicuous keel. Supradigital smooth, imbricate. Infradigital bi- to tri-carinate, numbering I: 13, II: 18, III:25, IV: 33, V: 19. Caudal scales smooth, a few slightly keeled, imbricated.

Coloration. In life, general dorsal background color tan. Dorsal head surface dark brown, grading to light tan in azygous and nuchal areas. Between occiput and rump, twelve irregular, transversal bands formed by four dark brown spots. Paravertebral spots more-or-less round, with a concave or notched posterior margin, distinctly outlined with white scales, the anterior margins indistinct. Lateral spots irregularly rectangulars, bordered with white or light tan scales, sometimes partially fused ventrally. Paravertebral spots are smaller but more marked than quadrangular lateral spots. Spots become less evident and apparently fused on neck and rump areas. On the tail, paravertebral spots become rounded and progressively fused, disappearing on the regenerated portion; lateral spots become rounded and lighter, and disappear in the first portion of the tail. On dorsal limbs areas an irregular pattern of rounded, irregular, sometimes fused dark spots on a light tan background. Ventral surfaces uniformly bluish gray. Immediately after capture, central body area with a light bright yellow dorsal coloration that disappeared after a few hours. Light yellow coloration on ventral areas of hindlimbs and cloacal apron faded after a few weeks in preservative. After several years in preservative, all coloration became faded.

Variation. Based on the paratypes (Table 2, Figure 2 ); in subadult–adult males: SVL 69–89 mm. Axilla groin distance 30.5–40.1 mm. Foot length 20.5–24.2 mm. Tibial length 14.9–18.3 mm. Arm length 19.6–24.1 mm. Head length 14.6–17.7 mm. Head width 11.9–15.3 mm. Head depth 8.7–10.9 mm. Midbody scales 67– 76. Dorsal scales between occiput to rump 74–78. Ventral scales 100–106. Supralabials 9. Infralabials 6. Fourth toe lamellae 29–30. Precloacal pores 6–9. In one subadult female: SVL 74.0 mm. Axilla-groin distance 35.1 mm. Foot length 21.5 mm. Tibial length 14.5 mm. Arm length 21.5 mm. Head length 14.7 mm. Head width 12.4 mm. Head depth 9.1 mm. Midbody scales 69. Dorsal scales between occiput to rump 75. Ventral scales 103. Supralabials 8. Infralabials 6. Fourth toe lamellae 28. Two small precloacal pores. Dorsal head scales variable in size, some irregular in shape, most convex but a few almost flat. Cloacal apron larger in males that in females. Scales of the cloacal apron slightly smaller that ventral scales in males, same size or slightly smaller in females. Postfemoral patch not present in females and variable in size in males between 18–25 scales. Only in a single lizard were 6 scales observed in contact with the mental scale; all others have 4 scales.

Dorsal background color brown to tan, recently captured adult males have a yellowish-green sheen. Dorsal head scales with brown, tan and white speckles. Most individuals boldly marked with holotype pattern, maintaining the pattern of transversal bands with variations in spots shape and distinctiveness. Variation more frequent on dorsal neck area by fusion of spots. Ventral surfaces of subadults tinged with gray, throat with light gray spots and light gray variegations.

Etymology. The specific name puelche , comes from the name of a group of aboriginal people that inhabited the mountain slopes and piedmont between the Barrancas and Diamante rivers, in southern Mendoza Province. Puelche culture apparently started to disappear two centuries ago when they mixed with other tribes invading from actual Chile.

Geographic distribution. Liolaemus puelche was collected only in an isolated locality along the National Road 40, north to the locality of Ranquil Norte, Malargüe Department, Mendoza Province, in western Argentina ( Figure 3 , 4 ).

Natural history. The holotype and the paratypes were found by active search, basking in the edges of spinose shrubs ( Schinus johnstonii ); the first lizard collected was found running from one shrub to another, but the others were usually found motionless in the bare ground below the vegetation branches where they blend in with patterns of light and shadows on the soil. When pursued they run to mammal burrows ( Ctenomys sp.) which are common beneath scattered bushes and shrubs of the area. Population density of this species in the area appears to be very low because no more than four individuals collected were found in two collecting trips, while other sympatric lizards were commonly found at the same time. Apparently suitable habitats for this species are scarce in the small area that is accessible by motorized vehicle. This species shares its habitat with other Liolaemus species ( L. austromendocinus , L. bibronii , L. darwinii ) and usually occupies the microhabitat below the spiny shrubs that grow as scattered “island clumps” of 5–10 m 2 separated by an open, loose sand substrate, while the other species were found in rocky patches or in the rocky-gravel accumulations of the road edges. Only a male of L. darwinii was observed sharing the same microhabitat of L. puelche . No data about reproduction or diet area available, but as in other related species, L. puelche is probably oviparous and feeds mainly in arthropods and some plant matter.

Remarks. A combined phylogenetic analysis using Maximum Parsimony, Maximum Likelihood and Bayesian Analyses of 141 species of Liolaemini with three mitochondrial (cytb, ND4, 12S = 2536bp) and two nuclear genes (GAPDH, C-mos = 834bp), recovered L. puelche as nested inside an arenicolous group of the donosobarrosi group (Morando, 2004). Here we reproduce the topology of the donosobarrosi group, where L. puelche (L. sp. 25, Fig. 5.2; Morando 2004) was recovered as the sister taxon of L. josei (L. sp. 24, Fig. 5.2; Morando 2004). This topology suggests that the group is still poorly known, and has at least six more “candidate species” ( Morando et al., 2003), including two in the cuyanus complex, two related to L. josei and L. puelche , and another two closely related to L. donosobarrosi . Although we do not have molecular data for L. mapuche, Abdala (2002) suggested that it is closely related to L. cuyanus ; if true, then, the donosobarrosi group includes five described and possibly seven undescribed species. These are under study by our research group and a detailed analysis of this group will be published elsewhere.

Figures

FIGURE 1. Liolaemus puelche, holotype adult male (MACN 38992), from south side of Ruta Nacional 40, 3 km N Ranquil Norte, Malargüe Department, Mendoza province, Argentina.

FIGURE 2. Dorsal and ventral color variation in the type series of Liolaemus puelche.

FIGURE 3. Type locality of Liolaemus puelche (black dot). Grey lines show main national and provincial roads, and open circles identify main towns in the region. Insert: the area in Argentina.

FIGURE 4. General view of the type locality of Liolaemus puelche from a northern perspective. Lizard activity was more intense under the large shrubs of Schinus johnstonii.