Liolaemus tirantii, Avila & Perez & Minoli & Medina & Sites & Morando, 2017

Avila, Luciano Javier, Perez, Cristian Hernán Fulvio, Minoli, Ignacio, Medina, Cintia Debora, Sites, Jack W. & Morando, Mariana, 2017, New species of Liolaemus (Reptilia, Squamata, Liolaemini) of the Liolaemus donosobarrosi clade from northwestern Patagonia, Neuquén province, Argentina, Zootaxa 4362 (4), pp. 535-563 : 543-549

Liolaemus tirantii sp. nov.

1996 Liolaemus donosobarrosi Avila, 1996 Cuadernos de Herpetología 9(2):109–110.

2006 Liolaemus donosobarrosi Avila et al. , Biological Journal of the Linnean Society 89(2):241–275.

Holotype. LJAMM-CNP 1 3464 ( Figs. 6 , 7 ), an adult male from a small unnamed sedimentary hill 2 km S Cerro Bandera, 1 km S National Road 22, 18.5 km W Cutral-Có, Zapala Department , Neuquén Province, Argentina (38º 58’ 06.0” S, 69º 27’ 08.7” W, 970 m), collected by J.A. Avila, I. Minoli, C. Medina and M. Kozykariski, 12 November 2010.

Paratypes. LJAMM-CNP 13465 (male),13467–13469 (females) from same locality as holotype, collected by J.A. Avila, I. Minoli, C. Medina and M. Kozykariski, 12 November 2010 . MLP. S 2625 (male), BYU 52427–52429 (juveniles), eastern outskirts La Amarga village , on the road to Petrified Forest, Zapala Department, Neuquén Province, Argentina (39º06’39.7” S, 69º34’29.2” W, 803 m), collected by L.J. Avila, M. Morando, D. Perez and J. Perez, 2 May 1998 .

LJAMM-CNP 2542 (female) from 2 km SE La Amarga village , Zapala Department, Neuquén Province, Argentina (39º06’39”S, 69º34’09”W), collected by L.J. Avila and M. Morando, 7 March 2000 . LJAMM-CNP 10409 (female), from 2 km E La Amarga village , 11 km E junction Provincial Road 34, Cerro Atravesado, Zapala Department, Neuquén Province, Argentina (39º06’39.70” S, 69º34’9.20” W, 803 m), collected by L.J. Avila, M. Morando and D. Perez, 26 February 2008 . LJAMM-CNP 11292–11293 , 11339 (females), from 2 km E La Amarga village , 11 km E junction Provincial Road 34, Zapala Department, Neuquén Province, Argentina (39º06’39.70”S, 69º34’9.20”W, 803 m), collected by L.J. Avila, N. Feltrin, M. Nicola, and L.E. Martinez, 17 December 2008 . LJAMM-CNP 11297 (female) from National Road , 1 km W junction Provincial Road 34, Cerro Atravesado, Zapala Department, Neuquén Province, Argentina (38º55’14”S, 69º42’13.2”W, 788 m), collected by L.J. Avila, N. Feltrin, M. Nicola, and L.E. Martinez, 17 December 2008 . LJAMM-CNP 247 (male), 248 (female) from 40 km E Zapala Yacimiento Cerro Bandera , Zapala Department, Neuquén Province, Argentina (38º57’22.47” S, 69º33’11.50” W, 811 m), collected by S. Tiranti.

Diagnosis. Liolaemus tirantii sp. nov. is a robust but small sized member of the clade of Liolaemus referred to as the boulengeri group by Etheridge (1995), and is a member of the Liolaemus donosobarrosi group ( Avila et al. 2006, 2007, Olave et. al 2014, 2016) that includes L. cuyanus , L. donosobarrosi , L. mapuche , L. josei , L. puelche and several other potential species still undescribed. Liolaemus tirantii sp. nov. can be distinguished from all other species of its group of Liolaemus by a combination of genetic and morphological characteristics.

Liolaemus tirantii sp. nov. can be differentiated from L. cuyanus by its smaller size (64.3 mm vs 92.5 mm maximum SVL), higher number of midbody scales (X = 73.5 vs X = 66.7; min.–max. 66–86 vs 60–76, Blum 2012) with overlap, higher number of dorsal scales (X = 87.2 vs 63.6; min.–max. 79–97 vs 57–74, Blum 2012) with no overlap, four scales in contact with mental vs six in L. cuyanus , lack of prescapular marks and ventral melanism, and a very different general coloration and coloration patterns.

Liolaemus tirantii sp. nov. is smaller than Liolaemus mapuche (64.3 mm vs 82.7 mm maximum SVL), has higher number of dorsal scales (X = 87.2 vs 76.2; min.–max. 79–97 vs 70–86) with some overlap, only four scales in contact with mental vs six in L. cuyanus (more common state, Abdala 2002), and lack of scapular marks, ventral melanism and a very different dorsal coloration characterized by a light blue head and scattered blue scales on a light green-blue background ( Abdala 2002).

Liolaemus tirantii sp. nov. is smaller than L. puelche (64.3 mm vs 89 mm maximum SVL), has a higher number of midbody scales (X = 73.5 vs 70.7; min.–max. 66–86 vs 67–76) with overlap and a higher number of dorsal scales (X = 87.2 vs 76.7; min.–max. 79–97 vs 74–80) with little overlap.

Liolaemus tirantii sp. nov. is slightly smaller than L. josei (64.3 mm vs 73.1 mm maximum SVL), has a higher number of midbody scales (X = 73.5 vs 69.1; min.–max. 66–86 vs 62–76), have higher number of dorsal scales (X = 87.2 vs X = 73.1; min.–max. 79–97 vs 67–81) with slight overlap, lacks the well-marked gular melanism of L. josei males as well as its more marked dorsal color pattern.

Liolaemus tirantii sp. nov. has a lower average number of scales around midbody than L. donosobarrosi (X = 73.5 vs 82.1; min.–max. 66–86 vs 74–89), middorsal scales (X = 87.21 vs 90.4; min.–max. 79–97 vs 86–101), and ventral scales (X = 94.0 vs 99.5; min.–max. 87–107 vs 88–109). Dorsal color pattern in L. tirantii sp. nov. is a series of rounded to kidney-shaped transversal marks bordered of white, usually fused along the body but more frequently on the limbs insertions, giving it a ¨wavy¨ appearance, whereas L. donosobarrosi has a series of halfmoon dots surrounding white spots, only in a few cases fused to each other, usually in the dorsal neck region; L. tirantii sp. nov. lacks of the black transversal marks observed on the head sides of L. donosobarrosi .

Liolaemus tirantii sp. nov. can be distinguished from L. donosobarrosi sp. 3 by its lack of a transverse yellow and brown bands along the dorsum in life, and presence of a series of rounded to kidney-shaped transversal marks bordered of white, usually fused along the body but more frequently on the limbs insertions, given it a ¨wavy¨ appearance.

Description of holotype. Adult male ( Fig. 6 , 7 ) 64.1 mm snout-vent length (SVL); tail length 97.4 mm. Axilla-groin distance 30.0 mm. Head length 13.3 mm (from anterior border of tympanum to tip of snout), 11.8 mm wide (at anterior border of tympanum), 7.9 mm high (at anterior border of tympanum). Snout length 4.2 mm (posterior margin of canthal to tip of snout). Interorbital distance 5.7 mm. Eye-nostril distance 2.5 mm. Orbitauditory meatus distance 4.2 mm. Forelimb length 17.1 mm. Tibial length 11.6 mm. Foot length 16.9 mm (ankle to insertion of claw on fourth toe). Dorsal head scales something bulged, smooth, 13 between occiput at level of anterior border of tympanum to rostral, pitted with numerous scale organs in the anterior region, and reducing to a single organ, or lack, in the posterior half of the head. Rostral scale wider (2.7 mm) than high (0.9 mm). Two postrostrals, together with anterior lorilabial, separate nasal scales from rostral. Nasal scales longer than wide, irregularly hexagonal; nostril one-half length of nasal, posterior in position. Scales surrounding nasals 8–7. Four elongated internasals. Frontonasals three, irregularly pentagonal. Prefrontals 5, a small pentagonal scale in the center (0.7 mm), and two pairs scales: a pair dorso-lateral larger scales, irregularly hexagonal (1.9 mm), and a pair lateral medium-sized scales (1.4 mm), approximately pentagonal. Three frontal scale, two anterior scales and one posterior scale. Interparietal pentagonal, surrounded by 7 scales; four smaller and irregular in front and sides, two larger in back. Parietal eye evident. Parietals some bulged, irregularly shaped, largest in size to interparietal (2.0 vs. 1.2 mm). Circumorbitals: 12–13. Transversally expanded supraoculars 6–7. Smaller lateral supraoculars: 18–17. Anterior canthal wider than long, separate from nasal by one postnasals. Posterior canthal longer than wide. Loreal scales bulged (3–5). Lorilabials longer than wide (9–7). Superciliaries eight on each side, flattened and elongated, anterior five broadly overlapping dorsally. Orbit with sixteen upper on each side and 16–14 lower ciliaries. Orbit diameter 3.4 x 1.8 mm. Preocular small, unfragmented, longer than wide. Subocular scale elongated, nine times longer than wide (4.1 x 0.6 mm). A well-marked longitudinal ridge along upper margin of preocular and subocular scales. Postocular small, slightly bulged, quarter superimposed to subocular, with a marked longitudinal ridge. Palpebral scales small granular and bulged. Supralabials 6–8, convex. Temporals smooth, convex, juxtaposed with one scale organ in the tip. Anterior auriculars smaller than adjacent posterior temporals, three slightly projecting outward. Posterior auriculars small and granular. External auditory meatus conspicuous, higher (2.0 mm) than wide (1.3 mm). Lateral scales of neck granular with inflated skin. Mental scale wider (2.2 mm) than high (1.3 mm), in contact with four scales. Mental followed posteriorly by two postmentals, and two rows of four chinshields on each side. Six 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, with melanophores. Scales of throat between chinshields slightly juxtaposed, becoming slightly imbricate toward auditory meatus. Thirty-four gulars between tympanum openings. Infralabials separated from chinshields by one to three rows of scales. Antehumeral well developed; longitudinal neck and postauricular distinctive; gular incomplete; rictal, dorsolateral, oblique unconspicuous.

Scales of dorsal neck region rhombals, imbricate, smooth. Eighty-nine dorsal scales between occiput and anterior surface of thighs. Dorsal body scales rhomboidal, imbricate, smooth to very slightly keeled towards the posterior half of body. Dorsal scales grade laterally into slightly smaller, smooth scales at midbody. Scales immediately anterior and posterior to forelimb and hindlimb insertion small, smooth, granular, and nonoverlapping. Body lateral scales grading smaller to larger at midbody. Ventral body scales rhombals, smooth, flat, imbricate, larger than dorsal scales. Seventy-one scales around midbody; scales between mental and precloacal pores 95. Scales of cloacal region about equal in size to ventral body scales. Eight conspicuous precloacal pores.

Anterior suprabrachials rhomboidal, imbricate, smooth, slightly larger to dorsal body scales. Postabrachials smaller, smooth, becoming granular near axilla. Supra-antebrachials similar to suprabrachial. Infra-antebrachials rhombals, imbricate, and smooth. Supracarpals imbricated, rhomboidal, smooth. Infracarpals strongly imbricate, rhomboidal, keeled, and slightly mucronate. Subdigital lamellae with three keels, each terminating in a short mucron, numbering: I: 9, II: 14, III: 18, IV: 19, V: 9. Claws robust, curved and sharp, opaque brown.

Suprafemorals as large as dorsal body scales, rhomboidal, imbricated, smooth. Postfemorals small, granular shape. Supratibial rhomboidal, imbricated, smooth, some very slightly keeled. Infrafemorals scale small, granular and smooth with enlarged and bulged scales patch, with thirty-six scales (femoral patch), mucronate. Supratarsal rhombals imbricated and smooth. Infratarsal small, rhomboidal, imbricate, smooth. Subdigital scales keeled, each terminating in 1–3 short mucrons, numbering: I: 10, II: 14, III: 18, IV: 23, V: 14. Claws robust, curved and sharp, opaque brown. Tail complete, non-regenerated. Dorsal and lateral caudal scales, quadrangular and moderately keeled. Ventral subtriangular and smooth.

Coloration. In life, general dorsal background color on head, neck, trunk, tail, and limbs Robin Rufous (30); ventral background coloration Cream White (52), in both cases at sunlight. Dorsal head surface uniform with some Cream White (52)/Burnt Umber (48) spots irregularly distributed on some scales, becoming more numerous on the nuchal region. A conspicuous solid transversal line of Burnt Umber (48) color in the fourth-sixth line of neck dorsal scales. Between occiput and rump, nine transversal series of four rounded, half-moon to roughly kidneyshaped Burth Umber (48) spots (approximately 7 scales x 3 scales). Each spots bordered posteriorly by a Cream White (52) irregular spot. Dorsal spots turn into an oval shape on rump and fuse on tail forming ¨V¨ shaped marks in the first third part of the tail, with lateral and posterior borders Cream White (52). On the tail, marks fade progressively becoming irregular rings to the tip of the tail. On the lateral sides of the head, clear areas are larger and Cream White (52), with some transversal Robin Rufous (30) marks. On the limbs, Robin Rufous (30) transversal bands intermix with slightly clear bands. Gular region variegated with Mikado Brown (42) marks, mandibular area without any mark. Upper belly and lateral midtrunk with some Dusky Brown (285) speckles. Gular region, lower belly and femoral-tibial hind limbs Cream Yellow (82). All bright coloration disappears after capture, and after several years in preservative, all coloration became faded.

Color in preservative. In preservative the general brightly color disappear or fade, but color pattern of whites, blacks and greys persist for several years.

Variation. Table 3 summarizes the variation of males and females of the type series for some scale and morphometric characters. Precloacal pores are present only in males. In some males, a narrow Robin Rufous (30) vertebral band became apparent in life. In few individuals dorsal transversal marks connected longitudinally, becoming lines of clear dots on the dorsum. Dorsal spots shape varies, from half-moon shaped, usually in the middle of the trunk to quadrangular, usually near rump. Half-moon spots usually fused in the first in the upper scapular and first half of the trunk. Females ( Fig. 8 ) have a Drab (19) dorsal background coloration, and dorsal spots are usually more half-moon shaped than in males, but usually the first two-three rows are wavy-fused marks bordered with White Cream (52) scales. Each spots have in front a Robin Rufous (30) area, and connecting laterally with dorsolateral spots, a Cream Color (12) zone. Ventral areas of females lack of any dark mark or speckle. A few females have a Robin Rufous (30) longitudinal dorsolateral band connecting spots between scapular region to rump. On regenerated tails, a conspicuous narrow dark dorsal line extends from the cut area to almost the tip of the tail.

Etymology. To honor Sergio Igor Tiranti, an enthusiastic biologist, colleague and friend, who first discovered and collected the species in early 1990 at the type locality, and to his father Ivan Nicolas Tiranti, professor of genetics at the National University of Río Cuarto, Argentina, who helped the first author on its doctoral studies and shared its experiences and friendship in the academic world.

Geographic distribution. Liolaemus tirantii sp. nov. is known from three localities in a small area in central Neuquén province ( Fig. 1 ): (1) the type locality ( Fig. 9 ), (2) outskirts of La Amarga village, and (3) flatlands near National Road 22 and Provincial Road 34 junction, all in the Zapala Department. Scrocchi et al. (2010) includes a photography epigraph of an adult male of Liolaemus tirantii sp nov. with the name Liolaemus cf. donosobarrosi from Cutral-Co, Neuquén, a locality 40 East of our type locality. In the text, they said that L. cf. donosobarrosi ¨Se distribuye en el centro este de Neuquén, hasta Zapala y en Rio Negro fue hallada en el Cerro Policía, meseta de Renteria¨. Scrocchi et al. (2010) did not mention any voucher specimens from these localities and we cannot confirm above statements with any other reference material.

Natural history. The holotype and the paratypes were found basking on small rocks or running between bushes. All localities are in an ecotonal area between Patagonian Steppe (Erial Patagónico) and Southern Patagonian Monte ( Roig 1998). This ecotone is an arid region in central Neuquén province, between 800 to 1000 m above sea level, characterized by small hills, folded sedimentary hills, small alluvial plains, basaltic plateaus, and sedimentary piedmonts plateau-shaped. Annual rain usually is lower than 200 mm and medium temperatures are between 10–12° C. Soils are poor, deep, with very few organic matter that limits the vegetation grow and have a very high natural erosion.

Vegetation is typical of ecotone of Patagonian Steppe (Erial Patagónico) and Southern Patagonian Monte ( Roig 1998); predominant vegetation formation is a low to medium shrubs steppe (bushes between 0.50 to 2.00 m). Soils are heavily eroded by rain and wind with intermixed gravel and sand areas or small stony outcrops without vegetation. Common plants are Grindelia chiloensis, Senecio filaginoides , Acantholippia seriphioides , Ephedra ochreata , Schinus roigii , Stillingia patagonica , Chuquiraga avellanedae , Larrea nitida , L. divaricata , L. cuneifolia , L. ameghinoi , Atriplex lampa , Prosopis denudans , Bougainvillea spinosa and endemic species such as Prosopis catellanosii . All sites where L. tirantii sp. nov. was collected are similar in physiography, normally small barren areas, with sandy soil mixed with gravel areas or more compacted soil. Clumps of shrub appear in areas with some degree of protection from the strong winds typical of Patagonia; these clumps are composed mainly of Prosopis denudans and smaller shrubs of Atriplex sp., Lycium chiliensis , and Chuquiraga avellanedae , all of which grow under the protection of this spiny mesquite. These clumps reach a maximum height of 1.8 m, and cover an area between two to four m2 and separated from each other by distances of two or three meters. This region is characterized by highly degraded soils (due to overgrazing or natural erosion), and shows a tendency towards desertification. Clumps seem to be the safest areas for lizards, as well as for small mammals and birds. The majority of the lizards were observed only inside or in the immediate vicinity of the spiny shrub clumps and in few cases basking on the top of stones or open areas. Usually lizards bask under sun in early morning (09:00 to 10:00) and in late afternoon (18:00 to 19:00) in the mid-summer, when they show a bimodal type of activity, but in early spring and late-summer they usually show continuous activity without a mid-day break. When days are cloudy or cold in summer they show an all-day round activity. In the type locality we collected other species of lizards living in syntopy with L. tirantii sp. nov., including L. darwinii , L. gracilis , L. gununakuna , and L. aff. mapuche , as well as a teiid lizard, Aurivela longicauda and a gecko, Homonota darwinii . In nearby localities we collected Bothrops ammodytoides , Philodryas patagoniensis and P. trilineata , all are known to eat lizards and are probable predators of L. tirantii . Liolaemus tirantii sp. nov. is oviparous as well others members of its clade, and feeds mainly in arthropods and some plant matter as a dissected stomach content show.

Figures

FIGURE 1. Map of southern Mendoza, eastern Neuquén, northwestern Río Negro and southwestern La Pampa provinces showing the complex landscape of this region with main orographic landmarks marked. White circles are main towns in the area, marked as reference, red lines are principal roads. Blue lines are main rivers, Colorado and Neuquén. Main mountain ranges, are marked.Inset:Regionin SouthAmérica.Line-dotted line show the political boundaries between four provinces cited above; other limits follow rivers and are not marked for simplicity. Localities sampled are shown with black (L. donosobarrosi, T.L.), red (L. donosobarrosi sp. 1), blue (L. donosobarrosi sp. 2) and green (L. donosobarrosi sp. 3) circles. Orange dotted line marks the approximate boundaries between the Patagonian Steppe and Southern Patagonian Monte.

FIGURE 6. Liolaemus tirantii sp. nov., holotype adult male from a small unnamed sedimentary hill 2 km S Cerro Bandera, 1 km S National Road 22, 18.5 km W Cutral-Có, Zapala Department, Neuquén Province, Argentina.

FIGURE 7. Liolaemus tirantii sp. nov., holotype adult male in dorsal and ventral view.

FIGURE 8. Liolaemus tirantii sp. nov., undetermined paratype adult female from a small unnamed sedimentary hill 2 km S Cerro Bandera, 1km S National Road 22, 18.5 km W Cutral-Có, Zapala Department, Neuquén Province, Argentina.

FIGURE9. Type locality of Liolaemustirantii sp. nov. Upper:generalview of the area.Below:close view of the microhabitat where lizards were collected.