Liolaemus morandae, Breitman, Maria Florencia, Parra, Micaela, Pérez, Cristian Hernán Fulvio & Sites, Jack Walter, 2011

Breitman, Maria Florencia, Parra, Micaela, Pérez, Cristian Hernán Fulvio & Sites, Jack Walter, 2011, Two new species of lizards from the Liolaemus lineomaculatus section (Squamata: Iguania: Liolaemidae) from southern Patagonia, Zootaxa 3120, pp. 1-28 : 7-13

Liolaemus morandae sp. nov.

( Figure 3–6 )

2001, Liolaemus lineomaculatus, Ibargüengoytía, N., Casalins, L., Schulte II, J.A., Amico, G.A. and Sympson, L., Herpetological Review, 32, 120.

Holotype. MLP.S 2626 ( Figure 3 ), an adult male from Provincial Road 37, 22.8 km SW junction National Road 3, Escalante department, Chubut province, Argentina (45°41’10,6” S; 67°53’49,9” W, 693 m) ( Figures 4 , 5 ); L.J. Avila, M. Kozykariski, M.F. Breitman and R. Martinez collectors, 12th March 2010.

Paratypes. LJAMM-CNP 13020, adult female and LJAMM-CNP 13021, juvenile; same locality as holotype. LJAM-CNP 9677-9679, adult females and LJAMM-CNP 9680, juvenile; from Holdich station, Escalante department, Chubut province, Argentina (45º58’00.1” S; 68º11’58.8” W, 761 m); L.J. Avila, C.H.F. Pérez, M.F. Breitman and N. Feltrin collectors, 2nd February 2008. LJAMM-CNP 10201, adult male and LJAM-CNP 10202, juvenile; from Provincial Road 37, 2.5 km W junction National Road 3, Escalante department, Chubut province, Argentina (45º37’43.4” S; 67º41’03.6” W, 637 m); L.J. Avila, C.H.F. Pérez, M.F. Breitman and N. Feltrin collectors, 2 nd February 2008.

Diagnosis. Liolaemus morandae sp. nov. is a member of the L. lineomaculatus section, included in the lineomaculatus group, and is characterized by the absence of precloacal pores in both sexes, and presence of dorsal trifid scales ( Etheridge 1995). Molecular evidence includes this species in the lineomaculatus group, closely related to the clade (L. sp. 2 + L. lineomaculatus ) Figure 1 . All the following differences are summarized in Tables 1 to 4 and in Figure 6 .

Relative to L. lineomaculatus , L. morandae sp. nov. has fewer dorsal scales (47–57, X = 51.67 vs. 52–68, X = 58.41; p <0.0001), fewer third finger lamellae (13–16, X = 15.33 vs. 14–19, X = 16,18; p <0.0001), shorter fourth toe length (13.4–16.8, X = 14.9 vs. 12.6–16.1, X = 14.09; p <0.0001) and a larger eye (eye height: 1.7–2.27, X = 1.99 vs. 1.5–2.21, X = 1.84, eye length: 2.91–3.42, X = 3.15 vs. 2.35–3.3, X = 2.83; p <0.0001 in both cases). Liolaemus morandae sp. nov. has more dorsal blotches than L. lineomaculatus , and the blotches in L. morandae sp. nov. have (in 50% of the cases) a yellow-orange coloration while L. lineomaculatus blotches have dark gray color.

In general, dorsal scales of L. morandae sp. nov. are more mucronated than those of L. lineomaculatus , and dorsal leg scales of L. morandae sp. nov. are less carinated with respect to L. lineomaculatus .

Compared to L. hatcheri , L. morandae sp. nov. has higher number of midbody scales (55–63, X = 59.83 vs. 43–57, X = 48.65 p <0.0001), larger number of scales from occiput to rump (47–57, X = 51.67 vs. 43–55, X = 48.75; p <0.0001), more ventral scales (79–85, X = 81.33 vs. 62–74, X = 67.8; p <0.0001), more fourth toe lamellae (18–23, X = 21.17 vs. 18–22, X = 19.45; p <0.0001), smaller body size (snout-vent length: 50–61, X = 55.67 vs. 55–69, X = 61.89, axilla-groin distance: 19.7–30.8, X = 25.9 vs. 19.5–35.5, X = 28.95; p <0.0001 in both cases), shorter forelimbs (elbow-wrist length: 4.69–5.56, X = 5.32 vs. 5.67–7.33, X = 6.44, third finger length: 6.69–9.14, X = 7.83 vs. 8.5–10.57, X = 9.15; p <0.0001 in both cases), smaller head (head width: 9.08–10.58, X = 9.84 vs. 9.7–13.05, X = 11.22, head length: 10.96–13.06, X = 11.96 vs. 11.43–14.65, X = 12.97; p <0.0001 in both cases), and smaller auditory meatus (auditory meatus height: 1.6–2.44, X = 1.95 vs. 1.85–2.61, X = 2.15; auditory meatus length: 0.86–1.81, X = 1.27 vs. 1.38–2.16, X = 1.77; p <0.0001 in both cases). Liolaemus morandae sp. nov. has more dorsal blotches and better defined vertebral and paravertebral lines than L. hatcheri ; ventral melanism is more pronounced in L. hatcheri , while L. morandae sp. nov. has fewer melanophores. Dorsal scales of L. morandae sp. nov. are less “bristly” than those of L. hatcheri . Dorsal and ventral limb scales are non-mucronate and carinate in L. morandae sp. nov. while in L. hatcheri they are mucronate and non-carinate. Belly scales are round-shaped in L. morandae sp. nov. and rhomboidal in L. hatcheri .

Liolaemus morandae sp. nov. differs from L. kolengh in the following traits: larger number of midbody scales (55–63, X = 59.83 vs. 44–55, X = 50.55; p <0.0001), larger number of scales from occiput to rump (47–57, X = 51.67 vs. 44–54, X = 49.35; p <0.0001), more ventral scales (79–85, X = 81.33 vs. 61–76, X = 68.58; p <0.0001), larger number of fourth toe lamellae (18–23, X = 21.17 vs. 16–22, X = 18.74; p <0.0001), shorter forelimbs (elbow-wrist length: 4.69–5.56, X = 5.32 vs. 4.6–7.03, X = 6.16, third finger length: 6.69–9.14, X = 7.83 vs. 8.31– 11.12, X = 9.65; p <0.0001 in both cases), smaller auditory meatus (auditory meatus height: 1.6–2.44, X = 1.95 vs. 1.63–2.95, X = 2.22, auditory meatus length: 0.86–1.81, X = 1.27 vs. 1.22–2.24, X = 1.67; p <0.0001 in both cases) and longer tail (61–79, X = 70.33 vs. 47–70, X = 60.52; p = 0.0019). Liolaemus morandae sp. nov. has more dorsal blotches and better defined vertebral and paravertebral lines than L. kolengh ; ventral melanism is stronger in L. kolengh than in L. morandae sp. nov.; the general background coloration of L. kolengh is dark gray or black, though in almost 50% of our samples the dorsal pattern cannot be recognized. In general, the dorsal scales of L. morandae sp. nov. are less “bristly” than those of L. kolengh . Dorsal and ventral limbs scales are non-mucronate and non-bristly in L. morandae sp. nov., but mucronate and “bristly” in L. kolengh .

Liolaemus morandae sp. nov. differs from L. silvanae in several characters, including: a smaller number of scales from occiput to rump (47–57, X = 51.67 vs. 52–64, X = 56; p <0.0001), higher number of ventral scales (79–85, X = 81.33 vs. 70–85, X = 74,59; p <0.0001), fewer third finger lamellae (13–16, X = 15.33 vs. 14–18, X = 16.24; p <0.0001), smaller body size (snout-vent length: 50–61, X = 55.67 vs. 65–78, X = 73, axilla-groin distance: 19.7–30.8, X = 25.9 vs. 25.1–39.7, X = 32.38; p<0.0001 in both cases), shorter forelimbs (elbow-wrist length: 4.69–5.56, X = 5.32 vs. 6.83–8.82, X = 7.71, third finger length: 6.69–9.14, X = 7.83 vs. 10.65–13.76 X = 12.23; p <0.0001 in both cases), smaller head (head height: 6.52–8.21, X = 7.23 vs. 6.84–9.06, X = 8.18, head width: 9.08–10.58, X = 9.84 vs. 11.31–14.71, X = 13.14, head length: 10.96–13.06, X = 11.96 vs. 12.47–15.9, X = 14.42; p <0.0001 in all cases), smaller eye (eye height: 1.7–2.27, X = 1.99 vs. 2.09–2.9, X = 2.46, eye length: 2.91–3.42, X = 3.15 vs. 3.59–4.18, X = 3.86; p <0.0001 in both cases), and smaller auditory meatus (auditory meatus height: 1.6–2.44, X = 1.95 vs. 1.95–2.95, X = 2.54; auditory meatus length: 0.86–1.81, X = 1.27 vs. 1.49– 2.57, X = 2.04; p <0.0001 in both cases). The general background coloration of L. silvanae is black; there is no recognizable dorsal pattern, and the venter is also strongly melanistic, whereas the venter is weakly melanistic in L. morandae sp. nov. Liolaemus morandae sp. nov. possesses well-defined vertebral and paravertebral lines that are not present in L. silvanae . In general, L. silvanae possesses a “bristly” appearance that is not present in L. morandae sp. nov. Limb scales are less mucronated in L. morandae sp. nov. than in L. silvanae .

Description of holotype. Adult male. Snout-vent length 56.0 mm. Tail length (complete, not regenerated) 79.0 mm. Axilla-groin distance 22.8 mm. Auditory meatus-eye distance 4.8 mm. External auditory meatus conspicuous, higher (2.1 mm) than wide (1.2 mm). Head length 13.0 mm (from anterior border of tympanum to tip of snout), 10.5 mm wide (at anterior border of tympanum), 7.8 mm high (at anterior border of tympanum). Snout length 4.0 mm (orbit-tip of snout distance). Interorbital distance 4.2 mm. Eye-nostril distance 3.1 mm. Forelimb length 16.7 mm. Tibial length 10.3 mm. Foot length 16.8 mm (ankle to tip of claw on fourth toe).

Dorsal head scales bulged, smooth, 15 between occiput, at the 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 (2.4 mm) than high (0.9 mm). Two postrostrals, together with anterior lorilabial, separate nasal scales from rostral, surrounded by six scales. Nasal scales longer than wide, irregularly hexagonal; nostril one-half length of nasal, posterior in position. Scales surrounding nasals 7 on the left side and 8 on the right side. Four internasals. Frontonasals six, irregular in size and position. Prefrontals 6, a small rhomboidal scale in the center (0.9 mm). Three dorso-lateral larger scales, one for the fragmentation of the right scale, and a pair lateral medium-sized scales (1.3 mm), approximately quadrangular. Two frontal scales. Frontoparietals in two rows, one anterior and one posterior scale, unfragmented. Interparietal pentagonal, surrounded by six scales; four smaller and irregular in front and sides, two larger in back. Parietal eye evident. Parietals slightly bulged, irregularly shaped, equal in size to interparietal (1.5 vs. 1.5 mm). Circumorbitals:13–10. Transversally expanded supraoculars 4–3. Smaller lateral supraoculars: 16–17. One canthal higher than wide, separated from nasal by one postnasal. Loreal scales bulged, two on the left side (by fusion of the posterior loreal scale with preocular scale) and three on the rigth side. Lorilabials longer than wide (8–6), approximately equal to labials. Superciliaries 7–7, flattened and elongated, anterior five broadly overlapping dorsally. Orbit with 15–17 upper and 12–13 lower ciliaries on each side. Orbit diameter 4.5 x 1.9 mm. Preocular small, unfragmented, longer than wide. Subocular scale elongated, approximately nine times longer than wide (3.7 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 9–10, convex. Temporals smooth, convex, juxtaposed with one scale organ in the tip. Anterior auriculars smaller than adjacent posterior temporals, slightly projecting outward (3–2). Posterior auriculars small and granular. External auditory meatus conspicuous, higher (2.1 mm) than wide (1.2 mm). Lateral scales of neck granular with inflated skin. Mental scale wider (2.9 mm) than high (1.4 mm), in contact with four scales. Mental posterior followed by two postmentals, and two rows of three chinshields on each side. Six infralabials on each side, first on each side quadrangular two times wider than supralabials, all others elongated, 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 gulars between tympanum openings. Infralabials separated from chinshields by one to two rows of scales.

Antehumeral, longitudinal and postauricular neck folds well developed; gular, rictal, dorsolateral and oblique not conspicuous.

Scales of dorsal neck region rhomboidal, imbricate and strongly keeled. Fifty dorsal scales between occiput and anterior surface of thighs. Dorsal body scales rhomboidal, imbricate, strongly keeled, mucronate, very few trifid scales. Dorsal scales grade laterally into slightly smaller, slightly keeled scales at midbody. Scales immediately anterior and posterior to forelimb and hindlimb insertion small, smooth, granular, and non-overlapping. Body lateral scales grading smaller to larger at midbody. Ventral body scales rhomboidal, smooth, flat, imbricate, larger than dorsal scales. Sixty-one midbody scales; scales between mental and precloacal pores 85. Scales of cloacal region about equal in size to ventral body scales; without precloacal pores.

Anterior suprabrachials rhomboidal, imbricate, smooth, slightly larger in size to dorsal body scales. Postabrachials smaller, smooth, becoming granular near axilla. Supra-antebrachials similar to suprabrachial. Infra-antebrachials rhomboidals, imbricate, smooth, toward the hand slightly mucronate. Supracarpals imbricated, rhomboidal, smooth. Infracarpals strongly imbricate, rhomboidal, slightly keeled, 3-mucronate. Subdigital lamellae with 2–5 keels, each terminating in a short mucron, 2–5 mucronate, numbering: I: 8, II: 12, III: 16, IV: 17, V: 11. Claws robust, curved and sharp, opaque brown.

Suprafemorals as large as dorsal body scales, rhomboidal, imbricated, smooth near the body, toward to the knee slightly keeled. Postfemorals small, granular shape. Supratibials rhomboidal, imbricated, keeled, some mucronate, smaller and smooth toward to the foot. Infrafemoral scales small, granular and smooth. Supratarsals rhomboidal, imbricated and smooth. Infratarsals small, rhomboidal, imbricate, smooth, mucronate, slightly keeled near the digit. Subdigital scales 1–3 keeled, 1–5 mucronate, numbering: I: 9, II: 14, III: 20, IV: 21, V: 14. Claws robust, curved and sharp, opaque brown. Tail complete, non-regenerated. Dorsal and lateral caudal scales, rhomboidal in the first half of the tail, becoming quadrangular toward the tip, strongly keeled. Ventrals subtriangular and smooth, toward posterior half moderate keeled.

Color of holotype in life. Dark grey dorsal background ( Figure 3 ) that becomes lighter on the lateral region, between axilla and groin. Dorsal pattern presents twelve paravertebral, quadrangular, black blotches, extending from the nuchal to the postcloacal region, those series fuse into a dark line that is present to the tip of the tail. Dorsolateral region, in between the occipital and the tip of the tail, formed by black and white-bordered blotches. Lateral region, between axilla and groin, is characterized by a black and white reticulate pattern. In between the series of blotches, one white vertebral line and two light orange paravertebral longitudinal lines appear, one and one and a half scale wide, respectively. Vertebral line extends from the nuchal region, while the paravertebral lines extend from the temporal region; all of them continue to the tip of the tail. Dorsal region of the limbs are dark gray with a black reticulation.

Dark gray head background. Lateral white line along the longitudinal folds is present from the top of the auditory meatus to the antehumeral fold. White subocular scale with black longitudinal ridge. Dorsal surface with black blotches, one longitudinal blotch is present in the anterior part of the head, crossing through internasals, postnasals and prefrontals; two transverse blotches are crossing through postfrontals and first circumorbitals, and extend back to the last circumorbitals, forming a cross with the longitudinal blotch. The interparietal scale is surrounded by three small dark-brown blotches and the occipitals show a white longitudinal band of the same color.

White ventral color with a black reticulate pattern over the entire body, most strongly marked in the pectoral region and throat, and extending to the adjacent malar and maxilla region. Ventral scales with melanophores. Ventral area of chest, belly, cloacal region and limbs tinged with a light yellowish color. Cloacal and postcloacal region with few black blotches. Gray tail.

Color of holotype in preservative. After one year in preservative, the dorsal coloration of the head, dorsum, body flanks and tail becomes darker while maintaining the contrast, but the two yellowish paravertebral lines turned gray. Ventral scales of throat, neck, chest, belly and forelimbs retain the same coloration as in life, and the distinctive light ventral yellowish tinge of chest, belly, cloacal region and limbs turns gray ( Figure 6 ).

Variation. Morphological and meristic character variation between males and females of Liolaemus morandae sp. nov., are shown in Table 6 . Females in life present basically the same dorsal and lateral pattern as present in males, but females generally have a lighter brown background coloration; vertebral and paravertebral lines are strongly defined, and in both sexes paravertebral lines vary from white to yellow. Ventrally, one male shows a light orange color on the belly, while in other males and all females the ventral region ranged from white to gray due to different degrees of melanism. All distinctive ventral light orange coloration changed to gray in preservative.

Etymology. The specific epithet of this species “ morandae ” refers to our colleague and the first author’s PhD advisor Dr. Mariana Morando, to honor her after more than ten years of research and teaching on the Patagonian herpetofauna.

Distribution. Liolaemus morandae sp. nov. is known only from the type locality and a few localities on the Pampa del Castillo Plateau; and from three localities from southern Chubut and northern Santa Cruz provinces. The holotype and some paratypes are from Provincial Road 37, 22.8 km SW junction National Road 3, Escalante department, Chubut province, Argentina ( Figures 4 , 5 ). The other paratypes are from Provincial Road 37, 2.5 km W junction National Road 3 in Holdich Station, Escalante department, Chubut province, Argentina ( Figures 4 , 5 ). The distance from the type locality to the first locality is more than 17.8 km airline, and more than 38.8 km airline to the second locality.

Natural history. Liolaemus morandae sp. nov. occurs in Patagónica province, Golfo de San Jorge District, in environments characterized as grasslands and steppe highlands ( Roig 1998), with vegetation dominated by the shrubs Stipa spp ., Senecio filaginoides , Mulinum spinosum, Nassauvia spp ., among others. This species was found in open substrates, sharing the habitat with Liolaemus kingii , L. bibronii, Diplolaemus bibronii and D. darwinii ( Figure 4 ). Based on the natural history of its close relative L. lineomaculatus , we hypothesize that L. morandae sp. nov. is herbivorous and likely to also be viviparous, perhaps giving birth to between three and six neonates per clutch ( Cei 1986; Espinoza et al. 2004).

Figures

FIGURE 1. Relationships between Liolaemus morandae sp. nov. and L. avilae sp. nov. within the L. lineomaculatus section, and selected species of the subgenus Eulaemus and Liolaemus sensu stricto. On the left, phylogenetic relationships recovered with Bayesian inference using seven nuclear genes (Cmos, ACM 4 tg, PRLR, LDA 8 F, LDA 1 D, LDA 9 C and LDA 9 E, 4180 bp total alignment length), bold branches represent well supported nodes (Pp> 0.95). On the right, the Bayesian tree (modified from Breitman et al. 2011) represents a concatenated analyses including the seven nuclear genes plus two mitochondrial genes (12 S and cyt- b; 5865 bp) and summarizes information from MP and ML methods. Nodes with high support from three methods (MP, jackknife and ML bootstrap> 0.70; Bayesian Pp> 0.95) are identified by bold branches; solid circles show nodes with weak MP support and open circles nodes with weak MP and ML support.

FIGURE 3. Upper, dorsal and ventral views of male holotype in life of Liolaemus morandae sp. nov. MLP. S 2626; lower, dorsal and ventral views of female (LJAMM-CNP 13020).

FIGURE 4. Type locality of Liolaemus morandae sp. nov., Patagonic phytogeographic province, del Golfo de San Jorge District (45 ° 41 ’ 10,6 ” S; 67 ° 53 ’ 49,9 ” W, 693 m).

FIGURE 5. Distribution map for species of the lineomaculatus group, with geographical localities sampled in this study superimposed on regional elevation (shading). Black circles: L. morandae sp. nov.; white circle: L. avilae sp. nov.; black squares: L. lineomaculatus; white square: L. kolengh; white triangles: L. hatcheri; black triangle: L. silvanae. Arrows indicate type localities. CH: Chubut province; SC: Santa Cruz province.

FIGURE 6. Morphological comparison between Liolaemus morandae sp. nov. (A), L. lineomaculatus (B) and L. avilae sp. nov. (C). Notice differences in color pattern on the tails, size of blotches on the limbs and differences in vertebral and paravertebral lines. Dorsal scales (shown on the right) are less mucronated in L. lineomaculatus. Dorsal scales of hindlimbs are less carinated in L. morandae sp. nov. than in the other species, while they are more mucronated in L. avilae sp. nov.