Liolaemus avilae Breitman, Parra, Pérez & Sites, 2011, sp. nov.
Creators: name: Breitman, Maria Florencia
name: Parra, Micaela
name: Pérez, Cristian Hernán Fulvio
name: Sites, Jack Walter
Publisher: name: Zenodo
Year: 2011
Subjects: Biodiversity
Taxonomy
Animalia
Chordata
Reptilia
Squamata
Liolaemidae
Liolaemus
Liolaemus avilae

description: Liolaemus avilae sp. nov.
(Figures 6, 7)
1971, Liolaemus lineomaculatus, Donoso-Barros, R. and Cei, J.M., Journal of Herpetology, 5, 89–95. 1975, Liolaemus lineomaculatus, Cei, J.M., Herpetologica, 31, 109 –116.
1982, Liolaemus lineomaculatus, Cei, J.M. and Scolaro, J.A., Journal of Herpetology, 16, 354–363. 1992, Liolaemus lineomaculatus, Scolaro, J.A., Acta zoologica lilloana, 41, 287 –293.
Holotype. MLP.S 2627 (Figure 7), an adult male from Lago Buenos Aires plateau, 18.7 SW Puesto Lebrun, Lago Buenos Aires department, Santa Cruz province, Argentina (47º05’29.0” S, 71º01’12.9” W, 1154 m) (Figures 5, 8), L.J. Avila, C.H.F. Pérez, M.F. Breitman and N. Feltrin collectors, 9th January 2008.
Paratypes. LJAMM-CNP 9250, 9253, 9274, adult males, LJAMM-CNP 9276-9277, 9399, adult females and LJAMM-CNP 9251, 9252, juveniles; from same locality as holotype, L.J. Avila, C.H.F. Pérez, M.F. Breitman and N. Feltrin collectors, 9 January 2008. LJAMM-CNP 9243, an adult male from Puesto Lebrun, 27.3 km W Estancia La Vizcaina, Lago Buenos Aires Plateau, Lago Buenos Aires department, Santa Cruz province, Argentina (46º57’51.8” S, 71º06’27.2” W, 1353 m), L.J. Avila, C.H.F. Pérez, M.F. Breitman and N. Feltrin collectors, 8th January 2008.
Diagnosis. Liolaemus avilae sp. nov., a member of the L. lineomaculatus section and specifically the lineomaculatus group, has dorsal trifid scales but lacks of precloacal pores in either sex (Etheridge 1995); molecular evidence places this species in the lineomaculatus group, as the sister species to L. lineomaculatus (Breitman et al. 2011; see Tables 1 to 4 and Figure 6).
Relative to L. morandae sp. nov., L. avilae sp. nov. has more scales in contact with the interparietal scale (7– 10, X = 8,13 vs. 6–7, X = 6.33; p = 0.0027), fewer supralabial scales (5–6, X = 5.25 vs. 5–10, X = 7.17; p = 0.0008), fewer midbody scales (53–59, X = 55.5 vs. 55–63, X = 59.83; p <0.0001), more third finger lamellae (15– 18, X = 16.25 vs. 13–16, X = 15.33; p <0.0001), a longer elbow-wrist distance (4.89–6.81, X = 5.76 vs. 4.69–5.56, X = 5.32; p <0.0001), and a shorter rostral-nasal distance (1.69–2.25, X = 1.95 vs. 2.02–2.42, X = 2.15; p <0.0001). Liolaemus avilae sp. nov. has fewer dorsal blotches than L. morandae sp. nov., and the blotches in L. morandae sp. nov. have (50% of cases) brownish coloration, while L. avilae sp. nov. blotches have dark gray color. Ventral hand and foot surfaces of L. morandae sp. nov. present mucronated and “bristly” scales that are less obvious in L. avilae sp. nov.
Liolaemus avilae sp. nov. differs from L. lineomaculatus in the following traits: more scales in contact with the interparietal (7–10, X = 8,13 vs. 6–10, X = 7.18; p = 0.0027), fewer supralabial scales (5–6, X = 5.25 vs. 5–8, X = 6.71; p = 0.0008), fewer midbody scales (53–59, X = 55.5 vs. 54–65, X = 60; p <0.0001), fewer scales from occiput to rump (51–58, X = 53.75 vs. 52–68, X = 58.41; p <0.0001), fewer ventral scales (72–91, X = 83 vs. 80– 95, X = 85.29; p <0.0001), larger eyes (eye height: 2–2.62, X = 2.2 vs. 1.5–2.21, X = 1.84, eye length: 2.63–3.32, X = 3.07 vs. 2.35–3.3, X = 2.83; p <0.0001 in both cases), and longer auditory meatus lengths (1.11–2.1, X = 1.51 vs. 0.87–1.39, X = 1.15; p <0.0001). Liolaemus avilae sp. nov. has fewer dorsal blotches than L. lineomaculatus. The vertebral line is less well defined in L. avilae sp. nov. than in L. lineomaculatus. In general, dorsal scales of L. avilae sp. nov. are more mucronated than in L. lineomaculatus. Liolaemus avilae sp. nov. possesses scales in the dorsal surface of the forelimbs that are less carinated than those of L. lineomaculatus.
Compared with L. hatcheri, L. avilae sp. nov. has more scales in contact with the interparietal (7–10, X = 8,13 vs. 5–10, X = 6.85; p = 0.0027), fewer supralabial scales (5–6, X = 5.25 vs. 6–8, X = 6.85; p = 0.0008), more midbody scales (53–59, X = 55.5 vs. 43–57, X = 48.65; p <0.0001), more scales from occiput to rump (51–58, X = 53.75 vs. 43–55, X = 48.75; p <0.0001), more ventral scales (72–91, X = 83 vs. 62–74, X = 67.8; p <0.0001), more third finger lamellae (15–18, X = 16.25 vs. 13–18, X = 14.85; p <0.0001), more fourth toe lamellae (20–23, X = 21.29 vs. 18–22, X = 19.45; p <0.0001), smaller body size (snout-vent length: 48–59, X = 55.38 vs. 55–69, X = 61.89, axilla-groin distance: 19.4–29.5, X = 25.5 vs. 19.5–35.5, X = 28.95; p <0.0001 in both cases), shorter forelimbs (elbow-wrist length: 4.89–6.81, X = 5.76 vs. 5.67–7.33, X = 6.44, third finger length: 7.27–9.52, X = 8.02 vs. 8.5–10.57, X = 9.15; p <0.0001 in both cases), and smaller head width (9.02–10.19, X = 9.59 vs. 9.7– 13.05, X = 11.22; p <0.0001). Liolaemus avilae sp. nov. possesses fewer dorsal blotches and a more defined vertebral line relative to L. hatcheri. Ventral melanism is more pronounced in L. hatcheri than in L. avilae sp. nov. (fewer melanophores). Dorsal scales of L. avilae sp. nov. are less “bristly” than those of L. hatcheri. In the new species, limb surface scales (fore-hind and dorsal-ventral) are non-mucronate and carinated (in L. hatcheri the majority are mucronated and non-carinate). Ventral scales are round-shaped in L. avilae sp. nov. and rhomboidal in L. hatcheri.
Compared with L. kolengh, L. avilae sp. nov. has fewer scales in contact with the interparietal (7–10, X = 8,13 vs. 4–8, X = 6.26; p = 0.0027), fewer supralabial scales (5–6, X = 5.25 vs. 5–8, X = 6.65; p = 0.0008), more midbody scales (53–59, X = 55.5 vs. 44–55, X = 50.55; p <0.0001), more scales from occiput to rump (51–58, X = 53.75 vs. 44–54, X = 49.35; p <0.0001), more ventral scales (72–91, X = 83 vs. 61–76, X = 68.58; p <0.0001), more third finger lamellae (15–18, X = 16.25 vs. 13–16, X = 14.6; p <0.0001), more fourth toe lamellae (20–23, X = 21.29 vs. 16–22, X = 18.74; p <0.0001), longer tails (64–77, X = 68.88 vs. 47–70, X = 60.52, p = 0.0019), shorter forelimbs (elbow-wrist length: 4.89–6.81, X = 5.76 vs. 4.6–7.03, X = 6.16, third finger length: 7.27–9.52, X = 8.02 vs. 8.31–11.12, X = 9.65; p <0.0001 in both cases), smaller head width (9.02–10.19, X = 9.59 vs. 8.95– 11.71, X = 10.73; p <0.0001), and smaller rostral scale (rostral height: 0.55–1.09, X = 0.85 vs. 0.87–1.2, X = 1.03, rostral length 2.31–2.69, X = 2.51 vs. 2.46–3.12, X = 2.77; p <0.0001 in both cases). Liolaemus avilae sp. nov. exhibits a light background coloration, while the general background coloration of L. kolengh is dark gray or black; however, the dorsal pattern could not be recognized in around 50% of our samples. Liolaemus kolengh exhibits more dorsal blotches than L. avilae sp. nov. Ventral melanism is stronger in L. kolengh than in L. avilae sp. nov. In general, the dorsal scales of L. avilae sp. nov. are less “bristly” than scales of L. kolengh. Limb scales (fore-hind and dorsal-ventral) are non-mucronate and non-“bristly” in L. avilae sp. nov. (mucronated and “bristly” in L. kolengh).
Liolaemus avilae sp. nov. differs from L. silvanae in several characters, including: a smaller number of scales in contact with the interparietal (7–10, X = 8,13 vs. 5–8, X = 6.94; p = 0.0027), fewer supralabial scales (5–6, X = 5.25 vs. 5–7, X = 6.18; p = 0.0008), fewer midbody scales (53–59, X = 55.5 vs. 55–65, X = 59.65; p <0.0001), fewer scales from occiput to rump (51–58, X = 53.75 vs. 52–64, X = 56; p <0.0001), more ventral scales (72–91, X = 83 vs. 70–80, X = 74.59; p <0.0001), smaller body size (snout-vent length: 48–59, X = 55.38 vs. 65–78, X = 73, axilla-groin distance: 19.4–29.5, X = 25.3 vs. 25.1–39.7, X = 32.38; p <0.0001 in both cases), shorter fourth toe length (13.82–15.98, X = 14.91 vs. 16.5–20.1, X = 18.46), shorter forelimbs (elbow-wrist length: 4.89–6.81, X = 5.76 vs. 6.83–8.82, X = 7.71, third finger length: 7.27–9.52, X = 8.02 vs. 10.65–13.76, X = 12.23; p <0.0001 in both cases), smaller head (head height: 6.43–7.67, X = 7.07 vs. 6.84–9.06, X = 8.18, head width: 9.02–10.19, X = 9.59 vs. 11.31–14.71, X = 13.14, head length: 10.94–12.9, X = 11.78 vs. 12.47–15.9, X = 14.42; p <0.0001 in all cases), smaller eyes (eye height: 2–2.62, X = 2.2 vs. 2.09–2.9, X = 2.46, eye length: 2.63–3.32, X = 3.07 vs. 3.59– 4.18, X = 3.86; p <0.0001 in both cases), smaller rostral scale (rostral height: 0.55–1.09, X = 0.85 vs. 1.01–1.35, X = 1.19, rostral length: 2.31–2.69, X = 2.51 vs. 2.61–3.47, X = 3.13; p <0.0001 in both cases), and smaller auditory meatus (auditory meatus height: 1.95–2.56, X = 1.95 vs. 1.95–2.95, X = 2.54, auditory meatus length: 1.11–2.01, X = 1.51 vs. 1.49–2.57, X = 2.04; p <0.0001 in both cases). Background coloration in L. silvanae is black, there is a no recognizable dorsal pattern, and ventral area is strongly melanistic, whereas the ventral area is weakly melanistic in L. avilae sp. nov. Liolaemus avilae sp. nov. has a defined vertebral and paravertebral lines that are not present in L. silvanae. In general, L. silvanae possesses a “bristly” appearance that is absent in L. avilae sp. nov. Scales of limbs (fore-hind and dorsal-ventral) are less mucronated in L. avilae sp. nov. than in L. silvanae.
Description of holotype. Adult male. Snout-vent length 59.0 mm. Tail length (complete, not regenerated) 77.0 mm. Axilla-groin distance 26.9 mm. Head length 12.3 mm (from anterior border of tympanum to tip of snout), 10.1 mm wide (at anterior border of tympanum), 7.6 mm high (at anterior border of tympanum). Snout length 4.2 mm (orbit-tip of snout distance). Interorbital distance 4.5 mm. Eye-nostril distance 3.5 mm. Auditory meatus-eye distance 4.9 mm. Forelimb length 16.3 mm. Tibial length 11.0 mm. Foot length 15.9 mm (ankle to tip of claw on fourth toe).
Dorsal head scales bulged, smooth, 15 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 absence at the posterior half of the head. Rostral scale wider (2.8 mm) than high (0.8 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, in a posterior position. Scales surrounding nasals 7 on the left side and 6 on the right side. Four internasals. Frontonasals four, irregular in size and position. Prefrontals 5, a small quadrangular scale in the center (1.1 mm), two dorso-lateral larger scales roughly hexagonal (1.7 mm), and a pair of lateral medium-sized scales (1.2 mm), approximately triangular. Three frontal scales. Frontoparietals in two rows, one anterior and two posterior scales. Interparietal pentagonal (1.3 mm), surrounded by seven scales; five in front and sides, irregular in shape and size, and two larger posterior scales. Parietal eye evident. Parietals slightly bulged, irregularly shaped, similar in size to interparietal. Circumorbitals: 12–10. Transversally expanded supraoculars 5– 4. Smaller lateral supraoculars: 16–19. One canthal higher than wide, separate from nasal by one postnasals. Loreal scales bulged, four on each side. Lorilabials longer than wide (7–6), approximately equal to labials. Superciliaries 7–8, flattened and elongated, anterior four broadly overlapping dorsally. Orbit with 12–15 upper and 13–11 lower ciliaries on each side. Orbit diameter 4.3 x 2.0 mm. Preocular small, unfragmented, square. Subocular scale elongated, approximately nine times longer than wide (3.8 x 0.6 mm). A well marked longitudinal ridge along upper margin of preocular and subocular scales. Postocular small, slightly bulged, ~ 25% superimposed onto subocular, with a marked longitudinal ridge. Palpebral scales small granular and bulged. Supralabials 6–5, convex. Temporals smooth, convex, juxtaposed with one scale organ in the tip. Anterior auriculars smaller than adjacent posterior temporals, projecting slightly outward (3–2). Posterior auricular scales small and granular. External auditory meatus conspicuous, higher (1.7 mm) than wide (1.2 mm). Lateral scales of neck granular with inflated skin. Mental scale wider (3.0 mm) than high (1.0 mm), in contact with four scales. Mental followed posteriorly by two postmentals, and two rows of three chinshields on each side. Infralabials 5-4 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-seven gulars between tympanum openings. Infralabials separated from chinshields by one to two rows of scales. Antehumeral and longitudinal neck folds well developed; postauricular, rictal, dorsolateral, oblique and gular not conspicuous.
Scales of dorsal neck region rhomboidal, imbricate, strongly keeled. Very few trifids scales. Fifty-five dorsal scales between occiput and anterior surface of thighs. Dorsal body scales rhomboidal, imbricate, strongly keeled. Dorsal scales grade laterally into slightly smaller and 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 rhomboidals, smooth, flat, imbricate, larger than dorsal scales. Fifty-eight midbody scales; scales between mental and precloacal pores 90. Scales of cloacal region similar 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 rhomboidal, imbricate, smooth. Supracarpals imbricated, rhomboidal, smooth. Infracarpals imbricate, rhomboidal, very slightly keeled. Subdigital lamellae with 2–3 keels, each terminating in a short mucron, 2–3 mucronate, numbering: I: 8, II: 14, III: 17, IV: 19, V: 12. Claws robust, curved and sharp, opaque brown. Suprafemorals larger as dorsal body scales, rhomboidal, imbricated, smooth upper thigh, toward to the lateral thigh keeled. Postfemorals small, granular shape. Supratibials rhomboidal, imbricated, moderately keeled, smaller and smooth toward to the foot. Infrafemoral scale small, granular and smooth. Supratarsals rhomboidal, imbricated and smooth. Infratarsals small, rhomboidal, imbricate, smooth, some keeled and mucronate near the digit. Subdigital scales have 3 keels, 3–4 mucronate, numbering: I: 9, II: 14, III: 19, IV: 22, V: 16. Claws robust, curved and sharp, opaque brown. Tail complete, non-regenerated. Dorsal and lateral caudal scales, rhombic in the first half of the tail, quadrangulars toward the tip, strongly keeled. Ventral subtriangular and smooth, toward posterior half moderatly keeled.
Color of holotype in life. Dark gray dorsal background (Figure 7) that becomes lighter on the lateral region between axilla and groin. Dorsal pattern presents nine paravertebral, quadrangulars, dark brown blotches, extending from the nuchal to the pelvic region, these series or blotches are combined at the beginning of the pelvic region into a pattern of transversal bands, which is present to the tip of the tail. Dorsolateral region between the occipital and the pelvic region is characterized by eight irregular, black and white-bordered blotches that are combined with the transversal bands of the tail. Lateral region, between axilla and groin, is characterized by a black and white reticulate pattern. In between the series of blotches a white vertebral and two yellowish paravertebral lines, onescale wide, are present. Vertebral line extends from the nuchal region, while the paravertebral lines extend from the temporal region; all are present to the pelvic region.
Dark gray head background. Two lateral white lines along the longitudinal fold are present from the top of the auditory meatus to the antehumeral fold. White subocular scale with a black longitudinal ridge. One longitudinal band is present in the anterior part of the head, crossing though internasals, postnasals, prefrontals, frontals, postfrontals and interparietal scales, ending in the interparietal scale that is surrounded by small brown spots; however, a lighter central region is observable in the frontal, postfrontal and interparietal scales. Anterior circumorbitals present a transverse dark brown band. Nasals scales dark brown. Small dark brown blotches on postocular, temporal and occipital region. Dorsal region of the limbs are dark gray with a dark brown reticulation.
Gray ventral color with a dark reticulate pattern over the entire body, it is strongly marked in throat, chest, tail and body laterals, and extend to the adjacent malar and maxilla region. Ventral scales with melanophores. Ventral area of chest, belly, cloacal region and limbs tinged with a light orange color. Gray tail.
Color of holotype in preservative. After three years in preservative, the dorsal coloration of the head, dorsum, body flanks and tail become 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 orange color of the chest, belly, cloacal region and limbs turns gray (Figure 6).
Variation. Morphological and meristic character variation between male and female Liolaemus avilae sp. nov., is shown in Table 6. Females in life present basically the same dorsal and lateral patterns as present in males, but females generally have a lighter brown background coloration; vertebral and paravertebral lines are variable, in some individuals they are strongly defined while in others they are less conspicuous; lines vary from white to yellow in females and from light orange to pink in males. Ventrally dark reticulate pattern in the entire body is present and is strongly marked in the throat, tail and lateral body scales. Ventral, ventro-lateral, limb, cloacal and postcloacal regions vary from males to females, in the former a ventral light orange coloration is present, while in one female (out of three) a light pink ventral coloration is present. All distinctive ventral and dorsal coloration changed from yellow, orange or pink to gray or white in preservative.
Etymology. The specific epithet of this species “ avilae ” refers to our colleague and the first author’s PhD advisor Dr. Luciano Javier Avila, to honor him after more than fifteen years of research and teaching on the Patagonian herpetofauna.
Distribution. Liolaemus avilae sp. nov. is known only from the type locality and surroundings, from Lago Buenos Aires Plateau in Puesto Lebrun and 18.7 SW from there, Santa Cruz province, Lago Buenos Aires department, Argentina (Figures 5, 8).
Natural history. Out of three females, only one presents a ventral pinkish coloration and a different one was pregnant; thus we infer that the pinkish coloration in females is not related to pregnancy. The species occurs in the Altoandina Phytogeographic province, Altoandino Austral district, in environments characterized as sparse grasslands and chamaephytes such as Empetrum rubrum, Nassauvia pygmaea and Azorrella ameghinoi, among others (Roig 1998); some Festuca spp. and Senecio spp. were also observed. This new species was found in open volcanic-sand substrates between patches of basaltic rocks, sharing the habitat with Liolaemus archeforus and Liolaemus silvanae (Figure 8). Based on the natural history of its sister species L. lineomaculatus, we hypothesize that L. avilae sp. nov. is herbivorous and likely to also be viviparous, perhaps giving birth to between three and six individuals per clutch (Cei 1986; Espinoza et al. 2004).
description: Published as part of 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, pp. 1-28 in Zootaxa 3120 on pages 16-21, DOI: 10.5281/zenodo.208130