Liolaemus antumalguen, Avila, Luciano Javier, Morando, Mariana, Perez, Daniel Roberto & Sites, Jack W., 2010
Avila, Luciano Javier, Morando, Mariana, Perez, Daniel Roberto & Sites, Jack W., 2010, A new species of the Liolaemus elongatus clade (Reptilia: Iguania: Liolaemini) from Cordillera del Viento, northwestern Patagonia, Neuquén, Argentina, Zootaxa 2667, pp. 28-42: 30-37
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Liolaemus antumalguen sp. nov.
Type material. Holotype. MACN 38985, an adult male from rocky boulders from the eastern piedmont of Domuyo volcano, around Chadileu Creek (36 º 39 ’ S, 70 º 20 ’’ W), Chos Malal Department, Neuquén Province, Argentina; collected by D. R. Perez; 23 Enero 2005.
Paratypes. MACN 38987, MLP.S 2592 – 3, (males); BYU 12592View Materials, MACN 38986, MLP.S 2594 – 5, LJAMM 6172 – 3 (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 unpublished).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 unpublished data).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 107.8 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 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, 96–110, 98– 115 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 102.6 vs 107.8 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 107.8 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 107.8 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, 60. 2 and 61.1 mm vs 102.7 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. 1View FIGURE 1, 2View FIGURE 2) 107.8 mm snout-vent length (SVL); tail length 87.3 .0 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: 14 – 10. Transversally expanded supraoculars 7 – 7. Smaller lateral supraoculars: 24 – 19. 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 15–16 upper and 12–13 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.
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.
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. 1View FIGURE 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.
Variation in paratypes. Based on the paratypes ( Table 2, Fig. 3View FIGURE 3): In three males: SVL (mean ± SD (range)): 95.5 ± 10.9 (86.6–107.8). Axilla –groin distance: 40.6 ± 4.3 (36.7–47.6). Head length: 20.2 ± 2.04 (18.7–22.5). Head width: 17.2 ± 1.95 (16.0– 19.5). Foot length: 30.0 ± 0.8 (29.1–30.8). Tibial length: 19.9 ± 1.0 (19.0–21.0). Hand length: 28.8 ± 0.7 (28.3–29.6). Midbody scales: 80.3 ± 2.0 (78–82). Dorsal scales: 75.0 ± 3.0 (72–78). Ventral scales: 111 ± 6.0 (107–118). Precloacal pores: 3–4. Scales around interparietal: 5–6 – 7. Supraoculars scales: 6 – 5. Scales around nasal 5–7. Supralabial scales: 7–8. Infralabial scales: 6. Fourth toe lamellae: 28–34. Third toe lamellae: 23–24. In six females: SVL (mean ± SD (range)): 95.9 ± 6.9 (84.0– 101.6). Axilla –groin distance: 44.7 ± 4.5 (36.7–47.6). Head length: 19.5 ± 1.0 (17.7–20.3). Head width: 16.7 ± 0.8 (15.2–17.3). Foot length: 27.7 ± 0.2 (27.5 –28.0). Tibial length: 18.5 ± 0.7 (17.7–19.7). Hand length: 28.3 ± 0.9 (26.8 –29.0). Midbody scales: 77.4 ± 3.8 (72–82). Dorsal scales: 72.8 ± 2.1 (70–76). Ventral scales: 108.2 ± 4.5 (105–116). 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: 27–32. Third toe lamellae: 21–23. 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 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.
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. 4View FIGURE 4, 5View FIGURE 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 40–90 % 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 .
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 known only from the nearby Tromen Volcano, is syntopic in same areas ( Avila and Perez, 2006), as well as Phymaturus verdugo (Avila et al., 2007 b). 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. 6View FIGURE 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.
|MACN 38986||MLPS 2594||MLPS 2595||LJAMM LJAMM BYU 6173 6172 12592||MACN 38987||MLPS 2592||MLPS 2593|
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