Two new species of the Liolaemuselongatus-kriegi complex (Iguania, Liolaemidae) from Andean highlands of southern Chile Author Troncoso-Palacios, Jaime Author Diaz, Hugo A. Author Esquerre, Damien Author Urra, Felix A. text ZooKeys 2015 500 83 109 http://dx.doi.org/10.3897/zookeys.500.8725 journal article http://dx.doi.org/10.3897/zookeys.500.8725 1313-2970-500-83 D250C8BF532A4767895FF1FA36D368B7 Taxon classification Animalia Squamata Liolaemidae Liolaemus zabalai sp. n. Fig. 6 Liolaemus kriegi , Donoso-Barros, 1974. Bol. Soc. Biol. Concepcion , 47: 287. Liolaemus kriegi (in part), Cei, 1986. Mus. Reg. Scien. Nat. Torino, 4: 230. Liolaemus sp?, Torres-Perez , 1997. Not. Biol., 5(4): 146. Liolaemus kriegi (in part), Pincheira-Donoso, 2001. Not. Mens. Mus. Nac. Hist. Nat., Chile, 346: 11. Liolaemus sp. A, Morando et al., 2003. Syst. Biol., 52: 179. Liolaemus kriegi (in part), Pincheira-Donoso & Nunez , 2005. Pub. Oc. Mus. Nac. Hist. Nat., Chile, 59: 289. Liolaemus kriegi (in part), Mella, 2005. Guia Camp. Rep. Chil. Zon. Cent., p. 64. Liolaemus sp. A, Medina et al., 2013. Cuad. Herp. 27(1): 27. Liolaemus sp. A, Medina et al., 2014. Biol. J. Linnean Soc. 113: 256. Holotype. SSUC Re 602 (Fig. 6). Near Los Barros, Laja Lagoon, Biobio Region, Chile. ( 37°31'S - 71°15'W , 1460 m). Collected by J. Troncoso-Palacios, F. Urra and H. Diaz . 07/01/2014. Figure 6. Liolaemus zabalai sp. n. A, B Holotype, male C, D Paratype, female E Paratype, male F Paratype, female. All from the type locality, near Los Barros, Laja Lagoon, Biobio Region, Chile. Paratypes. SSUC Re 598. Adult male. SSUC Re 597, 599, 600-01. Four adult females. The same data as the holotype (Figs 6 and 8). MZUC 35607, 39567. One male and one female. Malleco, Antuco Volcano, Los Barros. Unknown coll. Etymology. This species is named after Patricio Zabala, collection manager of the "Coleccion de Flora y Fauna Patricio Sanchez Reyes, Pontificia Universidad Catolica de Chile" (SSUC). We dedicate this species to him because of his support of herpetological research in Chile, allowing us to review and deposit material in SSUC, and especially for his friendship. Diagnosis. Liolaemus zabalai belongs to the kriegi clade of the elongatus - kriegi complex and is closely related to some undescribed species: Liolaemus sp. C and Liolaemus sp. D; being more distant from the currently described species Liolaemus buergeri , Liolaemus kriegi and Liolaemus tregenzai (Fig. 7). According to Medina et al. (2014) , in regards to the species of the kriegi clade Liolaemus zabalai is sympatric only with Liolaemus tregenzai at the Copahue Volcano. Figure 7. Phylogenetic position of Liolaemus zabalai sp. n. in the kriegi clade, based on cytochrome-b (cyt-b) locus according to Medina et al. (2014) . With respect to the species species of the kriegi clade, Liolaemus zabalai differs from Liolaemus tregenzai because the latter has 71-85 midbody scales and the males have no precloacal pores ( Pincheira-Donoso and Scolaro 2007 ), whereas Liolaemus zabalai has 90-104 midbody scales and the males have 3-5 precloacal pores. In addition, the green-bluish ventral color of Liolaemus tregenzai is completely absent in Liolaemus zabalai . The uncorrected pairwise difference (cyt-b) between the species is 3.09% ( Medina et al. 2014 ). Liolaemus zabalai differs from Liolaemus kriegi in that the latter reaches 101.1 mm SVL, has reddish cloacal coloration in both sexes and has an unringed tail ( Avila et al. 2003 ), whereas Liolaemus zabalai is smaller (max. SVL = 92.0 mm), has yellowish cloacal coloration in both sexes and has a ringed tail (in specimens with original tails). The uncorrected pairwise difference between these species is 3.79% ( Medina et al. 2014 ). Liolaemus zabalai differs from Liolaemus buergeri in that the latter has fewer dorsal scales (78-91; x = 84.1 +/- 4.4, n = 14) than Liolaemus zabalai (86-96; x = 89.4 +/- 3.2, n = 8) ( Mann-Whitney U = 19.5; P = 0.01, DF = 20). Liolaemus zabalai has more loreal scales between the nasal and the subocular (4-6; x = 4.3 +/- 0.6, n = 8) than Liolaemus buergeri (3-4; x = 3.3 +/- 0.5, n = 14) ( Mann-Whitney U = 11.0; P <0.01, DF = 20). Also, Liolaemus buergeri has a vertebral stripe on the tail, whereas Liolaemus zabalai has a ringed original tail. The limbs in Liolaemus zabalai are black with dispersed light brown spots, whereas Liolaemus buergeri has brown limbs with dispersed black spots (Fig. 8). Liolaemus zabalai and Liolaemus buergeri share basically the same dorsal coloration pattern, but this is noticeably more marked and darker in Liolaemus zabalai (Fig. 8, see discussion). Based on the cyt-b locus, the uncorrected average pairwise difference between Liolaemus zabalai and Liolaemus buergeri is 2.94% ( Medina et al. 2014 ), greater than the values reported for other Liolaemus widely accepted as valid species (see discussion). Also, Liolaemus zabalai can vocalize, a feature only documented for Liolaemus chiliensis in the entire genus Liolaemus ( Labra et al. 2013 ). Finally, although the ranges overlap, males of Liolaemus buergeri have 3-4 (x = 3.3) precloacal pores, whereas males of Liolaemus zabalai have 3-5 (x = 3.9) precloacal pores ( Medina et al. 2014 ). Figure 8. Comparison of the dorsal color pattern. A Liolaemus zabalai sp. n. with marked color pattern and B Liolaemus buergeri , diffuse color pattern. Compared to the other species of the elongatus - kriegi complex that occur near the known distribution of Liolaemus zabalai , the new species may be diagnosed as follows. Males of Liolaemus zabalai have precloacal pores, whereas males of Liolaemus flavipiceus and Liolaemus punmahuida lack them (Table 3). Liolaemus zabalai is larger than Liolaemus scorialis ; and Liolaemus zabalai has more midbody scales than Liolaemus antumalguen , Liolaemus burmeisteri and Liolaemus choique (Table 3). Description of the holotype. Adult male. SVL: 90.3 mm. Tail length: 92.3 mm (autotomized). Axilla-groin length 39.7 mm. Head length (from the posterior border of the auditory meatus to the tip of the snout) 22.2 mm. Head width (distance between the two ear openings) 16.5 mm. Head height (at the level of ear openings) 11.7 mm. Forelimb length 28.5 mm. Hindlimb length 47.1 mm. Foot length 23.4 mm. Rostral scale wider (4.5 mm) than high (2.2 mm). Two postrostrals. Four internasals. Heptagonal interparietal scale, with a central, small, and whitish central spot marking the position of the parietal eye. Interparietal smaller than right parietal, but bigger than left parietal, surrounded by eight scales: nine scales between the interparietal and the rostral; 14 scales between occiput and rostral; orbital semicircle complete on both sides (formed by 13 scales); 5 supraoculars on both sides; seven superciliary scales. Frontal area is divided into six scales (three posterior, one anterior-left, two anterior-rigth); 2 scales between nasal and canthal; preocular separated from the lorilabials by one loreal scale; nasal in contact with the rostral, surrounded by six scales. There is one row of lorilabials between the supralabials and the subocular. Seven supralabials, the fourth is curved upward without contacting the subocular. Five infralabial scales. The mental scale is pentagonal and is in contact with four scales. Four pairs of postmental shields, the second is separated by two scales. Temporal scales are subimbricated and smooth or slightly keeled. Nine temporal scales between the level of superciliary scales and the rictal level. Two projected scales on the anterior edge of the ear, which are small and do not cover the auditory meatus. There is no differentiated auricular scale. Forty-two gulars between auditory meatus. Well developed "Y" shaped lateral neck fold with antehumeral and posthumeral folds developed. Dorsolateral fold slightly developed. Midbody scales 90. Dorsal scales on the vertebral zone are lanceolate to rounded, subimbricate, keeled and without mucrons. Dorsal scales on the paravertebral fields are more rounded, subimbricate, smooth or with less developed keels, without mucrons and there are interstitial granules between them. Dorsal scales are smaller than the ventral scales. Dorsal scales 86. Ventral scales are rhomboidal, smooth, subimbricate, and with few interstitial granules. Ventral scales 122. There are three precloacal pores. The suprafemoral scales are rhomboidal, imbricate, and smooth or keeled. Infrafemoral scales are lanceolate to rhomboidal, smooth, and subimbricate and with few interstitial granules. Supra-antebrachials scales are rhomboidal to rounded, subimbricate, and keeled or smooth. Infra-antebrachials are rounded to rhomboidal, subimbricate, and smooth. The dorsal scales of the tail are lanceolate to rectangular, subimbricate, keeled or smooth and with few interstitial granules. The ventral scales of the tail vary from lanceolate to triangular, and are subimbricate and smooth. Lamellae of the fingers: I: 11, II: 16, III: 20, IV: 22 and V: 15. Lamellae of the toes: I: 12, II: 16, III: 21, VI: 27 and V: 18. Color of the holotype in life. Black head, with some light brown spots on the supraocular and snout areas. The scales located behind the orbital semicircles are light brown; but the interparietal scale, parietal scales and the scales in contact with the parietal scales are black. Superciliary scales are light brown with black spots. Temporal scales are light brown; cheeks light gray with some black spots. Subocular is gray with a black vertical line on the middle. Background color of the dorsum is light brown. Wide occipital band on the dorsum, formed by twelve transverse black bars. Very few whitish scales dispersed on the dorsum. Black lateral band bearing a few dispersed whitish scales, running from the tip of snout to the groin. Flanks below lateral band are light brown. Limbs black with dispersed light brown spots. Tail light brown with inconspicuous vertebral stripe in the regenerated zone; occipital black band ends in the first fifth of the tail, remainder with some dispersed black spots and a black vertebral stripe. Throat, belly and ventral surfaces of limbs whitish with dispersed inconspicuous dark dots. Rear portion of the belly and the thighs are yellowish. Ventrally, tail is whitish with a dark gray ventral stripe and diffuse dark gray rings from the cloaca to the midpoint of the tail. Precloacal pores orange. Variation. In three males: SVL: 72.6-90.3 mm. Axilla-groin distance: 32.7-38.6 mm. Head length: 17.6-22.2 mm. Head width: 14.2-16.5 mm. Head height: 9.2-11.7 mm. Foot length: 21.5-23.0 mm. Leg length: 42.1-47.2 mm. Arm length: 24.6-28.5 mm. Tail length: 102.0 mm in one specimen (autotomized in the rest). In three females: SVL: 71.8-90.2 mm. Axilla-groin distance: 32.9-42.7 mm. Head length: 17.9-19.5 mm. Head width: 13.9-16.6 mm. Head height: 9.4-11.1 mm. Foot length: 20.6-24.2 mm. Leg length: 41.5-48.8 mm. Arm length: 24.8-29.4 mm. Tail length: 105-115 mm (in two specimens without autotomized tails). The variation of the scalation in Liolaemus zabalai is as follows. Midbody scales: 90-104 (x = 94.3 +/- 4.8). Dorsal scales: 86-96 (x = 89.4 +/- 3.2). Ventral scales 116-122 (x = 119.5 +/- 2.1). Fourth finger lamellae: 19-22 (x = 20.9 +/- 1.0). Fourth toe lamellae: 26-27 (x = 26.8 +/- 0.5). Supralabial scales: 6-7 (x = 6.6, +/- 0.5). Infralabial scales: 4-5 (x = 4.6 +/- 0.5). Interparietal scale pentagonal, hexagonal or heptagonal, bordered by 5-8 scales (x = 7.3 +/- 1.1). Precloacal pores in males: 3-4. There is slight sexual dichromatism; males are slightly darker than females. In general, all specimens have the pattern and color described for the holotype. One female has rusty-colored scales dispersed on the flanks, paravertebral fields and groin. In all specimens, the ventral surface of the throat, belly and limbs are whitish with dark marked or inconspicuous dots dispersed; there is a fragmented midventral stripe on the belly of two specimens. Males and females have a yellowish coloration in the posterior portion of the belly and the thighs (faint in some females). The tail has black rings, marked or diffuse, with a fragmented vertebral stripe in all specimens with complete original tails. Males have orange precloacal pores. The coloration and pattern of the juveniles are unknown. Distribution and natural history. To our knowledge, in Chile this species is only found in the surroundings of the Laja Lagoon. The type locality is near Los Barros, Laja Lagoon, Biobio Region, Chile ( 37°31'S - 71°15'W , 1460 m, Fig. 9); but we also saw specimens (not collected) on the road to the Laja Lagoon at two localities ( 37°23'S - 71°23'W , 1320 m; 37°23'S - 71°22'W , 1390 m). The new species was found inhab iting areas of sandy soil with rocks of small and medium size. The vegetational cover is low, consisting mainly of Ephedra chilensis . It is an abundant lizard of saxicolous habits. This species was observed active between 11h00 and 18h00, taking refuge in cavities under the rocks. Near Los Barros, at its upper altitudinal limit (1460 m), this species was found in syntopy with Diplolaemus sexcinctus . At the lower altitudinal limit (1320 m), it was found in syntopy with Liolaemus scorialis , Phymaturus vociferator and Diplolaemus sexcinctus . Two specimens of Liolaemus zabalai vocalized (squealed) in several occasions in response to the manipulation. Figure 9. View of the type locality of Liolaemus zabalai sp. n. Liolaemus zabalai is also found in Argentina (where it has been called " Liolaemus sp. A") at several localities in Neuquen Province ( Morando et al. 2003 , Medina et al. 2013 , 2014 ). An analysis of the intestinal contents performed on one specimen, showed that this species is omnivorous, but feeds mainly on plants. At the time of capture (January) the females had no embryos, but three had several small oocytes.