A new species of Liolaemus related to L. nigroviridis from the Andean highlands of Central Chile (Iguania, Liolaemidae) Author Troncoso-Palacios, Jaime Author Elorza, Alvaro A. Author Puas, German I. Author Alfaro-Pardo, Edmundo text ZooKeys 2016 555 91 114 http://dx.doi.org/10.3897/zookeys.555.6011 journal article http://dx.doi.org/10.3897/zookeys.555.6011 1313-2970-555-91 E1446FEB806142E79C952CD6E56E6E9C Taxon classification Animalia ORDO FAMILIA Liolaemus uniformis sp. n. Fig. 2A, B Liolaemus altissimus altissimus (in part?), Mella. 2005 , Guia Camp. Rep. Chil. Zon. Cent., p. 38. Liolaemus monticola ?, Nunez et al. 2010 , Bol. Mus. Nac. Hist. Nat., p. 57. Holotype. SSUC Re 674. Adult male. Collected in the west shore of the Chepical Lagoon ( 32°15'S - 70°30'W ), approximately 30 km NE Alicahue, San Felipe de Aconcagua Province, Valparaiso Region, Chile. Collectors: J. Troncoso-Palacios and E. Alfaro. December, 2012. Paratypes (Fig. 2C, D, E, F). SSUC Re 675, male. SSUC Re 676-78, three females. SSUC Re 679, juvenile. The same data as the holotype. Figure 2. Liolaemus uniformis sp. n. A, B Holotype, male C, D Paratype, female E Paratype, male F Paratype, juvenile (unknown sex). All from the type locality. Etymology. The species name " uniformis " (Latin) refers to the lack of dorsal pattern and uniform color found for both males and females. Diagnosis. Liolaemus uniformis is larger than Liolaemus constanzae ( Mann-Whitney U = 0.5, P <0.01, Table 1). Liolaemus constanzae has sexual dichromatism, a feature absent in Liolaemus uniformis . Males of Liolaemus constanzae have a black vertebral line and black spots on the paravertebral fields (Fig. 3A), whereas Liolaemus uniformis has no dorsal pattern. Additionally, the southern distributional limit of Liolaemus constanzae in Agua Verde, Antofagasta Region, Chile ( Ortiz 1975 ), is more than 750 km north of the type locality recorded for Liolaemus uniformis . Figure 3. Chilean species of the nigroviridis group (with the exception of Liolaemus nigroviridis ), ordered from north to south. A Liolaemus constanzae , male from vicinity of San Pedro (picture by JTP) B Liolaemus melanopleurus , male from Atacama (picture by JTP) C Liolaemus isabelae , male from Montandon (picture by JTP) D Liolaemus juanortizi , unknown sex specimen from road to Negro Francisco (picture by F. de Grotee) E Liolaemus lorenzmuelleri , unknown sex specimen from Embalse La Laguna (picture by A. Labra) F Liolaemus maldonadae , male from vicinity of Alcohuaz (picture by JTP). Table 1. Scalation and morphological characteristics for the species of the nigroviridis group. Juvenile specimens examined are excluded. M = males; F = females. (*) Taken from Navarro and Nunez (1993) . (**) Examined specimen plus Young-Downey and Moreno (1991) data. (***) Taken from Young-Downey and Moreno (1991). (****) Counted only for one specimen.

Liolaemus constanzae (M = 14, F = 13)	Liolaemus isabelae (M = 4)	Liolaemus juanortizi (M = 1)	Liolaemus lorenzmuelleri (M = 3, F = 5)	Liolaemus maldonadae (M = 3)	Liolaemus melanopleurus (M = 2)	Liolaemus nigroviridis (M = 9, F = 4)	Liolaemus uniformis sp. n. (M = 2, F = 3)
SVL

Liolaemus uniformis differs from Liolaemus isabelae (Fig. 3C), because in the latter the nasal and the rostral scales are in contact only in 25% of specimens, whereas in Liolaemus uniformis , these scales are always in contact. Males of Liolaemus isabelae have black ventral coloration, a yellow dorsal color with a black vertebral line, black bars in the paravertebral fields, and a black lateral band, or some males have a completely black dorsal color; all traits that are absent in Liolaemus uniformis . Additionally, the southern distributional limit of Liolaemus isabelae in Salar de Pedernales, Atacama Region, Chile ( Pincheira-Donoso and Nunez 2005 ) is more than 650 km north of the type locality recorded for Liolaemus uniformis . Liolaemus uniformis resembles Liolaemus lorenzmuelleri (Fig. 3E) and Liolaemus juanortizi (Fig. 3D), species suggested as conspecific ( Pincheira-Donoso and Nunez 2005 ). However, the dorsal scales in Liolaemus lorenzmuelleri and Liolaemus juanortizi are noticeably larger than those of Liolaemus uniformis , and have a distinct "ovoid" shape. Liolaemus uniformis has more dorsal scales (60.0 +/- 2.9) than Liolaemus lorenzmuelleri (48.4 +/- 4.2) (t = -5.4, P <0.01). On the other hand, while only one specimen of Liolaemus juanortizi was examined, this one has 52 dorsal scales, which is below of the range for Liolaemus uniformis (Table 1). Liolaemus uniformis has more midbody scales (60.4 +/- 1.7) than Liolaemus lorenzmuelleri (54.9 +/- 4.5) (t = 2.6, P <0.05) and Liolaemus juanortizi (56.7 +/- 2.1) (t = 3.2, P <0.05). Liolaemus lorenzmuelleri has a dark vertebral line and dark transversal lines running from the paravertebral fields to the flanks, whereas Liolaemus uniformis has no dorsal pattern. The dorsal pattern of Liolaemus juanortizi is very similar to Liolaemus lorenzmuelleri , but some specimens have a black ventral coloration, a black lateral band, and the lack of a dark vertebral line, whereas Liolaemus uniformis has no black ventral color or black lateral band. Additionally, the southern distributional limit of Liolaemus lorenzmuelleri (Embalse La Laguna, Coquimbo Region, Chile) is more than 240 km north of the type locality recorded for Liolaemus uniformis ; and the southern distributional limit of Liolaemus juanortizi in Quebrada Contrabando, Atacama Region, Chile (MNHNCL collection catalog, unpublished) is more than 520 km north of the type locality recorded for Liolaemus uniformis . Liolaemus uniformis differs from Liolaemus melanopleurus (a species with only three known specimens from an undetermined location, Fig. 3B) in that the latter has a blue-gray dorsal coloration ( Philippi 1860 ) and a black lateral band running from the axilla to the midbody, features absent in Liolaemus uniformis . Although the type locality of Liolaemus melanopleurus is undetermined, the syntypes were collected by Philippi in his journey through the Atacama Desert, between the vicinities of Copiapo (27°23'S) and San Pedro de Atacama (22°54'S), more than 530 km north of the type locality recorded for Liolaemus uniformis . Liolaemus uniformis differs from Liolaemus maldonadae (Fig. 3F), because males of the latter have a yellowish or reddish dorsal color with black transverse dorsal and ventral bars and black lateral band, whereas Liolaemus uniformis has no dorsal pattern or black trans verse ventral bars. Dorsal scales in Liolaemus maldonadae are noticeably larger than found in Liolaemus uniformis , and they have an "ovoid" shape. Dorsal and ventral scale counts in Liolaemus maldonadae do not overlap with the same scale counts in Liolaemus uniformis (Table 1). Finally, the southern distributional limit of Liolaemus maldonadae in Los Molles ( Nunez et al. 1991 ) is more than 150 km north of the type locality of Liolaemus uniformis . Liolaemus uniformis is found in sympatry with Liolaemus nigroviridis (Fig. 4), but is larger than Liolaemus nigroviridis ( Mann-Whitney U = 8.0, P <0.05, Table 1). Liolaemus uniformis also has more dorsal scales (60.0 +/- 2.9) than Liolaemus nigroviridis (49.4 +/- 2.7) (t = 7.4, P <0.01). Liolaemus nigroviridis has strongly mucronated dorsal scales, whereas Liolaemus uniformis has no mucrons (Fig. 5). Liolaemus nigroviridis has sexual dichromatism, absent in Liolaemus uniformis . Males of Liolaemus nigroviridis have a bluish or yellowish green dorsal color with black reticulation, and females have a brown dorsal color with a black lateral band, black vertebral line, and black paravertebral spots. In contrast, Liolaemus uniformis has a brown dorsal color without any pattern. Figure 4. Variation in Liolaemus nigroviridis . A Male from Farellones (picture by H. Diaz ) B Male from Carpa Mountain (picture by JTP) C Male from Provincia Mountain (picture by JTP) D Female from Juncal (picture by JTP). Figure 5. Dorsal scales, 8 mm width of view. A Male of Liolaemus uniformis sp. n. B Liolaemus nigroviridis . Molecular data show that Liolaemus uniformis is not closely related to Liolaemus monticola (Fig. 1). Moreover, Liolaemus monticola is smaller (maximum SVL = 65.6 mm) than Liolaemus uniformis (max. SVL = 89.1 mm) (t = 3.9, P <0.01) according to our samples, and although Pincheira-Donoso and Nunez (2005) recorded a max. SVL = 67.3 mm for Liolaemus monticola , the difference between both species is marked. Moreover, Liolaemus monticola exhibit a characteristic black lateral band between the axilla and midbody (diffuse in females), and males have white dots dispersed on the dorsum and a series of small black spots on the dorsum (Fig. 6). All these traits are absent in Liolaemus uniformis . The upper altitudinal limit of Liolaemus monticola distributions is 2000 m a.s.l. ( Espinoza et al. 2004 , Fuentes and Ipinza 1979 ), whereas Liolaemus uniformis has a lower altitudinal distribution limit of 2820 m a.s.l. Figure 6. Variation in species probably confused with Liolaemus uniformis sp. n. A Liolaemus monticola from Salto de Apoquindo (picture by JTP) B Liolaemus monticola from La Cruz Mountain (picture by J. Abarca-Diaz ) C Liolaemus monticola from Provincia Mountain (picture by JTP) D Liolaemus bellii from La Parva (JR Martini) E Liolaemus bellii from Lagunillas (picture by JTP) F Liolaemus bellii from San Ramon Mountain (picture by JTP). Molecular data show that Liolaemus uniformis is not closely related to Liolaemus bellii (Fig. 1). Moreover, Liolaemus bellii is smaller (maximum SVL = 80.8 mm) than Liolaemus uniformis (max. SVL = 89.1 mm) (t = 2.7, P <0.05). Liolaemus uniformis has more midbody scales (60.4 +/- 1.7) than Liolaemus bellii (52.9 +/- 2.6) (t = 6.1, P <0.01); more dorsal scales (60.0 +/- 2.9) than Liolaemus bellii (43.3 +/- 3.1) (t = 10.2, P <0.01); and more ventral scales (96.2 +/- 4.8) than Liolaemus bellii (89.7 +/- 4.6) ( Mann-Whitney U = 10.5, P <0.05). Dorsal scales in Liolaemus bellii are strongly keeled and mucronated, whereas there are no mucrons in Liolaemus uniformis . Moreover , Liolaemus bellii exhibit a characteristic series of black dorsal "W" o "V" shaped spots (Fig. 6), whereas Liolaemus uniformis has no dorsal pattern. Description of the holotype. Adult male. SVL = 84.7 mm. Horizontal diameter of the eye: 4.3 mm. Subocular length: 4.5 mm. Length of the fourth supralabial: 4.1 mm. Head length (from the posterior border of the auditory meatus to the tip of the snout): 22.1 mm. Head height (distance between the two ear openings): 10.4 mm. Head width (at the level of ear openings): 15.8 mm. Neck width: 12.4 mm. Interorbital distance: 6.3 mm. Ear-eye distance: 7.5 mm. Internarine distance: 3.8 mm. Ear width: 2.5 mm. Ear height: 3.5 mm. Axillary-groin distance: 34.9 mm. Body width: 24.7 mm. Forelimb length: 25.7 mm. Hindlimb length: 46.1 mm. Length of the right hand: 10.4 mm. Length of the right foot: 22.4 mm. Tail length (not autotomized): 132.4 mm, with relation tail length/SVL = 1.56. Pentagonal rostral scale, wider (4.2 mm) than high (1.4 mm). Two postrostrals. Four internasals. Heptagonal interparietal, with a central, small, and whitish central spot marking the position of the parietal eye. Interparietal smaller than the parietals, surrounded by seven scales. Seven scales between the interparietal and rostral. Thirteen scales between the occiput and the rostral. Orbital semicircle incomplete on the right side and complete on the left (formed by thirteen scales). Three supraoculars on the left side and four on the right. Six superciliary scales. Frontal area divided into three scales (1 posterior and 2 anterior). Preocular separated from the lorilabials by one loreal scale. Two scales between nasal and canthal. Nasal in contact with the rostral, surrounded by six scales. One row of lorilabials between the supralabials and subocular. Four lorilabials in contact with the subocular. Six supralabials, the fourth is curved upward without contacting the subocular. Four infralabials scales. Pentagonal mental scale, in contact with four scales. Four pairs of post-mental shields, the second is separated by two scales. Temporal scales smooth or slightly keeled, imbricated. Six temporal scales between the level of superciliary scales and the rictal level. Four scales on the anterior edge of the ear, which do not cover the auditory meatus. Poorly differ entiated auricular scale, pentagonal and located at the upper part of the meatus. Thirty gulars between the auditory meatus. Lateral neck fold is "Y" shaped. Ventrolateral fold running from the neck to the groin. Dorsolateral fold slightly developed, running from the ear to the base of the tail. Midbody scales: 60. Dorsal scales are lanceolated, imbricated, keeled (without mucrons), with few interstitial granules. Dorsal smaller than the ventrals. Dorsal scales: 58. Ventrals scales are polymorphic (rounded, rhomboidal, pentagonal or hexagonal) smooth, imbricated, without interstitial granules. Ventrals: 91. Three precloacal pores. Supra-femoral scales lanceolate, imbricated, smooth or keeled. Infra-femoral scales lanceolate or rounded, smooth and imbricated. Supra-antebrachials scales are rounded or lanceolated, imbricated and smooth or keeled. Infra-antebrachials are rounded, imbricated and smooth. Dorsal scales of tail are pentagonal or rhomboidal, imbricated and keeled. Ventral tail scales are rounded or rhomboidal, smooth and imbricated. Lamellae of the fingers: I: 9, II: 13, III: 20, IV: 20 and V: 13. Lamellae of the toes: I: 11, II: 15, III: 21, VI: 27 and V: 17. Color of the holotype in life. The specimen is notable for its lack of pattern and uniform color. The head is brown and darker than the body. There are several white dots dispersed over the head and cheeks. The dorsum is coppery brown and has a few white-spotted scales that did not form a pattern. The subocular is brown and crossed by three white, vertical lines. The dorsal surface of the tail is light brown and without a pattern. The limbs are a dorsal-brown, similar to the dorsal surface, with white dots dispersed on the forelimbs and white transversal lines on the hindlimbs. The flanks are whitish with abundant dark brown scales. Ventrally, the hands, feet, thighs, vent, and tail are yellowish. The belly is whitish with dark dispersed spots and a dark ventral stripe. The throat is whitish with a dark thick reticulation. The precloacal pores are orange. Variation in the type series. Males are larger and more corpulent than females. In two males: SVL: 84.7-89.1 mm. Axilla-groin distance: 34.9-37.8 mm. Head length: 21.9-22.1 mm. Head width: 15.8-16.3 mm. Head height: 10.4-11.2 mm. Leg length: 45.4-46.1 mm. Arm length: 25.0-25.8 mm. Tail length: 132.4 mm in one specimen, with relation tail length/SVL = 1.56 (autotomized in the other). In three females: SVL: 67.7-73.1 mm. Axilla-groin distance: 33.1-35.7 mm. Head length: 17.8-20.0 mm. Head width: 11.8-13.3 mm. Head height: 7.5-8.3 mm. Leg length: 32.0-34.8 mm. Arm length: 19.2-21.3 mm. Tail length: 98.1 mm in one specimen, with relation tail length/SVL = 1.45 (autotomized in other). The variation of the scalation in Liolaemus uniformis is as follows. Midbody scales: 58-62 (60.4 +/- 1.7). Dorsal scales: 56-63 (60.0 +/- 2.9). Ventral scales 91-102 (96.2 +/- 4.8). Fourth finger lamellae: 17-20 (19.0 +/- 1.4). Fourth toe lamellae: 25-27 (26.4 +/- 0.9). Supralabial scales: 6. Infralabial scales: 4-5 (4.4 +/- 0.6). Interparietal scale pentagonal, hexagonal or heptagonal, bordered by 5-7 scales (6.0 +/- 0.7). Nasal and rostral always in contact. Precloacal pores in males: 3. Precloacal pores are absent in females. In general, all specimens have the pattern and color described for the holotype, with slight variations in shade. The male paratype has a dark brown throat. Two females have inconspicuous dark rings and an inconspicuous vertebral stripe on the dorsal surface of the tail. Also, two females have an olive hue on the snout. One female has a very inconspicuous series of dark crossbars on the paravertebral fields, which, while difficult to count, approximated eight. The juvenile has a similar pattern and color as the holotype, but it has an inconspicuous and fragmented dark vertebral line and inconspicuous dark spots on the paravertebral fields. Distribution and natural history. This species is currently only known from the type locality in the surroundings of the Chepical Lagoon, approximately 30 km NE of Alicahue, in the San Felipe de Aconcagua Province, Valparaiso Region, Chile (Fig. 7). Specimens were collected on the west shore of the Chepical Lagoon ( 32°15'S - 70°30'W , 3050 m a.s.l.). This new species was found inhabiting rocky areas with little shrubby vegetation composed mainly of high-Andean forbs, such as Chuquiraga oppositifolia and Azorella sp. (Fig. 8). This lizard was found in abundance and was observed to have saxicolous habits. It was active between 9:00 h and 18:00 h and took refuge under rocks. Moreover, this species was found in syntopy with Phymaturus alicahuense Nunez , Veloso, Espejo, Veloso, Cortes & Araya 2010. Specimens were also observed at lower altitudes ( 32°16'S - 70°30'W , 2820 m a.s.l.) in similar environments, altitudes at which this species was found in sympatry with a few specimens of Liolaemus nigroviridis . Figure 7. Distributional map for Liolaemus uniformis sp. n. along with geographically proximate species of the nigroviridis group. Red star: Liolaemus uniformis sp. n., Chepical Lagoon, type locality. Green circles: Liolaemus nigroviridis (1 = Manque, 2 = El Arpa, 3 = Juncal, 4 = Riecillo, 5 = La Campana, 6 = El Roble, without number = near Chepical Lagoon). Pink squares: Liolaemus maldonadae (1 = near Alcohuaz, 2 = Los Molles). Yellow triangles: Liolaemus lorenzmuelleri (1 = Banos del Toro, 2 = Embalse La Laguna). Figure 8. View of the type locality of Liolaemus uniformis sp. n., a high Andean environment. One of the collected specimens had a yellow flower inside of its mouth. An analysis of intestinal contents showed that Liolaemus uniformis is omnivorous; plant and Hymenoptera remains were found. A large quantity of nematodes from an unidentified species was found in the intestines. While the reproductive mode is yet unknown, at the time of sampling (December) no evidence of embryos was found but one female had several small oocytes. Comparisons with the reproductive modes of other species in the nigroviridis group would not be helpful as there is little available data. It is known that Liolaemus nigroviridis is viviparous ( Donoso-Barros 1966 ) and Liolaemus lorenzmuelleri is oviparous ( Cortes et al. 1995 ). Pincheira-Donoso and Nunez (2005) reported that Liolaemus maldonadae and Liolaemus isabelae are viviparous, but the source of this information is unclear (see Lobo et al. 2010 :4) since the reproductive mode was not mentioned in the original descriptions ( Navarro and Nunez 1993 , Nunez et al. 1991 ).