Two new Liolaemus lizards from the Andean highlands of Southern Chile (Squamata, Iguania, Liolaemidae) Author Troncoso-Palacios, Jaime Author Diaz, Hugo A. Author Puas, German I. Author Riveros-Riffo, Edvin Author Elorza, Alvaro A. text ZooKeys 2016 632 121 146 http://dx.doi.org/10.3897/zookeys.632.9528 journal article http://dx.doi.org/10.3897/zookeys.632.9528 1313-2970-632-121 0910B0A23CA94C37B6AD1DEB92ADDFFD Taxon classification Animalia Squamata Liolaemidae Liolaemus janequeoae sp. n. Figure 4 Holotype. SSUC Re 712 (Fig. 4). Male collected at Laguna Verde ( 38°12'S - 71°44'W , 1397 masl), approximately 13.5 km NW of the summit of the Tolhuaca Volcano, Araucania Region, Chile. Collected by J. Troncoso-Palacios and Edvin Riveros-Riffo. January 15, 2016. Figure 4. Liolaemus janequeoae sp. n. A and B Holotype, male C and D Paratype, female with dorsal black dots E and F Paratypes, typical females. Paratypes. SSUC Re 713-14. Two females (Fig. 4). Same data as the holotype. SSUC Re 715. Female. Collected at the locality of the holotype by Edvin Riveros-Riffo. February 18, 2015. SSUC Re 649-51, three females. Collected at the locality of the holotype by J. Troncoso-Palacios, F. Urra and H. Diaz . January 5, 2014 (Fig. 4). Diagnosis. Liolaemus janequeoae belongs to the Liolaemus elongatus clade. This species is characterized by 1) small size (maximum snout vent length = 69.6 mm), 2) lack of dorsal pattern, 3) high number of midbody scales (82-98), 4) precloacal pores present in males, and 5) absence of dark rings on the tail. We provide a differential diagnosis with regards to all species currently considered to be members of this clade, plus Liolaemus scorialis Troncoso-Palacios Diaz , Esquerre & Urra, 2015, the assignment of which is under study, but probably is related to the Liolaemus elongatus clade ( Troncoso-Palacios et al. 2015 ). Table 1 summarizes some of the diagnostic traits. Based on seven specimens. Table 1. Scale count and morphological characteristics for Liolaemus janequeoae sp. n. and the geographically proximate species of the Liolaemus elongatus clade plus Liolaemus leftrarui sp. n. Juvenile specimens examined are excluded. Source of data for non-examined species are: Liolaemus antumalguen ( Avila et al. 2010 ), Liolaemus burmeisteri ( Avila et al. 2012 ) and Liolaemus lonquimayensis ( Escobar-Huerta et al. 2015 ). M = males; F = females.

Liolaemus antumalguen	Liolaemus burmeisteri	Liolaemus carlosgarini (M = 4, F = 5)	Liolaemus cf. elongatus (M = 3, F = 3)	Liolaemus janequeoae sp. n. (M = 1, F = 6)	Liolaemus leftrarui sp. n. (M = 3, F = 4)	Liolaemus lonquimayensis	Liolaemus scorialis (M = 8, F = 3)
SVL

Liolaemus janequeoae is closely related to Liolaemus elongatus . However, Liolaemus janequeoae is smaller (maximum SVL = 69.6 mm, n = 7 adults, vs. max. SVL = 94.7 mm) and has more midbody scales (82-98 vs. 68-87) than Liolaemus elongatus from Argentina (Table 1). Dorsal color pattern in Liolaemus elongatus is highly variable from vertebral and lateral dark bands to complete melanism, whereas Liolaemus janequeoae never has black spots (only small black dots in one female). Interestingly, SVL of Liolaemus cf. elongatus from Llaima, Chile (SVL = 68.4 +/- 2.9 mm), is not significantly different compared with the SVL of Liolaemus janequeoae (SVL = 65.3 +/- 3.4 mm); but head height is lower in Liolaemus janequeoae than in Liolaemus cf. elongatus (6.8 +/- 0.5 mm vs 8.3 +/- 0.7 mm) (t = -4.6, DF = 11, P <0.01); the head is wider in Liolaemus cf. elongatus than in Liolaemus janequeoae (12.7 +/- 0.9 mm vs 11.0 +/- 0.4 mm) ( Mann-Whitney U = 0.001, P <0.01); Liolaemus janequeoae has more midbody scales than Liolaemus cf. elongatus (82-98 vs. 76-88) (t = 3.0, DF = 11, P <0.05), more dorsal scales (77-89 vs. 67-73) (t = 7.7, DF = 11, P <0.01) and more ventral scales (124-132 vs. 119-129) (t = 2.5, DF = 11, P <0.05). Additionally, PCA results show that Liolaemus janequeoae and Liolaemus cf. elongatus from Llaima occupy a different region of morphological space, without overlap (Fig. 3). Liolaemus janequeoae is smaller (SVL = 65.3 +/- 3.4 mm) than Liolaemus antumalguen (SVL = 95.0 +/- 6.2 mm) (t = -11.3, DF = 14, P <0.01); has a shorter axilla-groin distance (27.8 +/- 2.9 mm vs 43.0 +/- 4.4 mm) ( Mann-Whitney U, P <0.01); a shorter arm length (24.7 +/- 2.3 mm vs 28.4 +/- 0.7 mm) (t = -4.5, DF = 14, P <0.01); a lower head height (6.8 +/- 0.5 mm vs 10.0 +/- 0.6 mm) (t = -11.2, DF = 14, P <0.01); a narrower head (11.0 +/- 0.4 mm vs 16.6 +/- 0.8 mm) (t = -17.2, DF = 14, P <0.01); and has shorter foot length (19.4 +/- 1.4 mm vs 28.5 +/- 1.2 mm) ( Mann-Whitney U, P <0.01); whereas Liolaemus janequeoae has more midbody scales than Liolaemus antumalguen (t = 6.2, DF = 14, P <0.01, Table 1), more dorsal scales (t = 7.6, DF = 14, P <0.01, Table 1) and more ventral scales (t = 8.2, DF = 14, P <0.01, Table 1). Moreover, Liolaemus antumalguen has a very variable dorsal pattern of black spots to almost complete melanism, whereas Liolaemus janequeoae never has black spots (only small black dots in one female). Additionally, PCA results show that both species occupy a different region of morphological space, without overlap (Fig. 3). Liolaemus carlosgarini , Liolaemus scorialis and Liolaemus lonquimayensis have dark lateral and vertebral bands, features that distinguishes these from Liolaemus janequeoae . Additionally, Liolaemus janequeoae is larger than Liolaemus carlosgarini (SVL = 65.3 +/- 3.4 mm vs SVL = 60.2 +/- 5.1 mm) (t = 2.4, DF = 22, P <0.05); Liolaemus janequeoae has a larger axilla-groin length than Liolaemus carlosgarini (27.8 +/- 2.9 mm vs 24.8 +/- 2.9 mm) (t = 2.3, DF = 22, P <0.05); Liolaemus janequeoae has longer arms than Liolaemus carlosgarini (24.7 +/- 2.3 mm vs 21.8 +/- 1.8 mm) (t = 3.4, DF = 22, P <0.01); Liolaemus janequeoae has more dorsal scales than Liolaemus carlosgarini (t = 4.5, DF = 14, P <0.01, Table 1) and more ventral scales (t = 6.8, DF = 14, P <0.01, Table 1); whereas Liolaemus lonquimayensis has larger axilla-groin length (34.9 +/- 1.7 mm) than Liolaemus janequeoae ( Mann-Whitney U, P <0.05); Liolaemus lonquimayensis has a greater head height than Liolaemus janequeoae (8.3 +/- 0.1 mm vs 6.8 +/- 0.5 mm) (t = -4.8, DF = 8, P <0.01); whereas Liolaemus scorialis has the head wider than Liolaemus janequeoae (11.9 +/- 0.6 mm vs 11.0 +/- 0.4 mm) (t = -3.1, DF = 16, P <0.01); Liolaemus janequeoae has more midbody scales than Liolaemus scorialis (t = 3.6, DF = 16, P <0.01, Table 1) and more dorsal scales (t = 4.8, DF = 17, P <0.01, Table 1). Additionally, PCA results show that Liolaemus janequeoae does not overlap in the morphological space with Liolaemus carlosgarini and Liolaemus scorialis when ellipses are generated with the second and third PCs (Fig. 3). Liolaemus janequeoae is smaller (max. SVL = 69.6 mm) than Liolaemus shitan (max. SVL = 98.3 mm) and has more midbody scales (82-98 vs. 72-85). Dorsal color pattern in Liolaemus shitan is black, whereas only one female of our sample of Liolaemus janequeoae has small dorsal black dots. Liolaemus janequeoae is smaller (max. SVL = 69.6 mm) than Liolaemus choique (max. SVL = 90.7 mm). Moreover, Liolaemus choique has a very variable dorsal pattern of black spots to almost complete melanism, whereas Liolaemus janequeoae never has black spots (only small black dots in one female). Liolaemus janequeoae is smaller than Liolaemus crandalli (max. SVL = 69.6 mm vs max. SVL = 93.4 mm). Moreover, Liolaemus crandalli has dark lateral and vertebral bands with ringed tail, whereas all of these features are completely absent in Liolaemus janequeoae . According to Avila et al. (2015) , Liolaemus crandalli is the sister taxon of the pair Liolaemus smaug + Liolaemus choique , whereas in our phylogeny Liolaemus janequeoae is not closely related to Liolaemus smaug or Liolaemus choique . Liolaemus janequeoae is smaller than Liolaemus burmeisteri (max. SVL = 69.6 mm vs max. SVL = 85.2 mm) and has more midbody (82-98 vs. 70-81) and ventral scales (124-132 vs. 99-110). Moreover, Liolaemus burmeisteri has dark lateral bands. Liolaemus janequeoae has more midbody scales than Liolaemus smaug (82-98 vs 73-80). Moreover, Liolaemus smaug has dark lateral and vertebral band. In our phylogeny Liolaemus janequeoae and Liolaemus smaug are not sister taxa. Description of holotype. Adult male. SVL: 59.1 mm. Tail length: 42.0 mm (autotomized). Axilla-groin length: 21.8 mm. Head length: 13.1 mm. Head width (distance between the two ear openings): 10.5 mm. Head height (at the level of ear openings): 6.1 mm. Forelimb length: 21.1 mm. Hindlimb length: 36.0 mm. Foot length: 18.6 mm. Hand length: 9.8 mm. Rostral scale wider (2.36 mm) than high (0.8 mm). Subocular length: 4.2 mm. Fifth supralabial length: 1.6 mm. Neck width: 9.4 mm. Interorbital distance: 4.5 mm. Internasal distance: 1.5 mm. Body width: 13.7 mm. Meatus width: 1.4 mm. Meatus height: 2.1 mm. Two postrostrals. Four internasals. Hexagonal interparietal scale, with a central, small, and whitish '' parietal eye'' in the center. Interparietal smaller than the parietals, surrounded by other nine scales; ten scales between interparietal scale and rostral; seventeen scales between occiput and rostral (Hellmich Index); orbital semicircles are interrupted by one supraocular scales in both sides, but the rest is formed by ten scales on each side; 6-7 supraoculars (left-right); six superciliary scales. Frontal area is divided into three scales (one posterior, one middle and one anterior). Two scales between the nasal and the canthal. Preocular separated from the lorilabials by a single loreal scale. Nasal separated from rostral by one scale, surrounded by seven scales. One row of lorilabials between the supralabials and the subocular; seven supralabials, the fifth is curved upward without contacting the subocular; six infralabial scales. Mental scale is pentagonal, 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. Eleven temporal scales between the level of superciliary scales and the commissure of the mouth. Two projecting scales on the anterior edge of the ear, which do not cover the auditory meatus. Auricular scale is wide and restricted to the upper third of the meatus; 44 gulars between the auditory meatuses. Antehumeral fold and "Y" shaped lateral neck fold. Developed dorsolateral fold. Midbody scales: 94. Dorsal scales are rhomboidal, slightly keeled, without mucrons, subimbricate and with interstitial granules. Dorsal scales are similar in size than ventral ones. Dorsal scales: 89. Ventral scales are rhomboidal, smooth, imbricate, and without interstitial granules. Ventral scales: 124. Three precloacal pores. Hemipenial bulges are evident. The suprafemoral scales are lanceolate, imbricate, and slightly keeled. Infrafemoral scales are lanceolate to rounded, smooth, and imbricate. Scales of the dorsal surface of the forearm are lanceolate to rounded, imbricate, and slightly keeled or smooth. Scales of the ventral surface of the forearm are rounded, smooth, and subimbricate. The dorsal scales of the first third of the tail are rhomboidal to lanceolate, subimbricate or juxtaposed, keeled and with inter stitial granules. The ventral scales of the tail vary from rhomboidal to triangular, and are imbricate and smooth. Lamellae of the fingers: I: 10, II: 14, III: 22, IV: 24 and V: 15. Lamellae of the toes: I: 11, II: 16, III: 22, IV: 32 and V: 19. Coloration in life. Light brown head, with dark brown spots in the parietal area and in the posterior nasal area. The snout is olive. Temporal area is light brown. Subocular area and cheeks are slightly lighter than temporal area. The subocular is immaculate. Background color of the dorsum, limbs, and tail is light brown. The vertebral zone of the dorsum is slightly darker than rest, but without forming an occipital stripe. The only dorsal design is a series of white dots, formed by 1-3 white scales, running from the posterior half of the trunk to the first third of the tail. The tail is immaculate. Ventrally, the throat, belly, limbs and the tail are whitish pearly. Thighs and cloaca have a little yellowish coloration. Precloacal pores are orange. Variation. Despite four field campaigns, no additional males were found. Variation in measures refer to the six female paratypes: SVL: 66.2-69.6 mm. Axilla-groin distance: 27.4-30.2 mm. Head length: 13.5-15.1 mm. Head width: 10.7-11.4 mm. Head height: 6.4-7.6 mm. Foot length: 18.0-21.5 mm. Leg length: 36.5-44.7 mm. Hand length: 9.4-11.7 mm. Arm length: 21.1-26.7 mm. Tail length: 84-110 (n = 3; autotomized in the rest). Relation tail length/SVL = 1.2-1.7. Although more data on males are required, there is no sexual size dimorphism in the Liolaemus elongatus clade species ( Avila et al. 2012 ). Scale number variation in Liolaemus janequeoae (all specimens) is as follows. Midbody scales: 82-98 (91.6 +/- 5.5). Dorsal scales: 77-89 (85.0 +/- 4.2). Ventral scales 124-132 (128.6 +/- 3.5). Fourth finger lamellae: 22-24 (23.5 +/- 0.8). Fourth toe lamellae: 28-32 (29.5 +/- 1.4). Supralabial scales: 6-8 (7.4 +/- 0.8). Infralabial scales: 5-6 (5.3 +/- 0.5). Interparietal scale is pentagonal or hexagonal, bordered by 5-9 scales (6.6 +/- 1.7). The interparietal is smaller than the parietals. The nasal is in contact with the rostral in 28.6% of specimens. Females have a very similar color pattern to the male holotype but without dorsal white dots or yellowish coloration on the thighs and cloaca. One female has four series of black dots (formed by 1-3 black scales) on the dorsum: two on the paravertebral fields (running from the head to the first third of the tail) and two on the dorsolateral area (running from the head to the middle of the trunk). Etymology. This species is named after Janequeo, a prominent Lonko (tribal chief) of Mapuche-Pehuenche origins. She fought against colonial Spaniards in the Arauco war, carried out mainly in the Araucania Region where Liolaemus janequeoae was discovered. It is believed that she became involved in the war after her partner (Lonko Hueputan) was captured and tortured to death. She played a leading role in the Battle of Fort Puchunqui, then retreating to Villarrica, where she disappeared. Distribution and natural history. Only known from the type locality at Laguna Verde ( 38°12'S - 71°44'W ), approximately 13.5 km NW of the summit of the Tolhuaca volcano, Araucania Region, Chile (Fig. 5). Figure 5. Distribution map for Liolaemus janequeoae sp. n. with geographically proximate species of the Liolaemus elongatus clade. In the case of Liolaemus elongatus a sample for each locality was included in the phylogeny. Red star: Liolaemus janequeoae sp. n., Laguna Verde. Yellow triangles: Liolaemus smaug (1= near Las Lenas , 2= between Las Loicas and Peteroa Volcano, 3= near Las Loicas). Blue pentagon: Liolaemus carlosgarini (1= Maule Lagoon, 2= Lircay). Black hexagon: Liolaemus choique (Paso el Choique). Pink diamond: Liolaemus antumalguen (Domuyo Volcano). Brown asterisk: Liolaemus burmeisteri (Caepe Malal). Green cross: Liolaemus crandalli (Auca Mahuida Volcano). Gray squares: Liolaemus scorialis (1= Laja Lagoon, 2= La Mula Lagoon). White circle: Liolaemus lonquimayensis (Lonquimay Volcano). Pink circles: Liolaemus shitan (1= Estancia Piedras Blancas, type locality and 2= near Antonio del Cuy). Blue circle: Liolaemus cf. elongatus (Llaima Volcano). Green circles: Liolaemus elongatus (1= Pampa de Lonco Luan, 2= Primeros Pinos, 3= Portal La Atravesada, 4= Laguna Blanca, 5= near Ingeniero Jacobacci, 6= San Carlos de Bariloche, 7= Ojo de Agua, 8= El Maiten, 9= Esquel, 10= Tecka, 11= Gobernador Costa and 12= Los Manantiales). At Laguna Verde, Liolaemus janequeoae was found between 1336-1397 masl. It inhabits the deciduous highland Andean forest ( Gajardo 1994 ), consisting of Araucaria araucana and Nothofagus dombeyi (1397 masl). The shrubs are represented by Chusquea culeou , Desfontainia spinosa , Drimys andina and Pseudopanax laetevirens . At lower altitudes (1336 masl), the vegetation was dominated by Araucaria araucana and Nothofagus pumilio , with the presence of Azara alpine , Chusquea culeou , Colletia hystrix , Lomatia hirsuta , Maytenus disticha , Myrceugenia chrysocarpa and Pernettya myrtilloides . At lower altitudes where there are no Araucaria araucana , Liolaemus janequeoae was not found. It is a diurnal lizard of apparently low abundance. It was seen on rocks and climbing in trees. Liolaemus janequeoae was found in syntopy with Liolaemus septentrionalis Pincheira-Donoso and Nunez , 2005; Liolaemus tenuis ( Dumeril & Bibron, 1837); Pristidactylus torquatus (Philippi, 1861) and the second new species described below. In this zone, it was also recorded the presence of Tachymenis chilensis (Schlegel, 1837). The intestinal content of one specimen (paratype) was examined and remnants of insects and several nematodes were found. At the date of capture (January 5) two females had two and three embryos each. All other females have only several small oocytes.