Liolaemus leftrarui, Troncoso-Palacios, Jaime, Diaz, Hugo A., Puas, German I., Riveros-Riffo, Edvin & Elorza, Alvaro A., 2016

Troncoso-Palacios, Jaime, Diaz, Hugo A., Puas, German I., Riveros-Riffo, Edvin & Elorza, Alvaro A., 2016, Two new Liolaemus lizards from the Andean highlands of Southern Chile (Squamata, Iguania, Liolaemidae), ZooKeys 632, pp. 121-146 : 129-133

Taxon classification Animalia Squamata Liolaemidae

Liolaemus leftrarui sp. n. Figure 6

Holotype.

SSUC Re 646 (Fig. 6a, b). Male collected at Laguna Verde (38°12'S - 71°44'W, 1405 masl), approximately 13.5 km NW of the summit of the Tolhuaca volcano, Araucanía Region, Chile. Collected by J. Troncoso-Palacios, F. Urra and H. Díaz. January 5, 2014.

Paratypes.

SSUC Re 647-48, 716 (Fig. 6). Three females. Same data as the holotype. SSUC Re 732-734. Two males and one female. Near Lagunillas, Araucanía Region, Chile (38°12'S - 71°46'W, 1483 masl), approximately 4 km NW from the type locality. Collected by J. Troncoso-Palacios & E. Villarroel. September, 2016.

Diagnosis.

Liolaemus leftrarui is closely related to Liolaemus villaricensis . This species is characterized by 1) lack of precloacal pores in either sex, 2) large size Liolaemus (max. SVL = 81.8 mm), 3) high amount of midbody scales (80-88), 4) light blue dots on the dorsum, and 5) absence of ventral melanism. We provide a diagnosis in regards to Liolaemus villaricensis , plus four unrelated species that occur geographically near to Liolaemus leftrarui and that also feature the absence of precloacal pores. Based on seven specimens.

Liolaemus leftrarui has more dorsal scales than Liolaemus villaricensis (77-87 vs. 80-89) (t = -2.5, DF = 11, P <0.05). Moreover, Liolaemus villaricensis has a marked lateral black band and a fragmented vertebral stripe, whereas in Liolaemus leftrarui these two color features are inconspicuous or less marked than in Liolaemus villaricensis . Liolaemus villaricensis has no light blue dots, which are in all specimens of Liolaemus leftrarui . Finally, although they are sister species, the average uncorrected pairwise distance between the two taxa is 7.3%, more than double that value proposed for identification of candidate species in Liolaemus . Additionally, PCA results show that both species only marginally overlap in morphological space when ellipses are generated with the two first PCs (Fig. 3).

Liolaemus leftrarui is larger (max. SVL = 81.8 mm) than Liolaemus coeruleus (males SVL = 58.7 ± 3.2 mm; females SVL = 58.2 ± 2.8 mm) and Liolaemus neuquensis (males SVL = 57.4 ± 3.5 mm; females SVL = 58.2 ± 1.9 mm). Moreover, Liolaemus coeruleus males feature black ventral color and some Liolaemus neuquensis males also feature a black ventral color, a feature absent in Liolaemus leftrarui . Females of Liolaemus coeruleus and Liolaemus neuquensis have a brown dorsal color, but females of Liolaemus leftrarui have a bluish brown dorsal color. Finally, in our phylogeny Liolaemus neuquensis is not closely related to Liolaemus leftrarui and although we have no molecular data for Liolaemus coeruleus , this last species and Liolaemus neuquensis are probably conspecific ( Avila et al. 2003).

Liolaemus leftrarui has more midbody scales (80-88 vs. 67-81) than Liolaemus punmahuida . Dorsal color in Liolaemus punmahuida is ochre and this species is patternless, whereas Liolaemus leftrarui has brown dorsal color with dispersed light blue dots. Liolaemus punmahuida has reddish color around the cloaca, feature absent in Liolaemus leftrarui . The species are not closely related according to our phylogeny.

Liolaemus leftrarui differs from Liolaemus tregenzai in that this last species features black color on the throat, chest and abdomen of males and gray color on the throat, chest and abdomen of females, features totally absent in Liolaemus leftrarui . The species are not closely related according to our phylogeny.

Description of holotype.

Adult male. SVL: 81.7 mm. Tail length: 102.9 mm (not autotomized). Axilla-groin length: 35.4 mm. Head length: 20.1 mm. Head width (distance between the two ear openings): 16.9 mm. Head height (at the level of ear openings): 10.8 mm. Forelimb length: 26.5 mm. Hindlimb length: 46.0 mm. Foot length: 21.8 mm. Hand length: 13.6 mm. Rostral scale wider (4.3 mm) than high (1.6 mm). Subocular length: 5.7 mm. Fourth supralabial length: 3.4 mm. Neck width: 16.2 mm. Interorbital distance: 7.2 mm. Internasal distance: 3.0 mm. Body width: 27.2 mm. Meatus width: 1.0 mm. Meatus height: 3.3 mm.

Two postrostrals. Four internasals. Pentagonal interparietal scale, with a central, small, and whitish ‘‘ parietal eye’’ in the center. Interparietal scale is similar in size to parietal one, surrounded by other six scales; seven scales between interparietal scale and rostral; twelve scales between occiput and rostral; orbital semicircle is incomplete in the right side and complete in the left side (formed by 12 scales); 5-4 supraoculars (left-right); five superciliary scales. Frontal area is divided into three scales (two posterior and one anterior). Remarkably, only one scale between the nasal and the canthal. Preocular separated from the lorilabials by a single loreal scale. Nasal in contact with the rostral, surrounded by seven scales. One row of lorilabials between the supralabials and the subocular; six supralabials, the fourth is curved upward without contacting the subocular; five infralabial scales. Mental scale is pentagonal, in contact with four scales; five pairs of postmental shields, the second is separated by two scales. Temporal scales are subimbricate and smooth, very few are slightly keeled. Eight temporal scales between the level of superciliary scales and the level of the commissure of the mouth. Two enlarged 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; 42 gulars between the auditory meatuses. Antehumeral fold and “Y” shaped lateral neck fold. Present inconspicuous ventrolateral fold. Midbody scales 86. Dorsal scales are rounded to lanceolate, slightly keeled, without mucrons, imbricate and with some interstitial granules. Dorsal scales are smaller than ventral ones. Dorsal scales 81. Ventral scales are rhomboidal to rounded, smooth, imbricate, and without interstitial granules. Ventral scales 118. There are no precloacal pores. Hemipenial bulges are evident. The suprafemoral scales are lanceolate, imbricate, and smooth or slightly keeled. Infrafemoral scales are rounded, smooth, and imbricate. Scales of the dorsal surface of the forearm are rounded, imbricate, and slightly keeled or smooth. Scales of the ventral surface of the forearm are rounded, smooth, juxtaposed or subimbricate with interstitial granules. The dorsal scales of the tail are rhomboidal, imbricate, keeled and some with mucrons. The ventral scales of the tail vary from rhomboidal to triangular, and are imbricate and smooth. Lamellae of the fingers: I: 12, II: 14, III: 20, IV: 22 and V: 15. Lamellae of the toes: I: 11, II: 16, III: 21, IV: 27 and V: 18.

Coloration in life.

Brown head, with dispersed dark brown spots. Occipital area of the head is dark brown; temporal area is brown with three dark brown stripes and some dispersed light blue scales. Ocular area, snout and cheeks are light green. Subocular scale is light blue with two dark brown vertical lines, one in the middle and other in the anterior edge. Background color of the dorsum is brown. Inconspicuous dorsolateral light brown stripe (two scales of wide) running from the occiput level to the level of the axilla. Dark brown spots dispersed on the dorsum, without forming an occipital band, but forming three lines on the neck; one of which (middle) forms an inconspicuous vertebral stripe on the dorsum. Several light blue dots dispersed on the dorsum (each corresponds to one scale). Inconspicuous dark brown lateral band with dispersed light blue scales. Below lateral band, flanks are yellowish. Limbs are brown with light green and few dispersed light blue scales. Tail is brown with dispersed light green scales and dark brown vertebral line. Ventrally, the throat is dark green, darker towards the tip of the snout. Belly and the tail are light green. Rear portion of belly, cloaca, chest and thighs have a yellowish coloration. Palms are dark brown and soles are light brown.

Variation.

Variation in three males (including the holotype): SVL: 76.1-81.8 mm. Axilla-groin distance: 33.2-35.7 mm. Head length: 17.9-20.1 mm. Head width: 14.6-16.9 mm. Head height: 9.3-10.8 mm. Foot length: 20.2-21.8 mm. Leg length: 42.7-46.0 mm. Hand length: 12.0-13.6 mm. Arm length: 26.0-27.3 mm. Tail autotomized in all male paratypes. Variation in four female paratypes is as follows: SVL: 60.5-68.2 mm. Axilla-groin distance: 26.4-30.1 mm. Head length: 13.2-15.0 mm. Head width: 9.7-12.0 mm. Head height: 6.3-7.0 mm. Foot length: 17.4-17.9 mm. Leg length: 32.5-38.2 mm. Hand length: 10.1-11.1 mm. Arm length: 20.5-21.2 mm. Tail autotomized in all females.

Scale number variation in Liolaemus leftrarui (all specimens) is as follows. Midbody scales: 80-88 (84.3 ± 3.5). Dorsal scales: 77-87 (81.3 ± 3.6). Ventral scales 108-123 (115.3 ± 5.8). Fourth finger lamellae: 20-23 (21.9 ± 1.1). Fourth toe lamellae: 27-30 (28.1 ± 1.3). Supralabial scales: 6-7 (6.4 ± 0.5). Infralabial scales: 4-5 (4.7 ± 0.5). Holotype has only one scale between the nasal and the canthal, but paratypes have two, as usual in the genus Liolaemus . No precloacal pores in the males and no vestigial precloacal pores in the females, which is rare in Liolaemus . Interparietal scale is quadrangular, pentagonal, hexagonal or heptagonal, bordered by 5-7 scales (5.7 ± 0.8). The interparietal is similar size or smaller than the parietals. The canthal is in contact with the rostral in all specimens.

Paratype males have similar coloration pattern to the holotype with variation only in shade. Females have similar coloration pattern to the holotype, but with some differences such as: the dark brown color on the occipital area is less marked or absent; the dark brown lateral band (inconspicuous in the holotype) is marked in some females; the dark brown vertebral stripe of the tail is inconspicuous or absent in females; the ventral color is light green or light blue; the throat is reticulated in one female; the yellowish color on the rear portion of belly and the cloaca is less marked or absent in females.

Etymology.

This species is named after Leftraru, the most prominent Lonko (tribal chief) of the Mapuche people, who fought against colonial Spaniards in the Arauco war, carried out mainly in the Araucanía Region where we discovered Liolaemus leftrarui . He was captured when he was eleven by Pedro de Valdivia (Governor of the Kingdom of Chile) and became his personal servant. He learned the military strategy of the Spanish and then escaped. Later, he ambushed and killed Valdivia, and won the most remarkable victories over the Spaniards. Finally, he was surrounded and died in battle.

Distribution and natural history.

Known from two localities: 1) the type locality at Laguna Verde (38°12'S - 71°44'W), approximately 13.5 km NW of the summit of the Tolhuaca volcano, Araucanía Region, Chile (Fig. 7). At Laguna Verde, Liolaemus leftrarui was found between 1336-1397 masl. Vegetation is the same described for the habitat of Liolaemus janequeoae . At lower altitudes where there are no Araucaria araucana , Liolaemus leftrarui was not found. 2) Near Lagunillas (38°12'S - 71°46'W, 1483 masl), approximately 4 km NW from Laguna Verde, in the Araucaria araucana forest. It is probable that the distribution of Liolaemus leftrarui could extend to Lagunillas (1700 masl) but in September (date of collection) this area is covered with snow and no specimens were found. Remarkably, Liolaemus janequeoae was not found near Lagunillas. Liolaemus leftrarui is a diurnal lizard of apparently low abundance at both localities. It was seen on rocks and trees (in Laguna Verde), clambering to approximately 5 m aboveground in trees when threatened. Near Lagunillas it was seen only in fallen trees.

Liolaemus leftrarui was found in syntopy with Liolaemus septentrionalis , Liolaemus tenuis , Liolaemus janequeoae and Pristidactylus torquatus at the type locality. Near Lagunillas it was found in syntopy with Liolaemus septentrionalis and Liolaemus tenuis . In this zone the presence of Tachymenis chilensis was also recorded.

The intestinal contents of one specimen from the type locality was examined and revealed the remnants of insects. No plant remains were found. One specimen from near Lagunillas had several nematodes in the intestines. The females collected in January had several small oocytes but the female collected in September carried one embryo.