Liolaemus meraxes, 2020

LIOLAEMUS MERAXES QUINTEROS ET AL. View in CoL , SP. NOV.

( FIG. 10)

Liolaemus bibronii Schulte et al. (2000) View in CoL , L. bironii 8 Morando et al. (2007), L. sp. 10 Portelli & Quinteros (2018), ‘ L. sp. from Malargüe’.

L S I D: u r n: l s i d: z o o b a n k. o r g: a c t: E 5 F F B 1 1 B - 9120-4CB0-A439-03B24249184B

Holotype: FML 7207 View Materials . Male. Bardas Blancas. Malargüe, Mendoza Province. 35°45’04,0”S; 69°34’49,0”W. GoogleMaps

Paratypes: FML 7202–06 View Materials . 7208–13. Eight females and two males. Same data as the Holotype GoogleMaps .

Diagnosis: Liolaemus meraxes is a small, slender Liolaemus (Max SVL 53.14mm) that belongs to the L. alticolor – bibronii group, because it shares a set of character states as defined by Quinteros (2012, 2013) and Quinteros et al. (2014). Liolaemus meraxes was previously identified as a population of L. bibronii , but it has a combination of characters that allow us distinguish it from L. bibronii and from all the other members of the L. alticolor – bibronii group.

The number of scales between canthal and nasal is two in L. meraxes , being one in L. bibronii . The number of scales between frontal and rostral in L. meraxes is 6–7, being lower in L. bibronii (5). Number of supercilliaries is higher in L. meraxes (average) than in L. bibronii (average) . Liolaemus meraxes has enlarged scales on the anterior margin of the auditory meatus; these scales are absent in L. bibronii . The number of scales of circumorbital semicircles is higher in L. meraxes (average) than in L. bibronii (average) . Subocular scale in L. bibronii is white (lighter than loreal region), being of the same colour as the loreal region in L. meraxes . Paravertebral spots are rounded in L. bibronii , whereas in L. meraxes these spots are merged forming a longitudinal stripe. In L. bibronii there is a black line surrounding the interparietal that projects forward to the frontonasals; this line is absent in L. meraxes .

Liolaemus meraxes occurs geographically closely to L. yalguaraz , but L. yalguaraz has the upper temporal scales weakly keeled, which are smooth in L. meraxes . The number of temporal scales is higher in L. yalguaraz (8–9) than in L. meraxes (7). The dorsal scales of L. yalguaraz exhibit a tiny mucron, whereas in L. meraxes the dorsal scales have a large mucron. The subocular scale is of the same colour as the loreal region in L. meraxes , being white (lighter than loreal region) in L. yalguaraz . Specimens of L. yalguaraz (SVL mean = 55.8) are larger than specimens of L. meraxes (SVL mean = 47.8). Also, the colour pattern of L. meraxes distinguishes it from L. yalguaraz . The dorsal colour is golden-reddish in Liolaemus yalguaraz and brown-greyish in L. meraxes . Paravertebral spots in L. meraxes are merged forming an evident continuous line, whereas in L. yalguaraz , paravertebral spots are rectangular and faded in some specimen. The vertebral field is narrower in L. meraxes (2.9 mm) than in L. yalguaraz (4.6 mm).

Liolaemus meraxes can be distinguished from L. balerion and L. sp. from Comallo by the following characters: the subocular is white (lighter than loreal region) in L. balerion , being of the same same colour as the loreal region in L. meraxes . The paravertebral spots are linear in L. balerion , whereas in L. meraxes they are merged, forming a longitudinal stripe. Dorsolateral stripes in L. meraxes are bordered with a black line, whereas in L. balerion , this black line is absent. In L. balerion , 60% of specimens show contact between the fourth supralabial and subocular scales, whereas in L. meraxes , the fourth supralabial never contacts the subocular. Snout–vent length is larger in L. balerion than in L. meraxes . The lengths of the second and third toes are larger in L. balerion than in L. meraxes , whereas the length of the fourth toe is larger in L. sp. from Malargüe. The head is shorter in L. bibronii than in L. meraxes . The head of L. balerion is wider than the head of L. meraxes . The number of neck and gular scales is higher in L. balerion than in L. meraxes . The number of infradigital lamellae on second finger is higher in L. balerion than in L. meraxes . The number of infradigital lamellae on first, second, third and fourth toes is higher in L. balerion than in L. meraxes . The number of temporal scales in L. meraxes (average) is lower than in L. meraxes (average). The subocular scale is white (lighter than loreal region) in L. sp from Comallo, being of the same colour as the loreal region in L. meraxes . Paravertebral spots in L. meraxes are merged forming a longitudinal stripe, whereas in L. sp from Comallo, these spots are irregularly shaped and never merged. Dorsolateral stripes in L. meraxes are bordered by a black line, in L. sp. from Comallo dorsolateral stripes lack the black border. In the dorsum of the head of specimens of L. sp. from Comallo, there is a black line surrounding the interparietal scale, but this line is absent in L. meraxes . Snout–vent length is larger in L. sp. from Comallo than in L. meraxes . The fifth finger is longer in L. meraxes than in L. sp. from Comallo. The head is longer in L. meraxes than in L. sp. from Comallo, but wider in L. sp. from Comallo than in L. meraxes . The tibia of L. meraxes is longer than in L. sp. from Comallo. The numbers of neck and gular scales are smaller in L. meraxes than in L. sp. from Comallo. The number of infradigital lamellae on the second finger is larger in L. sp. from Comallo than in L. meraxes .

The presence of a vertebral line distinguishes L. meraxes from L. abdalai , L. bitaeniatus , L. exploratorum , L. fuscus , L. incaicus , L. lemniscatus , L. sanjuanensis and L. saxatilis . Males of L. chavin , L. pachacutec , L. wari and L. walkeri have a partial or total ventral melanism, absent in L. meraxes . Dorsal scales of L. meraxes have a mucron, which is absent in L. alticolor , L. chavin , L. pachacutec , L. paulinae , L. tacnae and L. tandiliensis . Temporal scales of L. meraxes are smooth, which distinguishes it from L. abdalai , L. alticolor , L. chaltin , L. chungara , L. curicensis , L. exploratorum , L. fuscus , L. incaicus , L. pagaburoi , L. pyriphlogos , L. ramirezae , L. tandiliensis L. variegatus and L. yanalcu (weakly keeled), and from L. aparicioi , L. bitaeniatus , L. cyaneinotatus , L. lemniscatus , L. sanjuanensis and L. saxatilis (markedly keeled). Neck scales of L. meraxes are laminar and smooth, differing from L. alticolor and L. yanalcu (laminar and weakly keeled), from L. abdalai , L. aparicioi , L. bitaeniatus , L. chaltin , L. curicensis L. incaicus , L. pagaburoi , L. paulinae , L. puna , L. pyriphlogos , L. ramirezae and L. variegatus (some scales keeled), L. cyaneinotatus , L. fuscus , L. lemniscatus , L. sanjuanensis , L. saxatilis and L. tandiliensis (keeled neck scales) and from L. chungara (both granular and laminar scales). The number of neck scales of L. meraxes (26–32) is lower than in L. exploratorum (34–41). The surface of the head is smooth in L. meraxes , whereas it is slightly rugose in L. exploratorum and rugose in L. bitaeniatus , L. lemniscatus and L. saxatilis . Liolaemus meraxes shows paravertebral spots, but they are absent in L. fuscus and L. gracilis .

Description of the holotype: Adult male. Snout– vent length (SVL) 54.21 mm; axilla–groin distance 26.39 mm. Head 12.35 mm long (from anterior border of auditory meatus to tip of snout), 9.56 mm wide (at anterior border of auditory meatus), 7.08 mm high. Interorbital distance (between postorbital semicircles) 6.73 mm. Eye–nostril distance 4.35 mm. Tibia length 9.06 mm. Foot length 16.12 mm (from ankle to tip of claw on fourth toe). Dorsal head scales smooth between rostral and anterior border of auditory meatus. Seven smooth temporals. Interparietal subpentagonal, larger than parietal in size, surrounded by six scales. Frontal azygous. Six scales between frontal and rostral. Supraorbital semicircles incomplete. Four enlarged supraoculars. Six scales between frontal and supercilliaries. Six flat, elongate, imbricate supercilliaries. Canthal separated from nasal by two scales. Loreal region flat. Seven scales, including rostral, surrounding nasals. Nasals contact rostral scale. Seven lorilabials, third to fifth in contact with subocular. Six enlarged supralabials. Fourth supralabial curved upward posteriorly, without contacting subocular. Four infralabials, slightly taller than supralabials. Orbit diameter 4.66 mm (measured between upper and lower ciliaries). Subocular scale elongate, length 3.94 mm. Preocular unfragmented. Longitudinal ridge along upper margin of the three ocular scales. Rostral scale 2.4 times as wide (3.02 mm) as high (1.23 mm). Mental 1.6 times as wide (2.62 mm) as high (1.59 mm), followed posteriorly by two rows of four chinshields. Two scales in contact with second infralabial. Scales of throat between chinshields subimbricate. Thirty-five gulars between auditory meatus. Four outward projecting laminar scales along anterior border of auditory meatus. Auditory meatus higher (2.21 mm) than wide (1.02 mm). Lateral scales of neck laminar and smooth. Antehumeral fold distinct. Thirty scales in neck (between posterior margin o auditory meatus and shoulder). Twenty scales between auditory meatus and antehumeral fold (counted along postauricular and longitudinal folds). Forty-seven dorsal scales between occiput and anterior surface of thighs. Dorsal body scales lanceolate, imbricate, keeled and mucronate. Fifty-three scales around midbody. Eighty-two ventrals between mental and the vent. Three precloacal pores.

Colour of holotype in alcohol: Dorsal background, light brown to grey, with a black, complete vertebral line that extends to the tip of the tail. Dorsolateral stripes present but faded. The paravertebral region presents the same colour as the vertebral region. Paravertebral spots are merged, forming a narrow, longitudinal line. Lateral field, light brown to grey, with faded, dark, black irregular spots. Ventrolateral line faded but evident. Head the same colour as the dorsum, with dark brown spots. Some black spots are evident on supra and infralabial scales. Subocular scale of the same colous as the loreal region. A light-grey to white line begins at the upper margin of the auditory meatus and extends to the shoulder along the auricular and longitudinal fold. Fore and hindlimbs are light grey with some dark brown spots. Ventrally, the throat and anterior region of the chest grey with some dark brown lines. Posterior region of chest and belly grey, spotless. Fore and hindlimbs are immaculate grey. The tail is dorsally light brown with a black vertebral line. Ventrally, light brown, speckled with linear dark brown spots. Lateral region of tail with dark brown spots.

Variation: Based on 11 paratypes (eight males and three females). Snout–vent length 45.10–53.14 mm (mean = 47.80; SD = 3.60) in males; 37.4–48.6 mm (mean = 4.3; SD = 4.6) in females. Head length 13.18–15.53 mm (mean = 14.10; SD = 1.20) in males; 9.8–14.7 mm (mean = 12.5; SD = 2.1) in females. Head width 7.58–8.60 mm (mean = 8.20; SD = 0.50) in males; 6.60–8.86 mm (mean = 7.50; SD = 0.70) in females. Head height 6.02–6.76 mm (mean = 6.50; SD = 0.39) in males; 4.74–6.66 mm (mean = 5.80; SD = 0.60) in females. Axilla–groin distance 18.80–23.12 mm (mean = 21.40; SD = 2.27) in males; 16.7–22.5 mm (mean = 20.2; SD = 1.8) in females. Tail length 70.05– 92.98 mm (mean = 77.01; SD = 7.17). Tibia length 7.54–9.57 mm (mean = 8.64; SD = 0.69). Foot length (from ankle to the tip of fourth claw) 15.17–21.22 mm (mean = 17.80; SD = 3.46) in males; 12.53–16.23 mm (mean = 13.08; SD = 1.04) in females. Midbody scales 45–53 (mean = 49.31; SD = 2.14). Dorsal scales 44–55 (mean = 51; SD = 2.58) between occiput and anterior surface of thighs. Dorsal head scales (Hellmich’s Index) 10–15 (mean = 12.54; SD = 1.27). Ventrals 70–86 (mean = 81.08; SD = 4.23). Scales around interparietal 6–7 (mean = 6.31; SD = 0.48). Three to four enlarged supraoculars (mean = 3.54; SD = 0.52). Preocular not divided, not fused to subocular. Subocular scale length 2.32–2.76 (mean = 2.52; SD = 0.13). Seven smooth temporals. Scales of neck smooth and laminar. Number of scales on neck (between auditory meatus and shoulder, along the horizontal fold) 26–34 (mean = 30.46; SD = 2.3). Scales between auditory meatus and antehumeral fold, 18–24 (mean = 21.23; SD = 1.83). Gulars 31–37 (mean = 33.46; SD = 1.94). Five to seven supralabials (mean = 6; SD = 0.41). Infralabials 5–6 (mean = 5.08; SD = 0.28). Fourth supralabial upturned, in contact with subocular scale in 8% of specimens. Seven scales around nasals. Six to seven scales between rostral and frontal (mean = 6.38; DS = 0.51). Two postrostrals, with 3–7 scale organs each (mean = 4.81; SD = 1.2). Six to eight lorilabials (mean = 6.62; SD = 0.77). Subdigital lamellae on fourth finger 17–21 (mean = 18.46; SD = 0.97); on fourth toe 22–27 (mean = 24.08; SD = 1.38). Two to three (mean = 2.67; SD = 0.58) precloacal pores in males, absent in females.

Colour in life ( Fig. 10): Dorsal background, greyishbrown. With a black, complete (fragmented in some specimens) vertebral line that extends to the tip of the tail. Dorsolateral stripes evident, bordered by a black line. The paravertebral region darker than the vertebral region. The paravertebral spots are merged forming a longitudinal stripe. Lateral field, light brown, with black, rounded spots. Ventrolateral line is evident. Head of the same colour as the dorsum, without a black line surrounding the interparietal scale, but speckled with black spots. These are more evident on supraocular scales. With black spots on supra and infralabial scales. Temporal region with a white to yellow line, which begins in the posterior margin of the eye and joins dorsolateral stripes. This line extends over the neck to the shoulder. A white to yellow line begins on upper margin of the auditory meatus, and extends to the ventrolateral line over the auricular and longitudinal folds. Fore and hindlimbs light brown with dark brown spots. Ventrally, throat presents a cream colour with some dark brown spots. These spots extend to the neck on some specimens. Chest and belly cream colour, without spots. Fore and hindlimbs, immaculate, almost white. Tail, dorsally brown with black vertebral line (sometime fragmented). Ventrally, light brown, speckled with line-shaped, dark brown spots. Lateral region of tail with dark brown spots. Males with ventral surface of thighs yellow. Most females show the same colour pattern described previously. One female exhibits a light pattern, vertebral line evident. Dorsolateral stripes are light grey and narrower than in the other specimens. Paravertebral region present the same colour as the vertebral region. Paravertebral spots are merged forming a longitudinal line, but narrower than the other specimens. Lateral field, light brown with a few faded spots. Ventrolateral line barely distinguishable. Ventrally, the throat, chest and belly are light grey. The throat has the same spots as the other specimens. Lateral regions of the belly and cloacal regions yellow.

Distribution ( Fig. 8): Liolaemus meraxes is known from its type locality at Bardas Blancas and nearby el Manzano, in Malargüe, South of Mendoza Province, Argentina.

Biology: Liolaemus meraxes inhabits places where bushes and pastureland are predominant. This species is associated with rocky places, where Festuca spp. grasses occur. In Bardas Blancas, the new species is sympatric with L. grosseorum Etheridge, 2001 , L. josei Abdala, 2005 and L. smaug Abdala et al., 2010 . In Manzano, it is found together with L. choique Abdala et al., 2010 .

Etymology: In George R. R. Martin’s epic A song of ice and fire, Meraxes, is one of the three mighty dragons who helped to conquer Westeros. It was mounted by Rhaenys.