Liolaemus balerion, 2020

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

( FIG. 9)

Liolaemus bibronii Lobo & Abdala (2001 View in CoL , 2002), L. bibronii View in CoL 7. Morando et al. (2007), L. sp. 7. In part. Quinteros (2013), L. sp. 9. Portelli & Quinteros (2018), ‘ L. sp. from Ojos de Agua’.

L S I D: u r n: l s i d: z o o b a n k. o r g: a c t: 6 E F 5 E 8 F 6- BB25-4445-A9B5-039EB8C0D976

Holotype: FML 21533 View Materials . Male. 1.5 km N de Ojos de Agua por Provincial Road 6, Pilcaniyeu, Río Negro, Argentina. 41°32’36,3”S; 69°51’36,3”W. 1104 M. COLS C. Abdala, S. Quinteros, J. C. Stazzonelli. GoogleMaps

Paratypes: FML 21532 View Materials , 21457–458 View Materials . Same data of Holotype; 16345–346: 19 km W of Ingeniero Jacobacci, Pilcaniyeu , Río Negro , Argentina; 10075–076: 3.4 km S of Ingeniero Jacobacci, Pilcaniyeu, Río Negro ,

Argentina; MACN 15194–196: Ingeniero Jacobacci, Río Negro, Argentina.

Additional specimens: MCN 490–93; 496: Ingeniero Jacobacci, Pilcaniyeu, Río Negro, Argentina.

Diagnosis: Liolaemus balerion is a small slender Liolaemus (max. SVL 59.83). It belongs to the L. alticolor – bibronii group because it exhibits a set of character states that defines this group (see: Quinteros, 2012, 2013; Quinteros et al., 2014). Populations of Liolaemus balerion were previously assigned to L. bibronii . However, there exist some character state combinations that allow us to distinguish them in two different taxa. Liolaemus balerion differs from L. bibronii in the contact between nasal and rostral, this contact being broad in L. bibronii and narrow in the new species. In L. bibronii , the nasal scale is separated from the canthal by one scale, whereas in L. balerion there are two scales between nasal and canthal. Moreover, there are five scales between frontal and supercilliaries in L. balerion , being four in L. bibronii . In specimens of L. bibronii , the fourth supralabial never contacts the subocular scale, whereas in 60% of specimens of L. balerion , the fourth supralabial scale contacts the subocular scale. Colour pattern between L. bibronii and L. balerion are similar, but some differences exist. Shape of paravertebral spots are line-shaped in L. balerion , being rounded in L. bibronii . On the dorsal head of specimens of L. bibronii , there exists a black line surrounding the interparietal scale; this line is absent in L. balerion . The ventral surface of the tail is immaculate in L. balerion ; which distinguishes the new taxon from L. bibronii , which has black to grey spots on the ventral surface of the tail.

Liolaemus balerion also differs from the other two new species described. Subocular is white (lighter than loreal region) in L. balerion , being of the same colour as the loreal region in L. sp. from Malargüe. Paravertebral spots are line-shaped in L. balerion , whereas in L. sp. from Malargüe they are merged, forming a longitudinal stripe. Dorsolateral stripes in L. sp. from Malargüe are bordered with a black line, whereas in L. balerion , this black line is absent. In L. balerion , 60% of specimens show contact between fourth supralabial and subocular scales, whereas in L. sp. from Malargüe, the fourth supralabial never contacts the subocular. Snout–vent length is larger in L. balerion than in L. sp. from Malargüe. Lengths of second and third toes are larger in L. balerion than in L. sp. from Malargüe, whereas the length of the fourth toe is larger in L. sp. from Malargüe. The head in L. balerion is shorter and wider than in L. sp. from Malargüe. The number of neck and gular scales is higher in L. balerion than in L. sp. from Malargüe. The number of infradigital lamellae on the second finger is higher in L. balerion than in L. sp. from Malargüe. The number of infradigital lamellae on the first, second, third and fourth toes is higher in L. balerion than in L. sp. from Malargüe. The number of temporal scales in L. balerion (average) is lower than in L. sp. from Comallo (average). On the dorsum of the head of specimens of L. sp. from Comallo, there exists a black line surrounding the interparietal scale, which is absent in L. balerion . The lengths of the first, second, third and fourth toes are larger in L. balerion than in L. sp. from Comallo. The number of neck and gular scales is lower in L. balerion than in L. sp. from Comallo. The number of infradigital lamellae on the second finger is higher in L. balerion than in L. sp. from Comallo. The number of infradigital lamellae on the first, second, third and fourth toes is higher in L. balerion than in L. sp. from Comallo.

Liolaemus balerion is geographically isolated (> 1400 km) from L. alticolor , L. aparicioi , L. bitaeniatus , L. chaltin , L. chavin , L. chungara , L. incaicus , L. pachacutec , L. pagaburoi , L. paulinae , L. puna , L. ramirezae , L. tacnae , L. variegatus , L. walkeri , L. wari and L. yanalcu . Lanceolate dorsal scales of L. balerion distinguish it from L. bitaeniatus and L. variegatus (rhomboidal dorsal scales). Dorsal scales of L. balerion are mucronate, while dorsal scales of L. chavin , L. pachacutec , L. paulinae and L. tacnae lack a mucron, and sclaes of L. yalguaraz have a tiny mucron. The number of scales around the midbody in L. balerion (45–57) is higher than in L. abdalai , L. gracilis and L. variegatus (33–45). Moreover, the number of dorsal scales between occiput and thighs is higher in L. balerion (45–53) than in L. abdalai , L. alticolor , L. aparicioi and L. pagaburoi (32–44). The dorsal surface of the head (smooth) distinguishes this new taxon from L. exploratorum , L. pagaburoi (slightly rugose) and from L. bitaeniatus , L. lemniscatus , L. saxatilis , L. tacnae and L. variegatus (markedly rugose). Specimens of L. balerion show smooth temporal scales, differing from L. abdalai , L. chaltin , L. chungara , L. curicensis , L. exploratorum , L. fuscus , L. incaicus , L. pagaburoi , L. pyriphlogos , L. ramirezae , L. tandiliensis , L. yalguaraz and L. yanalcu (weakly keeled), and from L. alticolor , L. aparicioi , L. bitaeniatus , L. cyaneinotatus , L. lemniscatus , L. sanjuanensis , L. saxatilis and L. variegatus (markedly keeled). Also, the neck scales of L. balerion are smooth, distinguishing it from L. abdalai , L. curicensis and L. puna (some scales are keeled), and from L. aparicioi , L. bitaeniatus , L. chaltin , L. cyaneinotatus , L. fuscus , L. incaicus , L. lemniscatus , L. paulinae , L. pagaburoi , L. pyriphlogos , L. ramirezae , L. sanjuanensis , L. saxatilis , L. tandiliensis and L. variegatus (markedly keeled). The number of neck scales of L. balerion (35–41) is bigger than of L. abdalai , L. gracilis , L. lemniscatus , L. saxatilis , L. tandiliensis and L. yalguaraz (18–32). The number of gular scales is also larger in L. balerion (35–41) than in L. alticolor , L. aparicioi , L. bitaeniatus , L. chaltin , L. incaicus , L. pagaburoi , L. tacnae and L. variegatus (20–34). A lack of precloacal pores in females distinguishes the new taxon from L. aparicioi , L. bitaeniatus , L. incaicus , L. ramirezae , L. variegatus and L. yanalcu (females exhibit precloacal pores). Specimens of L. balerion show paravertebral spots, absent in L. alticolor , L. chungara , L. fuscus , L. gracilis , L. chaltin , most of males of L. puna and L. walkeri . The presence of a vertebral line in L. balerion differentiates it from L. abdalai , L. bitaeniatus , L. exploratorum , L. fuscus , L. lemniscatus , L. gracilis , L. saxatilis , females of L. incaicus , L. paulinae and most males of L. puna . Specimens of L. alticolor , L. aparicioi , L. chaltin , L. pyriphlogos , L. puna , L. chavin , L. pachacutec and some specimens of L. wari have a black line surrounding the interparietal scale, while this line is absent in L. balerion . Presence of dorsolateral stripes in L. balerion differentiates this species from L. tacnae and L. yanalcu . Males of L. balerion (50.6–56.7 mm) are larger than males of L. alticolor , L. abdalai and L. tacnae (43–48). Also, females of L. balerion (47.7– 57.2 mm) are larger than females of L. abdalai and L. fuscus (36–45).

Description of the holotype: Adult male. Snout– vent length (SVL) 53.13 mm. Axilla–groin distance 27.03 mm. Head 11.18 mm long (from anterior border of auditory meatus to tip of snout), 9.01 mm wide (at anterior border of auditory meatus), 6.53 mm high. Interorbital distance (between postorbital semicircles) 6.71 mm. Eye–nostril distance 3.79 mm. Tibia length 7.52 mm. Foot length 16.75 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, lower than parietal in size, surrounded by six scales. Frontal scale azygous. Five scales between frontal and rostrals. Supraorbital semicircles complete. Four enlarged supraoculars. Six scales between frontal and supercilliaries. Seven flat, elongated, imbricate supercilliaries. Canthal separated from nasal by two scales. Loreal region flat. Seven scales, including rostral, surrounding nasals. Nasals narrowly contact rostral scale. Six lorilabials, third to fifth in contact with subocular. Six enlarged supralabials. Fourth supralabial curved upward posteriorly contacting subocular. Five infralabials, slightly taller than supralabials. Orbit diameter 3.34 mm (measured between upper and lower ciliaries). Subocular scale elongate, length 3.83 mm. Preocular unfragmented. Longitudinal ridge along upper margin of the three ocular scales. Rostral scale 2.8 times as wide (2.52 mm) as high (0.91 mm). Mental twice as wide (2.68 mm) as high (1.29 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. Three outward projecting laminar scales along anterior border of auditory meatus. Auditory meatus higher (2.8 mm) than wide (1.29 mm). Lateral scales of neck laminar and smooth. Antehumeral fold distinct. Thirty-three scales on the neck (between posterior margin of auditory meatus and shoulder). Twenty-two scales between auditory meatus and antehumeral fold (counted along postauricular and longitudinal folds). Fifty-five dorsal scales between occiput and anterior surface of thighs. Dorsal body scales lanceolate, imbricate, keeled and mucronate. Fifty-six scales around midbody. Ninety ventral scales between mental and the cloaca. Four precloacal pores.

Colour of holotype in alcohol: Dorsal background, light brown to grey. It has a black, complete vertebral line that extends to the tip of the tail. Dorsolateral stripes are absent. The paravertebral region has the same colour as the vertebral region. Two rows of linear paravertebral spots are distinct (nine between fore and hindlimbs). Lateral field, light brown to grey, with dark brown/black irregular spots. Ventrolateral line faded but evident. Head the same colour as the dorsum (light brown), without a black line surrounding the interparietal scale, but speckled with black spots. With dark brown spots on supra and infralabial scales. Temporal region with two white lines: the upper beginning at the posterior margin of the eye, and the lower at the posterior margin of subocular scale. The upper extends over the neck to the shoulder. The lower follows the anterior margin of the auricular meatus and extends to the ventrolateral line over the auricular and longitudinal fold. Fore and hindlimbs are light brown with dark brown spots. Two or three tiny white spots can be observed on the shoulder. Ventrally, the throat is grey with some dark brown lines. Chest and belly are grey, spotless. The region of chinshields has a few black spots. Fore and hindlimbs are immaculate grey, almost white. The tail is dorsally light brown with a black vertebral line and some paravertebral spots, and ventrally light brown, speckled with line-shaped dark brown spots. Lateral region of the tail has dark brown spots (which are a continuation of those of the lateral field of body).

Variation: Based on 16 paratypes (seven males and nine females) and five additional specimens. Snout–vent length 47.70–57.83 mm (mean = 53.80; SD = 4.60) in males; 47.46–59.15 mm (mean = 52.70; SD = 4.50) in females. Head length 10.12–12.28 mm (mean = 11.40; SD = 1.01) in males; 9.76–12.01 mm (mean = 10.90; SD = 0.80) in females. Head width 8.23–9.1 mm (mean = 8.72; SD = 0.40) in males; 7.50–9.79 mm (mean = 8.30; SD = 0.80) in females. Head height 5.54–6.57 mm (mean = 6.10; SD = 0.51) in males; 5.12–6.74 mm (mean = 5.90; SD = 0.59) in females. Axilla–groin distance 21.98–27.90 mm (mean = 28.90; SD = 2.66) in males; 23.23–30.75 mm (mean = 27.70; SD = 3.08) in females. Tail length 62.74– 88.22 mm (mean = 79.17; SD = 8.60). Tibia length 5.84–9.55 mm (mean = 8.30; SD = 1.16). Foot length (from ankle to the tip of fourth claw) 15.170–18.18 mm (mean = 16.90; SD = 1.30) in males; 13.62–16.14 mm (mean = 15.12; SD = 0.80) in females. Midbody scales 45–57 (mean = 512.88; SD = 4.2). Dorsal scales, 46–55 (mean = 50.67; SD = 4.12) between occiput and anterior surface of thighs. Dorsal head scales (Hellmich’s Index) 10–12 (mean = 10.9; SD = 0.6). Ventrals 75–95 (mean = 87.4; SD = 6.75). Scales around interparietal 5–7 (mean = 6; SD = 0.53). Three to four enlarged supraoculars (mean = 3.5; SD = 0.53). Preocular not divided, not fused to subocular. Subocular scale length 2.83–4.49 (mean = 3.67; SD = 0.5). Temporals 7–8 (mean = 7.5; SD = 0.53), smooth. Scales of neck smooth and laminar. Number of scales on neck (between auditory meatus and shoulder, along the horizontal fold) 29–35 (mean = 31.5; SD = 2.51). Scales between auditory meatus and antehumeral fold 19–23 (mean = 21.5; SD = 1.41). Gulars 35–38 (mean = 36.4; SD = 1.14). Six supralabials. Infralabials 4–5 (mean = 4.83; SD = 0.41). Fourth supralabial upturned, in contact with subocular scale in 60% of specimens. Six to eight scales around nasals (mean = 7; DS = 0.43). Five to six scales between rostral and frontal (mean = 5.2; DS = 0.42). Two postrostrals, with 3–7 scale organs each (mean = 4.81; SD = 1.2). Lorilabials 6–7 (mean = 6.25; SD = 0.5). Subdigital lamellae on fourth finger 16–22 (mean = 18.75; SD = 2.55); on fourth toe 22–27 (mean = 23.5; SD = 1.77). Three to four (mean = 3.67; SD = 0.6) precloacal pores in males, absent in females.

Colour in life ( Fig. 9): Dorsal background brown. In some females, vertebral and paravertebral region lighter than in males. With a black, complete (in some specimens segmented) vertebral line that extends to the tip of the tail. Dorsolateral stripes evident, but faint, beginning at the upper margin of the eye and extending to the forelimbs. The paravertebral region has the same colour as the vertebral region. Two rows of black line-shaped paravertebral spots are distinct. In some females, a yellowish colouration is observed in the paravertebral and vertebral regions. The lateral field is the same colour as the dorsum, with dark brown/ black spots. Some females show pink spots. In some specimens, the lateral field has rounded, white, bright spots. These spots are more evident on the shoulder. Ventrolateral line always evident. Head darker than dorsum, speckled with black spots. Temporal region with two white lines: the upper one beginning at the posterior margin of the eye and joining the dorsolateral stripes; and the lower one beginning at the posterior margin of subocular scale and finishing at the anterior margin of the auricular meatus. Between these white lines, the temporal surface is black. The subocular scale is white (lighter than loreal region). Fore and hindlimbs are the same colour as the dorsum, with black spots. Ventrally, throat, chest and belly light grey, without spots. Some specimens show a few dark grey spots in lateral region of throat. Females can show an orange or pale pink colouration on the sides of the belly. Tail dorsally brown with black vertebral line and two rows of paravertebral spots. Lateral region of tail with dark brown spots. Ventrally grey, speckled with dark grey/black spots. Females with proximal region of tail orange-pink. Males with ventral surface of thighs yellow.

Distribution ( Fig. 8): Liolaemus balerion is known from its type locality to the North of Ojos de Agua, and from a close locality: Ingeniero Jacobacci, both from Río Negro Province, Argentina.

Biology: Liolaemus balerion can be found in hills where bushes are the predominant vegetation. It inhabits in sintopy with species of Phymaturus Gravenhorst, 1838 ; in its type locality with P. excelsus, Lobo & Quinteros, 2005 and P. spurcus Barbour, 1921 , and in Ingeniero Jacobacci with P. spectabilis Lobo & Quinteros, 2005 . No more data about its biology are known.

Etymology: In George R. R. Martin’s epic story A song of ice and fire, Balerion, also known as the Black Dread, is one of the mighty dragons who helped king Aegon conquer Westeros.