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Liolaemus balerion QUINTEROS, RUIZ-MONACHESI & ABDALA, 2019

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Higher TaxaLiolaemidae, Iguania, Sauria, Squamata (lizards) 
Subspecies 
Common Names 
SynonymLiolaemus balerion QUINTEROS, RUIZ-MONACHESI & ABDALA 2019
Liolaemus bibronii — LOBO & ABDALA 2001
Liolaemus bibronii — LOBO & ABDALA 2002
L. bibronii 7 — MORANDO et al. 2007
L. sp. 7. — QUINTEROS 2013 (In part)
L. sp. 9. — PORTELLI & QUINTEROS 2018 
DistributionArgentina (Río Negro)

Type locality: 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 elevation.  
Reproduction 
TypesHolotype: FML 21533. Male. COLS C. Abdala, S. Quinteros, J. C. Stazzonelli.
Paratypes: FML 21532, 21457–458. 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. 
DiagnosisDiagnosis: 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). 
Comment 
EtymologyIn 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. 
References
  • Lobo F, Abdala CS. 2002. La información cladística de un set de datos morfológicos en lagartos del género Liolaemus (Iguania: Liolaemidae). Cuadernos de Herpetología 16: 137–150
  • Lobo, Fernando 2001. A Phylogenetic Analysis of Lizards of the Liolaemus chiliensis Group (Iguania: Tropiduridae). The Herpetological Journal 11 (4):137-150
  • Morando, Mariana; Luciano J. Avila, Cameron R. Turner and Jack W. Sites Jr. 2007. Molecular evidence for a species complex in the patagonian lizard Liolaemus bibronii and phylogeography of the closely related Liolaemus gracilis (Squamata: Liolaemini). Molecular Phylogenetics and Evolution 43 (3): 952-973 - get paper here
  • Portelli SN, Quinteros AS. 2018. Phylogeny, time divergence, and historical biogeography of the South American Liolaemus alticolor–bibronii group (Iguania: Liolaemidae). PeerJ 6: e4404
  • Quinteros, A S; M R Ruiz-Monachesi, C S Abdala 2019. Solving the Liolaemus bibronii puzzle, an integrative taxonomy approach: redescription of L. bibronii and description of three new species (Iguania: Liolaemidae). Zoological Journal of the Linnean Society zlz113 - get paper here
  • QUINTEROS, ANDRÉS SEBASTIÁN 2013. A morphology-based phylogeny of the Liolaemus alticolor–bibronii group (Iguania: Liolaemidae). Zootaxa 3670: 1-32 - get paper here
 
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