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Liolaemus smaug ABDALA, QUINTEROS, SCROCCHI & STAZZONELLI, 2010

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Higher TaxaLiolaemidae, Iguania, Sauria, Squamata (lizards)
Subspecies 
Common Names 
SynonymLiolaemus smaug ABDALA, QUINTEROS, SCROCCHI & STAZZONELLI 2010
Liolaemus elongatus elongatus CEI 1974
Liolaemus elongatus — CEI 1975
Liolaemus elongatus — CEI 1998
Liolaemus choique ABDALA, QUINTEROS, SCROCCHI & STAZZONELLI 2010
Liolaemus choique — VRDOLJAK et al.2020 
DistributionArgentina (Mendoza)

Type locality: between Las Loicas and Volcán Peteroa Provincial Road 186 (35°39'51.3''S 70°12'0.9''W), 1688 m elevation, Malargüe Department, Mendoza, Argentina.

choique: Argentina (Mendoza); Type locality: Paso el Choique, on provincial road 221 (36°22'01.1''S, 69°48'07.2''W), 2407 m elevation, Malargüe Department, Mendoza Province, Argentina.  
Reproductionviviparous (not imputed, fide Zimin et al. 2022) 
TypesHolotype: FML 22449
Holotype: FML 22453 [choique] 
DiagnosisDiagnosis (smaug): A slender, medium size Liolaemus (Max SVL 71.25 mm), which belong to the L. chiliensis group, particularly in the L. elongatus group (sensu Lobo et al. 2010). Inside this group the new species is distributed far away south from the members of the L. capillitas group (Lobo, 2005) who inhabit in Northwestern Argentina and differs of them because the lack of red coloration on cloacal region and tiny spots spread in shoulder region (both character sates are synapomorphies of the L. capillitas group, sensu Lobo, 2005). Liolaemus smaug differs from all members of the L. petrophilus group in having a distinct color pattern and in character states of lepidosis. It differs from the southern members of the L. petrophilus group (L. austromendocinus, L. elongatus, L. gununakuna (Avila, Morando, Perez, and Sites, 2004), L. petrophilus, and L. thermarum) because it has a small SVL (max SVL 71.25 mm in L. smaug vs. 81 – 103 mm). Dorsal scales have a distinct keel in L. smaug, while in L. austromendocinus the dorsal scales are weakly keeled. Temporal scales are keeled in L. elongatus, L. petrophilus, and L. thermarum while in L. smaug the temporal scales are smooth. Liolaemus smaug have lower number of scales around midbody than L. gununakuna and L. thermarum (73- 80 in L. smaug vs. 84-97). The number of dorsal scales in L. smaug is lower than in L. gununakuna but higher than in L. elongatus (62-68 in L. elongatus; 69-83 in L. smaug; 83-90 in L. gununakuna). Liolaemus gununakuna has a lower number of ventral scales than L. smaug (108-112 in L. gununakuna vs. 119-131 in L. smaug). Precloacal pores are present in males of Liolaemus smaug, character state that differs with males of L. thermarum where the precloacal pores are absent. Sexual dichromatism is evident in L. smaug, being absent in L. austromendocinus, L. elongatus, L. gununakuna, L. parvus, L. petrophilus, and L. thermarum. It differs from the member of the L. kriegi group (sensu Morando et al., 2003) in a lower number of scales around midbody (73-80 vs 85-110 in L. burgeri and L. kriegi); lower number of dorsal scales (71-83 vs 87-110 in L. kriegi). Liolaemus burgeri exhibit a red coloration in cloacal region, absent in L. smaug. The max SVL in L. smaug is 71.3 mm, being 75-108 mm in L. burgeri and in 73-98 mm L. kriegi.

Diagnosis (choique): A large sized Liolaemus (Max SVL 90.7 mm), which belongs to the L. chiliensis group. Inside this group, it belongs to the L. elongatus group (sensu Lobo et al., 2010). The new species is distributed far away south from the members of the L. capillitas group (Lobo, 2005) – included in the L. petrophilus group – which inhabit in Northwestern Argentina and differs by the absence of the synapomorphies of the L. capillitas group (see Lobo, 2005). The new species differs from L. austromendocinus, L. elongatus, L. gununakuna, L. parvus, L. petrophilus, L. shitan, L. thermarum, and L. smaug, in the different dorsal color pattern. Liolaemus choique is smaller (Max SVL 90.7 mm) than L. austromendocinus, L. petrophilus and L. shitan (98 -103 mm), but larger than L. parvus and L. smaug (71-77 mm). The temporal scales are slightly keeled in L. choique, these keels are absent in the temporal scales of L. austromendocinus and L. smaug. In L. choique the dorsal scales exhibit a conspicuous keel, in L. austromendocinus and L. thermarum the dorsal scales are weakly keeled. The number of ventral scales (118-135, mean = 124) in L. choique is larger than in L. elongatus, L. gununakuna, and L. parvus, (96-105, 108-112 and 96- 113 respectively). The number of dorsal scales in L. choique (65-81) is lower than in L. gununakuna (83-90). The males of L. choique exhibit precloacal pores, absent in L. thermarum. Liolaemus smaug exhibit sexual dichromatism, absent in L. choique. The presence of ringed pattern in the tail in L. austromendocinus, L. gununakuna, and L. petrophilus is not shared by L. choique (tail without ringed pattern). It differs from the member of the L. kriegi group (Morando et al., 2003) in the lower number of dorsal scales (65-81 vs 87-110 Diagnosis: A large sized Liolaemus (Max SVL 90.7 mm), which belongs to the L. chiliensis group. Inside this group, it belongs to the L. elongatus group (sensu Lobo et al., 2010). The new species is distributed far away south from the members of the L. capillitas group (Lobo, 2005) – included in the L. petrophilus group – which inhabit in Northwestern Argentina and differs by the absence of the synapomorphies of the L. capillitas group (see Lobo, 2005). The new species differs from L. austromendocinus, L. elongatus, L. gununakuna, L. parvus, L. petrophilus, L. shitan, L. thermarum, and L. smaug, in the different dorsal color pattern. Liolaemus choique is smaller (Max SVL 90.7 mm) than L. austromendocinus, L. petrophilus and L. shitan (98 -103 mm), but larger than L. parvus and L. smaug (71-77 mm). The temporal scales are slightly keeled in L. choique, these keels are absent in the temporal scales of L. austromendocinus and L. smaug. In L. choique the dorsal scales exhibit a conspicuous keel, in L. austromendocinus and L. thermarum the dorsal scales are weakly keeled. The number of ventral scales (118-135, mean = 124) in L. choique is larger than in L. elongatus, L. gununakuna, and L. parvus, (96-105, 108-112 and 96- 113 respectively). The number of dorsal scales in L. choique (65-81) is lower than in L. gununakuna (83-90). The males of L. choique exhibit precloacal pores, absent in L. thermarum. Liolaemus smaug exhibit sexual dichromatism, absent in L. choique. The presence of ringed pattern in the tail in L. austromendocinus, L. gununakuna, and L. petrophilus is not shared by L. choique (tail without ringed pattern). It differs from the member of the L. kriegi group (Morando et al., 2003) in the lower number of dorsal scales (65-81 vs 87-110 in L. kriegi). Red coloration in cloacal region is evident in L. burgeri, absent in L. choique. 
CommentSynonymy: Medina et al. 2017 found that DNA from L. choique nested within L. smaug and L. antumalguen, but for convenience we list it with L. smaug here. Both L. smaug and L. choique were described based on 7 type specimens each although the text mentions 16 and 8 specimens used to determine variation, respectively. Although Abdala et al. 2010 showed pictures of the 3 new species (smaug, shitan, choique), no details of characters are shown. Similarly, Ruiz et al. 2019 revalidated L. choique based on morphological data from 8 specimen from unspecified locations (compared to 9 specimens of L. smaug from equally unspecified locations), without showing morphological details. The specimens are probably from around the type localities which are ~60 km apart (by air).

KWET 2012 says that the ABDALA paper was published in 2011. 
Etymologynamed after Tokien’s “Smaug”, the Golden, the last of the Middle Earth dragons. The species also exhibits a golden color.

L. choique was named after the type locality. 
References
  • Abdala, C.S.; Quinteros, A.S.; Scrocchi, G.J. & Stazzonelli, J.C. 2010. Three new species of the Liolaemus elongatus group (Iguania: Liolaemidae) from Argentina. Cuadernos de Herpetología 24 (2): 93-109 - get paper here
  • AVILA, LUCIANO JAVIER; CINTIA DEBORA MEDINA, CRISTIAN HERNAN FULVIO PEREZ, JACK W. SITES, JR. & MARIANA MORANDO 2015. Molecular phylogenetic relationships of the lizard clade Liolaemus elongatus (Iguania: Liolaemini) with the description of a new species from an isolated volcanic peak in northern Patagonia. Zootaxa 3947 (1): 067–084 - get paper here
  • AVILA, LUCIANO JAVIER; CRISTIAN HERNAN FULVIO PEREZ, CINTIA DEBORA MEDINA, JACK WALTER SITES, JR & MARIANA MORANDO 2012. A new species of lizard of the Liolaemus elongatus clade (Reptilia: Iguania: Liolaemini) from Curi Leuvu River Valley, northern Patagonia, Neuquén, Argentina. Zootaxa 3325: 37–52 - get paper here
  • AVILA, LUCIANO JAVIER; LORENA ELIZABETH MARTINEZ & MARIANA MORANDO 2013. Checklist of lizards and amphisbaenians of Argentina: an update. Zootaxa 3616 (3): 201–238 - get paper here
  • Kwet, Axel 2012. Liste der im Jahr 2011 neu beschriebenen Reptilien. Terraria-Elaphe 2012 (3): 46-57 - get paper here
  • Ruiz, Maria Soledad; Ruiz Monachesi, Mario Ricardo; Abdala, Cristian Simón 2019. Revalidación de Liolaemus choique (Abdala, Quinteros, Scrocchi y Stazzonelli, 2010) Iguania: Liolaemidae. Cuad. herpetol. 33 (1): 33-38 - get paper here
  • Troncoso-Palacios J, Díaz HA, Esquerré D, Urra FA 2015. Two new species of the Liolaemus elongatus-kriegi complex (Iguania, Liolaemidae) from Andean highlands of southern Chile. ZooKeys 500: 83-109. doi: 10.3897/zookeys.500.8725 - get paper here
  • Troncoso-Palacios J, Esquerré D, Urra FA, Díaz HA, Castro-Pastene C, Ruiz MS. 2018. The true identity of the new world iguanid lizard Liolaemus chillanensis Müller and Hellmich 1932 (Iguania: Liolaemidae) and description of a new species in the Liolaemus elongatus group. Zoological Studies 57:22; doi:10.6620/ZS.2018.57-22 - get paper here
  • Vrdoljak, Juan; Kevin Imanol Sanchez, Roberto Arreola-Ramos, Emilce Guadalupe Diaz Huesa, Alejandro Villagra, Luciano Javier Avila, Mariana Morando 2020. Testing repeatability, measurement error and species differentiation when using geometric morphometrics on complex shapes: a case study of Patagonian lizards of the genus Liolaemus (Squamata: Liolaemini). Biological Journal of the Linnean Society, blaa079, - get paper here
  • Zimin, A., Zimin, S. V., Shine, R., Avila, L., Bauer, A., Böhm, M., Brown, R., Barki, G., de Oliveira Caetano, G. H., Castro Herrera, F., Chapple, D. G., Chirio, L., Colli, G. R., Doan, T. M., Glaw, F., Grismer, L. L., Itescu, Y., Kraus, F., LeBreton 2022. A global analysis of viviparity in squamates highlights its prevalence in cold climates. Global Ecology and Biogeography, 00, 1–16 - get paper here
 
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