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Liolaemus zabalai TRONCOSO-PALACIOS, DÍAZ, ESQUERRÉ & URRA, 2015

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Higher TaxaLiolaemidae, Iguania, Sauria, Squamata (lizards)
Subspecies 
Common Names 
SynonymLiolaemus zabalai TRONCOSO-PALACIOS, DÍAZ, ESQUERRÉ & URRA 2015
Liolaemus kriegi — DONOSO-BARROS 1974: 287 (in part)
Liolaemus kriegi — CEI 1986: 230 (in part)
Liolaemus sp? — TORRES-PÉREZ 1997: 146
Liolaemus kriegi — PINCHEIRA-DONOSO 2001: 11 (in part)
Liolaemus sp. A — MORANDO et al., 2003: 179
Liolaemus kriegi — PINCHEIRA-DONOSO & NÚÑEZ 2005: 289 (in part)
Liolaemus kriegi — MELLA 2005: 64 (in part)
Liolaemus sp. A — MEDINA et al. 2013: 27
Liolaemus sp. A — MEDINA et al. 2014: 256
Liolaemus zabalai — TRONCOSO-PALACIOS et al. 2019 
DistributionChile (Biobío Region)

Type locality: Near Los Barros, Laja Lagoon, Biobío Region, Chile. (37°31'S – 71°15'W, 1460 m elevation).  
Reproductionviviparous (not imputed, fide Zimin et al. 2022) 
TypesHolotype: SSUC Re 602 (Fig. 6). Collected by J. Troncoso-Palacios, F. Urra and H. Díaz. 07/01/2014. Paratypes. SSUC Re 598. Adult male. SSUC Re 597, 599, 600–01. Four adult females. The same data as the holotype (Figs 6 and 8). MZUC 35607, 39567. One male and one female. Malleco, Antuco Volcano, Los Barros. Unknown coll. 
DiagnosisDiagnosis: Liolaemus zabalai belongs to the kriegi clade of the elongatus-kriegi complex and is closely related to some undescribed species: Liolaemus sp. C and Liolae- mus sp. D; being more distant from the currently described species L. buergeri, L. kriegi and L. tregenzai (Fig. 7). According to Medina et al. (2014), in regards to the species of the kriegi clade L. zabalai is sympatric only with L. tregenzai at the Copahue Volcano.
With respect to the species of the kriegi clade, Liolaemus zabalai differs from L. tregenzai because the latter has 71–85 midbody scales and the males have no precloacal pores (Pincheira-Donoso and Scolaro 2007), whereas L. zabalai has 90–104 midbody scales and the males have 3–5 precloacal pores. In addition, the green-bluish ventral color of L. tregenzai is completely absent in L. zabalai. The uncorrected pairwise diffe- rence (cyt-b) between the species is 3.09% (Medina et al. 2014).
Liolaemus zabalai differs from L. kriegi in that the latter reaches 101.1 mm SVL, has reddish cloacal coloration in both sexes and has an unringed tail (Avila et al. 2003), whereas L. zabalai is smaller (max. SVL = 92.0 mm), has yellowish cloacal coloration in both sexes and has a ringed tail (in specimens with original tails). The uncorrected pairwise difference between these species is 3.79% (Medina et al. 2014).
Liolaemus zabalai differs from L. buergeri in that the latter has fewer dorsal scales (78–91; x = 84.1 ± 4.4, n = 14) than L. zabalai (86–96; x = 89.4 ± 3.2, n = 8) (Mann– Whitney U = 19.5; P = 0.01, DF = 20). Liolaemus zabalai has more loreal scales be- tween the nasal and the subocular (4–6; x = 4.3 ± 0.6, n = 8) than L. buergeri (3–4; x = 3.3 ± 0.5, n = 14) (Mann–Whitney U = 11.0; P < 0.01, DF = 20). Also, L. buergeri has a vertebral stripe on the tail, whereas L. zabalai has a ringed original tail. The limbs in L. zabalai are black with dispersed light brown spots, whereas L. buergeri has brown limbs with dispersed black spots (Fig. 8). Liolaemus zabalai and L. buergeri share basi- cally the same dorsal coloration pattern, but this is noticeably more marked and darker in L. zabalai (Fig. 8, see discussion). Based on the cyt-b locus, the uncorrected average pairwise difference between L. zabalai and L. buergeri is 2.94% (Medina et al. 2014), greater than the values reported for other Liolaemus widely accepted as valid species (see discussion). Also, L. zabalai can vocalize, a feature only documented for L. chil- iensis in the entire genus Liolaemus (Labra et al. 2013). Finally, although the ranges overlap, males of L. buergeri have 3–4 (x = 3.3) precloacal pores, whereas males of L. zabalai have 3–5 (x = 3.9) precloacal pores (Medina et al. 2014).
Compared to the other species of the elongatus-kriegi complex that occur near the known distribution of Liolaemus zabalai, the new species may be diagnosed as follows. Males of L. zabalai have precloacal pores, whereas males of L. flavipiceus and L. punmahuida lack them (Table 3). L. zabalai is larger than L. scorialis; and L. zabalai has more midbody scales than L. antumalguen, L. burmeisteri and L. choique (Table 3). 
Comment 
EtymologyThis species is named after Patricio Zabala, collection manager of the “Colección de Flora y Fauna Patricio Sánchez Reyes, Pontificia Universidad Católica de Chile” (SSUC). We dedicate this species to him because of his support of herpetological research in Chile, allowing us to review and deposit material in SSUC, and especially for his friendship. 
References
  • Cei, J.M. 1986. Reptiles del centro, centro-oeste y sur de la Argentina; herpetofauna de las zonas aridas y semiaridas. Mus. Regionale di Sci. Natur., Torino, Monogr. 4: 527 pp.
  • Demangel, Diego 2016. Reptiles en Chile. Fauna Nativa Ediciones, Santiago, 619 pp - get paper here
  • Demangel, Diego 2016. Guía de Campo - Reptiles del centro sure de Chile. Corporación Chilena de la Madera, Concepción, Chile 187 pp. - get paper here
  • DONOSO-BARROS, R., 1974. Notas herpetológicas. Bol. Soc. Biol. Concepción. 47: 285-287. - get paper here
  • Medina, Cintia Débora; Avila, Luciano Javier; Morando, Mariana 2013. Hacia una Taxonomía Integral: poniendo a prueba especies candidatas relacionadas a Liolaemus buergeri Werner 1907 (Iguania: Liolaemini) mediante análisis morfológicos. Cuadernos de Herpetología 27 (1): - get paper here
  • Mella JE 2005. In: Peñaloza APG, Novoa F, Contreras M (Eds) Guía de Campo Reptiles de Chile: Zona Central. Ediciones del Centro de Ecología Aplicada Ltda
  • Morando, M., L. J. Avila, and J. W. Sites Jr. 2003. Sampling strategies for delimiting species: genes, individuals, and populations in the Liolaemus elongatus-kriegi complex (Squamata: Liolaemidae) in Andean-Patagonian South America. Systematic Biology 52: 159–185. - get paper here
  • Pincheira-Donoso D 2001. Distribución geográfica de Liolaemus buergeri Werner, 1907 (Reptilia- Tropiduridae) en Chile y Argentina nuevos datos y comentarios. Noticiario Mensual del Museo Nacional de Historia Natural, Chile 346: 8–11
  • PINCHEIRA-DONOSO, Daniel & HERMAN NÚÑEZ 2005. The Chilean species of the genus Liolaemus Wiegmann, 1834 (Iguania, Tropiduridae, Liolaeminae). Taxonomy, systematics and evolution [in Spanish] Mus Nac Hist Nat Chile Publ Ocas 59: 1-486 - get paper here
  • Reyes-Olivares, C. and Labra, A. 2017. Emission of sounds by native lizards of Chile: the state of the art. Boletín Chileno de Herpetología. 4: 1-9 - get paper here
  • Torres-Pérez F 1997. Antecedentes para dos nuevas formas de saurios del Parque Nacional Laguna del Laja (VIII Región). Noticiero de Biología 5(4): 146
  • 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, Jaime; Yery Marambio-Alfaro, Diego Ramírez-Alvarez, Jorge Valdés Saavedra 2019. Phylogenetic position of two species of the Liolaemus elongatus-kriegi Complex and a new northern limit for L. buergeri (Squamata: Liolaemidae). Phyllomedusa 18 (1): 115-121 - 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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