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Liolaemus pantherinus PELLEGRIN, 1909

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Higher TaxaLiolaemidae, Iguania, Sauria, Squamata (lizards)
Common NamesE: Panther Tree Iguana
S: Lagartija pantera 
SynonymLiolaemus pantherinus PELLEGRIN 1909: 324
Liolaemus annectens orientalis MÜLLER 1924: 81
Liolaemus pantherinus — DONOSO-BARROS 1966: 327
Liolaemus pantherinus — PETERS et al. 1970: 192
Liolaemus multiformis simonsii — PETERS & DONOSO-BARROS 1970: 187
Liolaemus orientalis — CEI 1993
Liolaemus orientalis — DIRKSEN & DE LA RIVA 1999
Liolaemus simonsii — DIRKSEN & DE LA RIVA 1999
Liolaemus signifer (fide LAURENT 1992)
Liolaemus pantherinus — DIRKSEN & DE LA RIVA 1999
Liolaemus (Eulaemus) orientalis — SCHULTE et al. 2000
Liolaemus (Eulaemus) pantherinus — RAMIREZ LEYTON & PINCHEIRA-DONOSO 2005: 139
Liolaemus pantherinus — PINCHEIRA-DONOSO & NÚÑEZ 2005: 164
Liolaemus orientalis — ABDALA et al. 2008
Liolaemus pantherinus — LANGSTROTH 2021 
DistributionPeru, Bolivia (Cochabamba, Oruro, Tarija), Chile

Type locality: only generally given as “Hauts-Plateaux péruviens et boliviens“ (= Lake Titicaca region of the Altiplano i.e., high plateaus of the Andes of Peru and of Bolivia, at an altitude of around 4,000 meters.). Corrected to ““the environs of Pulacayo, Potosí Department, Bolivia, approximately 4200 m asl” by Langstroth 2021: 143.

orientalis: Bolivia (Chuquisaca, Potosí, Tarija), Argentina (Jujuy); Type locality: “Oberer Pilcomayo, zwischen Tarija und San Francisco, Bolivien”, corrected to “eastern Andes of the Upper Pilcomayo Basin of Bolivia, above 2,800 m a.s.l” by Langstroth 2021: 145.  
TypesSyntypes: MNHN-RA 1905.0343-0344
Holotype: ZMB 26405 [orientalis] 
DiagnosisDiagnosis: Liolaemus pantherinus Pellegrin 1909 can be distinguished from all other species of the L. montanus group found north of the Tropic of Capricorn by the following combination of characters: 1) adults with robust body and reaching SVL of greater than 95 mm (maximum adult SVL less than 90 mm in L. audituvelatus, L. chiribaya, L. erroneus, L. evaristoi, L. fittkaui, L. hajeki, L. insolitus, L. manueli, L. multicolor, L. nazca, L. poconchilensis, L. pulcherrimus, L. stolzmanni, L. schmidti, L. torresi, and L. victormoralesii); 2) absence of contrasting blue, cyan, green, yellow, orange, or red scales on the flanks or dorsum of adult males (present in L. annectens, L. chiribaya, L. erroneus, L. etheridgei, L. evaristoi, L. fittkaui, L. forsteri, L. lenzi, L. multicolor, L. multiformis, L. nazca, L. poconchilensis, L. pulcherrimus, L. schmidti)13; 3) absence of light dorsolateral lines or stripes in juveniles or adults (often present in L. fittkaui, , L. multicolor, L. multiformis, L. pulcherrimus, L. schmidti); 4) DSOT fewer than 70 (typically greater than 70 in L. erroneus, L. fabiani, L. forsteri, L. lenzi, L. multicolor, L. multiformis, L. pleopholis, L. schmidti) and greater than 40 (often fewer than 40 in L. jamesi and L. pachecoi); 5) absence of palpebral fringes (notable in L. audituvelatus, L. insolitus, L. manueli, L. poconchilensis, L. stolzmanni, L. torresi); and 8) geographically restricted to Cordillera Central and Cordillera Oriental of Bolivia (Oruro, Potosí, Chuquisaca, and Tarija departments) and eastern Jujuy Province of Argentina (all Peruvian endemics including L. annectens, L. chiribaya, L. etheridgei, L. melanogaster, L. nazca, L. ortizi, L. polystictus, L. robustus, L. thomasi, L. victormoralesii and L. williamsi, are separated by the northern Altiplano, the Cordillera Real, and the ranges of L. lenzi).
DSOT counts of 44 specimens (41 from Bolivia and 3 from Argentina) here assigned to L. pantherinus, yield a range of 45–67 and mean of 54.7, including the MNHN syntypes (DSOT of 51 and 58), all examined BMNH L. annectens from Bolivia, the L. annectens orientalis type series, and others from collections and field photographs. As noted above, Laurent’s (1993 “1991”) sample of 50 L. orientalis orientalis (11 from Bolivia and 39 from Argentina) yielded a range of 48–68 DSOT, which corresponds remarkably well with the counts reported here.
Abdala et al. (2008) reported 56–70 dorsals from a sample of 21 Argentine specimens identified as L. orientalis; this could be an indication that Laurent’s (1993 “1991”) earlier suspicions regarding the conspecifity of Bolivian and Argentine populations may have been warranted. Table 2 summarizes DSOT count data from a total of 190 individuals from 14 relevant species of the L. montanus group (i.e., those documented from Bolivia as of December 2018: L. chlorostictus, L. erroneus, L. fittkaui, L. forsteri, L. jamesi, L. lenzi, L. multiformis, L. pachecoi, L. puritamensis, and L. schmidti, plus L. annectens and the L. tropidonotus syntypes), including W and p-values for comparisons of each against L. pantherinus and L. multiformis. The only Bolivian species of the L. montanus group that cannot be reliably separated from L. pantherinus based on DSOT count distributions is L. fittkaui (p= 0.1503).
Of the Peruvian species of the Liolaemus montanus group, L. melanogaster Laurent 1998 is morphologically most similar to L. pantherinus and L. victormoralesii considering their low DSOT counts, robust form of the adults, and dorsal pigmentation patterns. However, L. melanogaster possesses a distinctive ventral melanism (i.e., black pigmentation vs. darker grey in some L. pantherinus) in most adults and is geographically restricted to a small area of the Pacific drainage of the Ayacucho Department, very remote from the range of L. pantherinus. Liolaemus melanogaster belongs to a well-supported clade consisting of species endemic to Peru, including L. polystictus, L. robustus, and L. williamsi, but not L. annectens or L. multiformis (Aguilar et al., 2017). My examination of the specimens (ZMH R10990–91) reported from southern Peru (“Puna, between Ajno and Tayapampa”) by Mertens (1952) as L. pantherinus reveal that they in fact belong to L. melanogaster14. Liolaemus victormoralesii differs from L. pantherinus its smaller known maximum SVL (89 mm vs >100 mm in L. pantherinus), the absence of strongly keeled dorsal scales, the absence of vestigial cloacal pores in females and its restricted range in the Ayacucho Region of Peru.
As noted above, the only Bolivian species of the Liolaemus montanus that cannot be distinguished from L. pantherinus solely based on DSOT is L. fittkaui (Table 2). However, these two species are otherwise very distinctive morphologically and ecologically. While L. pantherinus is typically drab, large-bodied (e.g., ZMB 26405 measures 101 mm SVL, Etheridge, in litt.) lizard of the drier interior ranges of Oruro, Potosí, Chuquisaca, Tarija, and Jujuy, L. fittkaui lives in humid puna on the north slope of the Cordillera Oriental in Cochabamba (Fig. 6) and is a small-bodied species (50-65 mm SVL; Jiménez-Robles et al., 2016) with distinctive male and female color patterns and strongly keeled, mucronate, lanceolate scales, suggestive of species of the alticolor group of the subgenus Liolaemus. Amongst the members of the L. montanus group, L. fittkaui is most similar morphologically to L. ortizi, as originally noted by Laurent (1986), which both seem to occur similar humid puna environments on the Amazonian fringes of the eastern Andes.
Other robust-bodied and large-scaled species of the Liolaemus montanus group from Bolivia, northern Chile, and northwestern Argentina such as Liolaemus jamesi, L. aymararum, L. puritamensis, L. pachecoi, and L. scrocchii have been assigned to a putative L. dorbignyi group (Díaz Gómez, 2007; Lobo et al., 2010a; Abdala et al., 2020). Most of these species are relatively robust and can reach SVL at least 90 mm but tend to have even larger and fewer dorsal scales than L. pantherinus (see Abdala et al., 2008). The members of the “L. jamesi group” can be distinguished from L. pantherinus by a combination of external morphological characters such as generally less imbricate (in adults), larger, and fewer dorsals, distinctive coloration patterns, and by their mutually exclusive geographic ranges15. No species of the L. jamesi group are known to be sympatric with L. pantherinus but the geographically closest species are L. pachecoi and L. puritamensis, which are found to the west of the known range of L. pantherinus.
Despite having been originally described by Laurent (1993 “1991”) as a subspecies of Liolaemus orientalis, L. chlorostictus is rather distinct from most L. pantherinus based on general morphology and coloration of the adults, particularly in males. However, the juveniles of L. pantherinus and L. chlorostictus are very similar. While the trunk scales of L. chlorostictus tend to be smaller and smoother than in L. pantherinus, the DSOT distributions shows considerable overlap—56–65 (median= 62) in L. chlorostictus vs. 45–67 (median= 54) in L. pantherinus. Also, some L. chlorostictus have keeled dorsals (e.g., topotype SDSU 3518), but not as strongly keeled as in some L. orientalis. However, double-sided Wilcoxon tests permit rejection of the null hypothesis of the equal DSOT distributions (W= 63.5, p= 0.0002) among L. pantherinus and L. chlorostictus. Adult male L. chlorostictus are dominated by yellow to greenish scales with varying degrees of black scales interspersed, sometimes weakly patterned (in younger individuals), quite distinct from male L. pantherinus. Adult female L. chlorostictus can be essential unicolor as many adult L. pantherinus but often retain distinctive dorsal blotches, and may be similar to some L. schmidti or L. multiformis at first glance, often showing reddish orange markings or reddish gravid coloration. As typical within the L. montanus group, juvenile L. pantherinus and juvenile L. chlorostictus are quite similar in general aspect and pigmentation (Langstroth 2021: 147). 
CommentSynonymy: PINCHEIRA-DONOSO & NUNEZ (2003) synonymized L. islugensis with L. pantherinus although most recent authors recognize it as valid.

Distribution: not in Chile (Altiplano) islugensis is considered a separate, valid species. If islugensis were considered a synonym of pantherinus, the latter would be in Chile. See map in Langstroth 2021: 124 (Fig. 6).

Subspecies: Liolaemus orientalis chlorostictus LAURENT 1993 is now considered as a valid species.

Synonymy: Langstroth 2021 recognized Liolaemus pantherinus Pellegrin 1909 as a valid species that includes L. annectens orientalis Müller 1924 and L. multiformis simonsii.

NCBI taxonomy ID: 109468 [orientalis] 
  • Abdala CS, Aguilar-Kirigin AJ, Semhan RV, Bulacios Arroyo AL, Valdes J, Paz MM, et al. 2019. Description and phylogeny of a new species of Liolaemus (Iguania: Liolaemidae) endemic to the south of the Plurinational State of Bolivia. PLoS ONE 14(12): e0225815 - get paper here
  • Abdala, C.S.; S. Quinteros & R.E. Espinoza 2008. Two new species of Liolaemus (iguania: Liolaemidae) from the Puna of Northwestern Argentina. Herpetologica 64 (4): 458–471 - get paper here
  • Avila, L. J.; Martinez, L. E.; Morando, M. 2010. Lista de las lagartijas y anfisbaenas de Argentina: una actualización [en línea]. Ver.1.0. 1 Diciembre 2010 [Checklist of lizards and amphisbaenians of Argentina: An update]. Centro Nacional Patagónico CENPAT-CONICET. Puerto Madryn, Chubut, Argentina. - 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
  • Dirksen, L. & De la Riva, I. 1999. The lizards and amphisbaenians of Bolivia (Reptilia, Squamata): checklist, localities, and bibliography. Graellsia 55: 199-215 - get paper here
  • Donoso-Barros, R. 1966. Reptiles de Chile. Santiago: Univ. Chile, 458 + cxlvi pp.
  • Jiménez‐Robles, O, De la Riva, I. 2019. Lizards in the mist: Thermal niches constrained by habitat and microclimates in the Andes of southern Bolivia. J Biogeogr. 46: 1676– 1686 - get paper here
  • Langstroth, R.P. 2005. Adiciones probables y confirmadas para la saurofauna boliviana. Kempffiana 1 (1): 101-128
  • Langstroth, Roberto Plotkin 2021. The identities of Liolaemus signifer (Duméril & Bibron 1837), L. pantherinus Pellegrin 1909, L. schmidti (Marx 1960), and L. erroneus (Núñez & Yáñez 1984 “1983-1984”) (Squamata: Liolaemidae). Cuadernos de Herpetología 35 (S1): 111-168
  • Laurent R F 1992. On some overlooked species of the genus Liolaemus Wiegmann (Reptilia Tropiduridae) from Peru. Breviora (494): 1-33 - get paper here
  • Laurent R F. 1992. Sobre algunas especies nuevas y poco conocidas de Liolaemus (Iguanidae) de la provincia de Jujuy, Argentina. Acta Zoologica Lilloana 40 (2): 91- 108 [1991]
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  • Lobo, F.; Espinoza, R.E. and S. Quinteros 2010. A critical review and systematic discussion of recent classification proposals for liolaemid lizards. Zootaxa 2549: 1–30 - get paper here
  • Mella-Ávila, Jorge E. & Michael Venegas Ponce. 2019. Distribución, frecuencia y abundancia de reptiles en distintos ambientes de la Región de Tarapacá, norte de Chile. Boletín Chileno de Herpetología. 6: 23-33 - get paper here
  • Müller, L. 1924. Ueber neue oder seltene Mittel- und Südamerikanische Amphibien und Reptilien. Mitt. Zool. Mus. Berlin 11 (1): 75-93 [1923]
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  • Ortiz J C; Marquet P 1987. Una nueva especie de lagarto altoandino: Liolaemus islugensis (Reptilia-Iguanidae). GAYANA ZOOLOGIA 51 (1-4): 59-63 - get paper here
  • Pellegrin,J. 1909. Description de cinq lézards nouveaux des hauts-plateaux du Pérou et de la Bolivie, appartenant au genre Liolaemus. Bull. Mus. natl. Hist. nat. Paris 6: 324-329 - get paper here
  • Pincheira-Donoso, D. & Núñez, H. 2002. Situación taxonómica de Liolaemus ornatus Koslowsky 1898 y Liolaemus pantherinus Pellegrin, 1909 (Sauria, Tropiduridae, Liolaeminae). Noticiario del Museo Nacional de Historia Natural de Chile 350: 38-42
  • 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
  • QUINTEROS, ANDRÉS SEBASTIÁN & CRISTIAN SIMÓN ABDALA 2011. A new species of Liolaemus of the Liolaemus montanus section (Iguania: Liolaemidae) from Northwestern Argentina. Zootaxa 2789: 35–48 - get paper here
  • Ramírez Leyton, G. & Pincheira Donoso, D. 2005. Fauna del Altiplano y Desierto de Atacama. Phrynosaura Ediciones, 395 pp.
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  • SCHULTE II, JAMES A.; J. ROBERT MACEY, ROBERT E. ESPINOZA AND ALLAN LARSON 2000. Phylogenetic relationships in the iguanid lizard genus Liolaemus: multiple origins of viviparous reproduction and evidence for recurring Andean vicariance and dispersal. Biological Journal of the Linnean Society 69: 75–102 - get paper here
  • VALLADARES, J. PABLO; RICHARD ETHERIDGE, JAMES SCHULTE II, GERMAN MANRIQUEZ & ANGEL SPOTORNO 2002. Nueva especie de lagartija del norte de Chile, Liolaemus molinai (Reptilia: Liolaeminae). Revista Chilena de Historia Natural 75: 473-489 - get paper here
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