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Phymaturus extrilidus LOBO, ESPINOZA, SANABRIA & QUIROGA, 2012

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Higher TaxaLiolaemidae, Iguania, Sauria, Squamata (lizards)
Common Names 
SynonymPhymaturus extrilidus LOBO, ESPINOZA, SANABRIA & QUIROGA 2012 
DistributionArgentina (San Juan)

Type locality: Argentina, Provincia de San Juan, Departamento Ullum, Reserva Natural Don Carmelo, hillside (Sierra La Invernada) behind the field station, 30°55’91’’S, 69°04’98’’W, 3133 m elevation.  
Reproductionviviparous (not imputed, fide Zimin et al. 2022) 
TypesHolotype: MCN-UNSa 2657, adult male, R. E. Espinoza, F. Lobo, L. Quiroga, and E. Sanabria, 12 December 2007. Paratypes.—MCN-UNSa 2655–56, 2665–66, 2669–71, 2673, 2721–35, 2737 (MCN-UNSa 2656, 2665–66 are skeletons, 5 adult males (Fig. 2A, B), 15 adult females, 1 juvenile), same data as holotype; MCN-UNSa 2709–20 (MCN-UNSa 2713: skeleton), 6 adult males, 1 adult female, 2 juvenile males, 3 neonate females (Fig. 2C, D, in part), Argentina, Provincia de San Juan, Departamento Ullum, Reserva Natural Don Carmelo, Aguada la Pinchagua, 30u589660S, 69u059210W, 3122 m, same collectors as holotype, 13 December 2007. 
DiagnosisDiagnosis: Phymaturus extrilidus can be differentiated from P. calcogaster, P. castillensis, P. ceii, P. desuetus, P. etheridgei, P. excelsus, P. felixi, P. indistinctus, P. manuelae, P. nevadoi, P. patagonicus, P. payuniae, P. somuncurensis, P. spectabilis, P. spurcus, P. tenebrosus, P. videlai, and P. zapalensis (i.e., members of the P. patagonicus group; Table 1) because these species have (1) an unfragmented subocular, (2) no separation between the subocular and supralabials, (3) elongate, laminar, and imbricate superciliaries, (4) fewer ventral, gular, and upper ciliaries, (5) unenlarged midvertebral scales, and (6) smooth dorsal caudals. In contrast, P. extrilidus has (1) a fragmented subocular, (2) a separation between the subocular and supralabials (or a preocular–lorilabial row), (3) square, nonimbricate superciliaries, (4) a high number of ventral, gular, and upper ciliaries, (5) a midvertebral band of slightly enlarged scales, and (6) rugose dorsal caudals with strongly projected mucrons (i.e., a spiny tail). Phymaturus extrilidus can be differentiated from P. alicahuense, P. darwini, P. dorsimaculatus, P. maulense, P. querque, P. roigorum, P. palluma, P. verdugo, and P. vociferator (i.e., non-Puna clade members of the P. palluma group; Table 1), which have black dorsal reticulations and bright yellow to orange tails in males (brown to light brown in P. extrilidus), yet lack the usually dispersed, brown dorsal spots, which sometimes merge to form irregular line segments (Figs. 2A, 3) of the new species and P. antofagastensis, P. laurenti, P. mallimaccii, P. paihuanense, and P. punae (i.e., members of the Puna clade; Table 1). Finally, P. extrilidus can be differentiated from other members of the Puna clade in the following ways: P. antofagastensis has enlarged scales in the center of the chest, which are absent in P. extrilidus. The new species has a conspicuous, yellow scapular spot (Fig. 3), which is not present in P. antofagastensis and P. punae. Most male and female P. mallimaccii and P. punae have light brown vertebral stripes, which are not present or are rare in male P. extrilidus. Melanism of the dorsal neck is complete in P. mallimaccii, but incomplete in P. extrilidus. The antehumeral fold is bordered by enlarged scales in P. extrilidus, but these scales are not enlarged in P. punae. Male P. laurenti have enlarged postcloacal scales (Lobo et al., 2010a: fig. 15B), enlarged scales in the center of chest, slightly keeled tarsal scales, orange flank color in females, no scapular spots, and similarly sized preocular and canthal scales. In contrast, male P. extrilidus lack enlarged postcloacal scales and enlarged scales in the center of chest, have strongly keeled tarsal scales, females lack orange flank color, and this new species has conspicuous scapular spots and larger preoculars than canthals. In contrast to the new species, Phymaturus paihuanense (Núñez et al. 2010) lack scapular spots, have light yellow to cream background color, lack melanistic heads, and only exhibit melanism in the lateral region of neck. 
CommentAbundance: only known from the type locality (Meiri et al. 2017). 
EtymologyThe specific epithet extrilidus is a Latin adjective for unterrified or dauntless, which is in reference to the approachability of this new species in nature. The boldness of these lizards is in contrast to the authors’ experience with most other species of Phymaturus, which tend to seek refuge in rock crevices at the slightest disturbance—even when approached slowly from a distance. 
  • 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
  • Beninato, V A; Pérez, M A & Borghi, C E; 2019. Phymaturus extrilidus Predation. Herpetological Review 50 (3): 580 - get paper here
  • LOBO, FERNANDO; CRISTIAN ABDALA & SOLEDAD VALDECANTOS 2012. Morphological diversity and phylogenetic relationships within a South-American clade of iguanian lizards (Liolaemidae: Phymaturus). Zootaxa 3315: 1–41 - get paper here
  • Lobo, Fernando; Robert E. Espinoza, Eduardo A. Sanabria, and Lorena B. Quiroga 2012. A New Phymaturus (Iguania: Liolaemidae) from the Southern Extreme of the Argentine Puna. Copeia 2012 (1): 12-22. - get paper here
  • Meiri, Shai; Aaron M. Bauer, Allen Allison, Fernando Castro-Herrera, Laurent Chirio, Guarino Colli, Indraneil Das, Tiffany M. Doan, Frank Glaw, Lee L. Grismer, Marinus Hoogmoed, Fred Kraus, Matthew LeBreton, Danny Meirte, Zoltán T. Nagy, Cristiano d 2017. Extinct, obscure or imaginary: the lizard species with the smallest ranges. Diversity and Distributions - get paper here
  • Pérez, Mauricio Armando, Carlos Eduardo Borghi and Verónica Alejandra Beninato. 2013. Phymaturus extrilidus predation. Herpetological Review 44 (4): 679 - get paper here
  • Ramallo, Geraldine, Bursey, Charles R., Goldberg, Stephen R., Castillo, Gabriel and Acosta, Juan Carlos 2017. Phymaturus extrilidus Endoparasites. Herpetological Review 48 (1): 198 - get paper here
  • Troncoso-Palacios, Jaime and Damien Esquerré 2014. A new species of Phymaturus of the P. mallimaccii Group from the Andes of central Chile (Iguania: Liolaemidae). Phyllomedusa 13 (1):3–15 - 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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