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Podarcis guadarramae (BOSCÁ, 1916)

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Higher TaxaLacertidae, Lacertinae, Sauria, Lacertoidea, Squamata (lizards)
Subspecies 
Common NamesE: Guadarrama Wall Lizard 
SynonymLacerta muralis guadarramae BOSCÁ 1916
Podarcis guadarramae guadarramae — GENIEZ et al. 2014
Podarcis guadarramae — SPEYBROECK et al. 2020
Podarcis guadarramae — CAEIRO-DIAS et al. 2021 
DistributionCW Spain (Central System mountains, including Sierras de Guadarrama, Béjar, Gredos, Peña de Francia and Gata and the smaller sierras around these main massifs, and in siliceous plains with Pinus pinaster forests north of the Central System mountains)

Type locality: village of Lozoya, Sierra de Guadarrama (province of Madrid, Spain (neotype locality), WGS84 = 40.951°N, 3.792°W.  
Reproductionoviparous 
TypesNeotype: MNHN-RA 2012.0262; designated by Geniez et al. (2014). Lost holotype by monotypy was an adult male from “San Ildefonso” (= La Granja de San Ildefonso, Sierra de Guadarrama, province of Segovia, Spain). 
DiagnosisDiagnosis (guadarramae). This is the lineage referred to as Podarcis hispanicus “type 1B” by Pinho et al. (2006, 2007), Carretero (2008) and Kaliontzopoulou et al. (2011, 2012). A typical Iberian wall lizard of the Podarcis hispanicus complex characterized by the following features (see also Figs. 4 & 5): large proportion of black elements in body coloration; medium body size (average of adult male SVL: 53.8 mm, range 41 to 64, with 83% of adult males above 50 mm SVL, average of adult female SVL: 51.6 mm, range 43 mm to 59 mm, with 67% of adult females above 50 mm SVL); head distinctly flattened, body slender and flattened; vertebral stripe generally absent or vestigial, and then usually limited to the anterior part of the dorsum; pale dorsolateral stripes in males bright and contrasting, narrower than the dark supradorsolateral stripes, and reaching the anterior part of the tail, usually made of series of elongated whitish spots on a very pale background; many females with pale dorsolateral stripes straight and uninterrupted, without irregular pattern (called here “guadarramae striped pattern”, cf. Fig. 5B), other females with pale dorsolateral stripes fragmented in series of elongated spots like males (Fig. 5A); juveniles with similar pattern (Figs. 5E & 5F); pileus strongly spotted with black; ground color of the dorsum generally brown, rarely with green hue except in mountains (Fig. 4F), with a pale area between the two dark supra-dorsolateral bands usually moderately wide (more than 5 scales in 77% of males and 84% of females); throat normally whitish, exceptionally pink, with numerous black points especially in males; ventral face whitish, pink or salmon, sometimes brick red (never yellow or yellowish), frequently with the anterior part of the underside whitish becoming progressively reddish toward the belly (Fig. 5C), underside often white in females, infrequently so in males (Fig. 5D); marginal ventral plates, and more rarely medium and central plates, with a quadrangular or roundish, more rarely triangular, black mark; iris whitish to pale orange; masseteric shield generally small, absent in 34% of males and 43% of females; numerous longitudinal rows of dorsal scales at midbody (52 to 72 for males, average: 59.9 and 48 to 68 for females, average 57.9); large number of femoral pores (15 to 23 for males, average 18.5, 15 to 20 for females, average 17.7). Diagnostic positions in the DNA sequence of the mitochondrial NADH dehydrogenase subunit 4 (ND4) gene relative to other lineages of the P. hispanicus complex include a C at position 10888, T at position 10929, A at position 11097, A at position 11365 and A at position 11418 (positions numbered according to the P. muralis mitochondrion complete genome GenBank accession number NC_011607). 
CommentDistribution: see map in Geniez et al. 2014: 8 (Fig. 2), Caeiro-Dias et al. 2018: Fig. 1, Caeiro-Dias et al. 2021.

Species delimitation: Caeiro-Dias et al. 2021 provide evidence that supports the recognition of lusitanicus as a valid, but cryptic species. P. lusitanicus and P. guadarramae are the only (sub-) species of the Podarcis hispanicus complex that even experienced observers cannot (yet) reliably distinguish based on morphology. Based on their analysis P. bocagei is more closely related to guadarramae than it is to lusitanicus, hence elevation to species level would avoid the paraphyly with P. bocagei (although all 3 could be also considered subspecies of bocagei). 
EtymologyNamed after the type locality. 
References
  • Ayres, Cesar; Miguel Domínguez-Costas 2021. Arboreal behavior in the Lusitanian Wall Lizard, Podarcis guadarramae (Boscá 1916). Reptiles & Amphibians 28 (1): 13-14 - get paper here
  • Băncilă, Raluca I. and Jan W. Arntzen 2016. Isolation of lizard populations measured with molecular genetic data – Podarcis guadarramae in the Ria de Arosa archipelago. Amphibia-Reptilia, DOI: 10.1163/15685381-00003066 - get paper here
  • Boscá, E. 1916. Dos observaciones a propósito de la Lacerta muralis en España. Bol. R. Soc. Española Hist. Nat. 16: 327-330. - get paper here
  • Caeiro-Dias, Guilherme, Carla Luís, Catarina Pinho, Pierre-André Crochet, Neftalí Sillero and Antigoni Kaliontzopoulou 2018. Lack of congruence of genetic and niche divergence in Podarcis hispanicus complex. J Zool Syst - get paper here
  • Caeiro-Dias, Guilherme, Sara Rocha, Alvarina Couto, Carolina Pereira, Alan Brelsford, Pierre-André Crochet, and Catarina Pinho. 2021. Nuclear Phylogenies and Genomics of a Contact Zone Establish the Species Rank of Podarcis lusitanicus (Squamata, Lacertidae). Molecular Phylogenetics and Evolution 164: 107270 - get paper here
  • GENIEZ, PHILIPPE; PAULO SÁ-SOUSA, CLAUDE P. GUILLAUME, ALEXANDRE CLUCHIER, PIERRE-ANDRÉ CROCHET 2014. Systematics of the Podarcis hispanicus complex (Sauria, Lacertidae) III: valid nomina of the western and central Iberian forms. Zootaxa 3794: 1–51 - get paper here
  • Kwet, Axel 2014. Neues von den Iberischen Mauereidechsen um Podarcis hispanicus. Terraria-Elaphe 2014 (5): 56-57 - get paper here
  • Kwet, Axel & Benny Trapp 2014. Liste der Reptilien Europas. Draco 15 (60): 72-79 - get paper here
  • MALKMUS, R. 2017. Beobachtungen zur Kopulation von Podarcis guadarramae lusitanicus GENIEZ, SÁ-SOUSA, GUILLAUME, CLUCHIER, & CROCHET, 2014. Die Eidechse 28 (2): 59-62 - get paper here
  • Malkmus, Rudolf 2018. Quer durch die Serra de Montemuro. Terraria-Elaphe 2018 (5): 40-48 - get paper here
  • Ortega et al., 2014. Altitudinally divergent adult phenotypes in Iberian wall lizards are not driven by egg differences or hatchling growth rates. Oecología 177: 357-366 - get paper here
  • Pinho, Catarina; Antigoni Kaliontzopoulou, Carlos A Ferreira, João Gama 2022. Identification of morphologically cryptic species with computer vision models: wall lizards (Squamata: Lacertidae: Podarcis) as a case study. Zoological Journal of the Linnean Society, 2022;, zlac087, - get paper here
  • Sillero, Neftalí; Verónica Gomes 2016. Living in clusters: the local spatial segregation of a lizard community. Basic and Applied Herpetology 30: 61-75 - get paper here
  • Speybroeck, Jeroen; Wouter Beukema, Christophe Dufresnes, Uwe Fritz, Daniel Jablonski, Petros Lymberakis, Iñigo Martínez-Solano, Edoardo Razzetti, Melita Vamberger, Miguel Vences, Judit Vörös0, Pierre-André Crochet 2020. Species list of the European herpetofauna – 2020 update by the Taxonomic Committee of the Societas Europaea Herpetologica.. Amphibia-Reptilia 41: 139-189 - get paper here
 
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