Podarcis virescens sp. nov.

Holotype: Muséum National d’Histoire Naturelle MNHN 2012.0264, formerly BEV.1898, holotype by present designation; an adult male caught by P. Geniez, P.-A. Crochet and O. Chaline on 23rd June 2001, 1 km past Villanueva de los Escuderos towards Cuenca by the road CUV-7037 (Spain, province of Cuenca) [40.0436°N / 2.2916°W], 1,014 m. a.s.l. (Fig. 12). Paratypes: BEV.1899, 1901, males, BEV.1900, female from Villanueva de los Escuderos, in the village (Spain, province of Cuenca) [40.0417°N / 2.3025°W] (see Fig. 14 for BEV.1900); BEV.1909, 1911-1912, males, BEV.1910, female, from Albalate de Zorita, in the village (NNE. Tarancón, Spain, province of Guadalajara) [40.308°N / 2.845°W] (see Fig. 13 for BEV.1911); BEV.7525, male from the motorway service area 4 km SW. of Ciempozuelos (between Aranjuez and Valdemoro, Spain, province of Madrid) [40.1341°N / 3.6563°W] (see Fig. 15); BEV.10940-941, males from 2.4 km NW. of Torrelaguna (province of Madrid) [40.8404°N / 3.5634°W]; BEV.10979, male from San Andres del Congosto, cultivated plaine 500 m east of the village (province of Guadalajara) [40.9994°N / 3.0213°W], 840 m. above sea level.

Etymology. The epithet virescens is a participle derived from the Latin verb “ viresco ” meaning “turning green, becoming green”.

Diagnosis. This is the lineage referred to as Podarcis hispanicus “ type 2” 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 (Figs. 12, 13 & 14): moderate size (adult males 40 to 62 mm, mean 54.3, adult females 41 mm to 63 mm, mean 53.5); head and body relatively robust and not particularly flattened especially in males (Figs. 12 & 13); vertebral stripe often absent or, if present, usually limited to the anterior part of the dorsum; light dorsolateral stripes in males variably marked, usually visible but varying from only marginally paler than the ground color to nearly white, and fragmented or continuous but with irregular dark border, often including series of paler spots, when fragmented these pale dorsolateral lines are generally not broken by any intrusion from the black supradorsolateral stripe but by the brown ground color of the body; in females (Fig. 14) the light dorsolateral stripes are usually continuous with straight edges, their color varying from whitish to nearly the same color as back, some females bear rows of pale ocelli instead of continuous pale dorsolateral stripes (Fig. 14 E); black supradorsolateral stripes in males highly fragmented, of same width or wider than the pale dorsolateral stripes, sometimes vestigial or absent, and carrying on the anterior part of the tail; in females dark supradorsolateral stripes narrower than the pale dorsolateral stripes or of same width, usually straight, continuous or interrupted; sometimes vestigial or absent; flanks in males dark with a row of pale ocelli often present along the flanks, in females flanks are dark bordered below by a pale stripe; pileus often uniform or weakly dark spotted, (sometimes strongly spotted); ground coloration of the dorsum in males frequently tinted green or greenish in spring, especially on the posterior part, this green coloration disappearing during summer; throat whitish, sometimes yellow, yellowish or orange, with black dots especially in males; ventral face whitish, yellowish or orange (Figs. 15 A & B), sometimes brick red, with marginal ventral plates, more rarely medium and central plates, with a black quadrangular or roundish mark, rarely triangular mark; underside of the tail and rear body usually with a distinctly yellower coloration (Fig. 15 A); masseteric shield generally medium-sized, sometimes absent (in 12.6% and 24.6% of males and females respectively); very numerous dorsal scales (average of the number of longitudinal rows counted at midbody 62.3 and 59.7 for males and females respectively, minimum 49 and maximum 74 for both sexes); iris pale orange to high orange (Fig. 9 C). 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 T at position 11181 and T at position 11274; the combination of C at position 10911 and A at position 11033 is also diagnostic (positions numbered according to the P. muralis mitochondrion complete genome GenBank accession number NC_ 011607).

Range and ecology. Podarcis virescens occurs in a large area of south-central Spain from west of Sigüenza along the southern foothills of the Central System to the south side of the Tagus River valley in Spanish Extremadura and in western Portugal (Carretero 2008, pers. obs.). From there it spreads north along the Atlantic plains to Espinho (south of the Duero River mouth, see Pérez-Mellado 2010); in Portugal it reaches south as far as the whole of the Algarve coast (Loureiro et al. 2010). The southern limits of its range in Spain are still imperfectly documented due to confusions with P. vaucheri: it runs through the sierras north of the Guadalquivir river valley (see Pinho et al. 2008, Kaliontzopoulou et al. 2011) to Cazorla (Renoult et al. 2009). From there the eastern limit runs approximately through Albacete and the western foothills of the Serranía de Cuenca to Cifuentes and a few kilometers west from Sigüenza (while Sigüenza city is inhabited by P. liolepis). It mainly inhabits plains and low plateaus where it often lives in open, agricultural landscapes. Like many species of Iberian Podarcis, it is mainly linked to human habitats (villages, isolated traditional buildings, bridges, etc…) in these open landscapes. It also inhabits more natural habitats in woody hills with rocky outcrops, and enters mountainous massifs especially in the south of its range (for example Sierra de Cazorla, Sierra Morena and other Andalusian sierras north of the Guadalquivir River). Its ecology is similar to many species of Iberian Podarcis, with a rather wide niche but a strict dependency upon hard substratum and micro relief (walls, outcrops, etc.), often in relatively humid situations.

Geographical variation. No clear geographically structured morphological variations, but genetic data indicate substantial mitochondrial diversity (Pinho et al. 2006, Kaliontzopoulou et al. 2011).

Situation in contact zones with other taxa. Podarcis virescens is widely sympatric with P. carbonelli in coastal Portugal (Carretero 2008). In the north part of its distribution, it is largely parapatric with P. g. lusitanicus (= Portuguese populations of type 1, Harris & Sá-Sousa 2001, Sá-Sousa 2000) with at most limited morphological introgression (Sá-Sousa et al. 2002, see also above). Along the southern foothills of the Spanish Central System, for example around Madrid or in the north of the province of Guadalajara, west of Sigüenza, P. virescens and P. g. guadarramae replace each other abruptly again with no evidence of morphological introgression or syntopic populations (pers. obs.). The BEV collection, however, holds one female with the “ guadarramae striped pattern” in a sample from Torrelaguna (Madrid, immediately south of the Central System foothills) which is suggestive of limited morphological introgression as this pattern has not been found in any other P. virescens populations (pers. obs.). Around Mandayona (province of Guadalajara), populations of typical virescens phenotypes were found only 2 km from populations of typical liolepis phenotype. Further south, P. v i re s c en s and P. hispanicus replace each other abruptly in the Sierra de Cazorla (the former on the north side and the latter on the south side). Further south along the Guadalquivir River it is replaced by P. vaucheri but we have no information on interaction in contact zones even if both species have been identified by their mtDNA in close proximity near Jaén and Córdoba (pers. obs., Pinho et al. 2006, Kaliontzopoulou et al. 2011).

Comparison with other species. See guadarramae account (above) for references dealing with separation from members of the Podarcis hispanicus complex not considered here. Males of P. virescens are usually quite easy to separate from P. g. guadarramae and P. g. lusitanicus (that are better treated together here as they are so similar: see also Sá-Sousa et al. 2002, Glandt 2010): less flattened, pileus usually less dark-spotted (uniform or with smaller dots but exceptions occur), pale dorsolateral stripes made of rounder pale dots (more elongated in guadarramae and lusitanicus), dark supradorsolateral stripes either narrow or, when wide, usually highly fragmented and appearing as rows of dark dots, in spring often a distinct green coloration on the rear dorsum sometimes the whole back, underparts almost always with a distinctly yellower coloration at the rear, under the thighs and the tail, iris usually more orange (often yellow or whitish in guadarramae or lusitanicus). Females of P. virescens differ from females of P. g. guadarramae and lusitanicus by their generally paler coloration, narrower dark supradorsolateral stripes (of usually the same width as the pale dorsolateral stripes), pale dorsolateral stripes usually with well visible round pale spots inside (more elongated in P. g. guadarramae and lusitanicus, or very straight and continuous in the “ guadarramae striped pattern”) and often more uniform flanks. Podarcis virescens is extremely similar to P. v a uc h e r i, and often very difficult to separate from this species without genetic data. In the lowland populations, P. vaucheri is generally larger (up to 69 mm SVL instead of 63 mm for P. virescens), slightly more robust and the back usually lacks green coloration, exhibiting at most greenish hues on the rear of the back. In Baetic Mountains, males of P. vaucheri are smaller but are usually bright green on the back (duller green in P. virescens). In addition, the yellow coloration of the underside in P. virescens is more intense under the tail, the hind legs and the rear belly, whereas it is more uniform, or more intense under the throat and chest, in P. v a uc h e r i.

Description of the holotype (Fig. 12): adult male measuring 53.5 mm of snout-vent length, 13.4 mm of pileus length, 6.22 of pileus width, 5.80 of head high (giving a high / length head ratio of 0.43), and having the following scalation features: 51 longitudinal rows of dorsal scales at mid-body, 28 gular scales counted along a line from the contact between the fourth pair of maxillary scales to the collar, 25 transversal rows of ventral plates from the collar to the anal plates, 18 femoral pores on each side, 23 subdigital lamellae beneath the fourth toe, 4 supralabials in front of the subocular, 67 scales on the temporal area, one large masseteric shield on each side. Coloration of the live animal: iris orange; pileus light brown with well-marked black spots; dorsum light greyish brown with slight greenish coloration on its posterior parts, finely black-spotted without vertebral band, with fragmented dark supradorsolateral bands much narrower than the pale area between them but wider than the pale dorsolateral stripes, pale dorsolateral stripes whitish and fragmented; a dark longitudinal band on each flank with rufous spots; tail very wide at its basis, regenerated, with a distinct orange-yellow coloration; throat white with very few small black dots on the sides; belly whitish turning yellowish on its posterior area, with undersides of thighs and tail distinctly yellow tinged; marginal ventral plates alternatively turquoise and yellowish, some of them marked with small round dark spots. Preserved specimen has the same pattern but the green coloration on the back has disappeared, and the yellow portions of the underparts are duller and less contrasting.