Podarcis guadarramae guadarramae ( Boscá, 1916 ), Bosca, 1916

Geniez, Philippe, Sá-Sousa, Paulo, Guillaume, Claude P., Cluchier, Alexandre & Crochet, Pierre-André, 2014, Systematics of the Podarcis hispanicus complex (Sauria, Lacertidae) III: valid nomina of the western and central Iberian forms, Zootaxa 3794 (1), pp. 1-51 : 8-14

Podarcis guadarramae guadarramae ( Boscá, 1916)

Lacerta muralis guadarramae Boscá, 1916 , Boletín de la Real Sociedad Española de Historia Natural 16: 330.

Name-bearing type: Muséum National d’Histoire Naturelle MNHN 2012.0262, neotype by present designation ( Fig. 3 ). Originally an adult male from “San Ildefonso” (= La Granja de San Ildefonso, Sierra de Guadarrama, province of Segovia, Spain), holotype by monotypy, currently lost. Type locality: Village of Lozoya, Sierra de Guadarrama, province of Madrid ( Spain) [WGS84 = 40.951°N / 3.792°W] after present neotype designation. Originally “San Ildefonso”.

Diagnosis. 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. 5 B), other females with pale dorsolateral stripes fragmented in series of elongated spots like males ( Fig. 5 A); juveniles with similar pattern ( Figs. 5 E & 5F); pileus strongly spotted with black; ground color of the dorsum generally brown, rarely with green hue except in mountains ( Fig. 4 F), 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. 5 C), underside often white in females, infrequently so in males ( Fig. 5 D); 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).

Range and ecology. Podarcis guadarramae guadarramae occurs only in Spain, in the Spanish 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. Range limits to the north-west are still poorly known due to difficulty of morphological separation from P. g. l u s i t an i c u s, north-westernmost localities confirmed with genetic analysis (mtDNA sequencing) are Alba de Tormes and Ciudad Rodrigo in province of Salamanca ( Kaliontzopoulou et al. 2011) while western limit is currently La Alberca, Salamanca (Pinho et al. 2007). Further east, populations assigned to P. g. guadarramae by their typical morphology and their location reach the north of the province of Segovia where they abruptly meet P. liolepis (see contact zones below). Eastern and southern limits better known as the adjacent P. virescens and P. liolepis are reasonably easy to separate morphologically (see below). The limit of the distribution south of the Spanish Central System seems to follow closely the transition between the dry, sparsely vegetated plateau of Central Spain (inhabited by virescens ) and the more vegetated foothills of the Central System massifs (pers. obs.). Podarcis guadarramae guadarramae is mostly a rock lizard adapted to rupicolous habitats including granite rocks, artificial stone walls, rubbles and detritus etc., mostly on siliceous substratum. It reaches at least 2,000 m a.s.l. in the Sierra de Gredos (pers. obs.). In parts of its range where there is no competitors (e.g. P. muralis or P. carbonelli ), it can also live on the ground, especially in pine Pinus pinaster forests on siliceous sandy soils. An (apparently) isolated population inhabits the Castillo de Trujillo (province of Cáceres, Spain), surrounded by localities where only type 2 has been documented; first photographed by V. Joubert in 1995 (photos in PG photo collection, see Appendix 1), this occurrence was later confirmed by genetic data ( Pinho et al. 2006, specimen Trj1, collected exactly at the same location as confirmed by Pinho pers. com.). This occurrence seems to be linked with the ecological situation of Trujillo which is built on a rocky outcrop emerging from the surrounding plains. In addition, Kaliontzopoulou et al. (2011) report an isolated occurrence of P. g. guadarramae in Torrejón de la Calzada (province of Madrid), a locality surrounded by P. virescens populations and in a typical ecological situation for that species. Further sampling would be needed to fully exclude a mistake in sample labeling or sample processing in the lab.

Situation in contact zones with other taxa. Podarcis guadarramae guadarramae can coexist (sometimes in syntopy) with two other Podarcis species: P. muralis (e.g. Sierra de Guadarrama) and P. carbonelli (e.g. Sierra de la Peña de Francia). It is parapatric with P. virescens south of the Central System (cf. below), and with P. l i o l e p i s along a contact zone that runs from the north of the province of Segovia (Villaverde de Iscar, Remondo, Chañe, Arroyo de Cuéllar, Campo de Cuéllar, Valleruela de Sepúlveda et Duruelo) to the easternmost foothills of the Central Spanish System south of Atienza (province of Guadalajara). The contact zone in the province of Segovia runs precisely along the border between cultivated calcareous plateau in the north (home of P. l i ol e pi s) and sandy siliceous plains with Pinus pinaster woods (home of P. g. guadarramae ). We have identified very few localities where P. g. guadarramae coexists with these species, as populations are usually a few kilometres apart. Within the abrupt contact zone with P. liolepis in the north of the province of Segovia pure populations of either species were found only tens to hundreds of meters apart and local syntopy has even been observed in one spot between cultivated lands on calcareous substratum and pinewoods on sand ( Geniez 2001) (see Fig. 6 ). No visible morphological introgression has been observed there or in any other population close to the contact zones with P. virescens or with P. liolepis , althoug detailed analysis of morphological variation accross contact zones remain to be done. We have no personal information on contact zones with lusitanicus (but see “systematic ranks of the new taxa” above).

Geographical variation. none evident to us but mountain populations tend to have a more contrasted coloration with larger proportion of dark elements, more contrasted pale dorsolateral lines and sometimes a green hue on the dorsum.

Comparison with other species within the Podarcis hispanicus complex. Separation of Podarcis bocagei and P. carbonelli from the other forms of the P. hispanicus complex has been well treated in e.g. Engelmann et al. (1993), Salvador & Pleguezuelos (2002), Kwet (2009), Arnold & Ovenden (2010), Glandt (2010). Separation of P. hispanicus has been covered in Geniez et al. (2007) and Glandt (2010), separation from P. liolepis in Renoult et al. (2010a). See P. virescens account below for separation from that species. Podarcis guadarramae guadarramae is most similar to P. g. lusitanicus but many populations can be separated by coloration patterns of adult males. Average differences in pholidosis and morphometry (see Kaliontzopoulou et al. 2012 and lusitanicus account below) are of little use for identification, even if lusitanicus is often a visibly more flattened lizard. In P. g. lusitanicus , pale spots in the light dorsolateral stripes are usually more obvious and better separated from each other because they stand out from a very dark background; the dark supradorsolateral stripes are broader and more fragmented, often reducing the pale background coloration to a narrow mid-dorsal band (dark supradorsolateral bands are narrower with more regular inner edges in guadarramae ), in some males the dark coloration invades the whole dorsum, pale coloration being limited to isolated round spots or ocelli, green-backed males more frequent than in guadarramae ; north-eastern populations of lusitanicus (Cantabria, Asturias, north of the province of León, see “geographical variation” in lusitanicus account) are better differentiated as they are paler with less developed, sometimes strongly reduced dark supradorsolateral stripes. Many specimens of the wholly allopatric P. v a uc h e r i can be separated from both P. g. guadarramae and P. g. lusitanicus by their greener back (especially males from mountain populations), less flattened head and body, narrower dark supradorsolateal stripes, and rounder pale dorsolateral spots (more elongated in P. guadarramae ), although some specimens can bear an almost identical dorsal pattern; P. vaucheri generally exhibits some obvious yellow coloration on the underside which P. guadarramae always lacks.

Description of the neotype. Adult male showing the following morphological features ( Fig. 3 ): 59 longitudinal rows of dorsal scales at midbody, 28 gular scales counted along a longitudinal fictive line from the contact between the fourth pair of maxillary scales to the collar, 27 transversal rows of ventral plates from the collar to the anal plates, 16 and 17 femoral pores (left and right), 29 subdigital lamellae beneath the fourth toe, 4 supralabials in front of the subocular but a small further plate inserted between the third and the fourth supralabials, 87 scales on the temporal area, one small masseteric shield on each side, snout-vent length 58 mm, tail length 107 mm, pileus length 15.0 mm, pileus width 6.8 mm, pileus high 6.4 mm, pileus strongly marked by 12 large black blotches, dorsum grey brown with two continuous paler dorsolateral stripes, each bordered on their inner edge by a wide supra-dorsolateral black stripe with light ocelli, reaching the base of the tail; no vertebral stripe, throat whitish with a few dark dots on the sides; belly uniform pale pinkish in life (with no dark markings, a rare character in males of this form but matching the description of the holotype of Lacerta muralis guadarramae given by Boscá).

Figures

FIGURE 3. Neotype of Lacerta muralis guadarramae Boscá 1916 MNHN 2012.0262 from the village of Lozoya, Sierra de Guadarrama (province of Madrid, Spain). Note the well flattened head, the strongly black-spotted pileus, the lack of vertebral stripe, the wide light dorsolateral and black supradorsolateral stripes on the dorsum, continuing on tail, and the light orange iris. Photos P. Geniez.

FIGURE 4. Variation in males of Podarcis guadarramae guadarramae. For GPS coordinates (WGS 84 in decimal degrees), see Appendix 1. A) Spain, Sierra de Gredos, 1.5 km N. Laguna Grande, 1,897 m (province of Ávila), 17.08. 2007 (PGe. 1107). Photo O. Buisson. B) Spain, Sierra de Guadarrama, Boca del Asno, 7 km past La Granja de San Ildefonso towards Puerto de Navacerrada (province of Segovia), 19.08. 1990 (PGe. 1028). Photo P. Geniez. C) Spain, Hiendelaencina, in the village, Plaza Major, 1,080 m (province of Guadalajara), 17.07. 2010 (BEV. 10983). Photo P. Geniez & P. - A. Crochet. D) Spain, San Martín de Valdeiglesias (province of Madrid), 27.06. 2001 (BEV. 2021). Photo P. Geniez. E) Spain, El Tiemblo (province of Ávila), 27.06. 2001 (BEV. 2009). Photo P. Geniez. F) Specimen with greenish tint on the back. Spain, Sierra de Gredos, 4 km NE. Laguna Grande, 2000 m (province of Ávila), 16.07. 1992 (PGe. 1035). Photo P. Geniez.

FIGURE 5. Variation in females, undersides and juveniles of Podarcis guadarramae guadarramae. A) Female. Spain, El Tiemblo (province of Ávila). 27.06. 2001 (BEV. 2007). Photo P. Geniez. B) Female with the “ guadarramae striped pattern ”. Spain, Navalmoral (province of Ávila), 27.6. 2001 (BEV. 1999). Photo P. Geniez. C) Typical underside of a male, Spain, Navalilla, in the village (province of Segovia), 13.05. 1996 (BEV. 4487). Photo P. Geniez. D) Male with a white underside. Spain, San Martin de Valdeiglesias (province of Madrid), 27.06. 2001 (BEV. 2018). Photo P. Geniez. E) Juvenile male. Spain, San Martin de Valdeiglesias (province of Madrid), 27.06. 2001 (BEV. 2025). Photo P. Geniez. F) Juvenile female. Spain, La Alberca, Iglesia de Nuestra Señora de la Asunción (province of Salamanca), 18.09. 2007 (PGe. 790). Photo P. - A. Crochet.

FIGURE 6. Podarcis liolepis BEV. 4122 (above) and Podarcis guadarramae guadarramae PGe. 1006, two males found in strict syntopy. Spain, Road C. 603, 1.4 km past San Miguel de Bernuy towards Cantalejo (province of Segovia), 7.05. 1986. Photos P. Geniez. Note in P. liolepis the complete vertebral stripe, the unbroken pale dorsolateral stripes, the less flattened head, the big masseteric shield and the less minute temporal and dorsal scales.