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Pelodytes hespericus sp. nov. (Fig. 12) Identity and diagnosis. Populations of this species have been considered as P. punctatus by Sánchez-Herráiz et al. (2000), and as lineage C by Díaz-Rodríguez et al. (2015). Assigned to the genus Pelodytes based on high morphological similarity to P. punctatus, the type species of the genus; vertical pupil; network of dark lines in the skin of tadpoles; tadpoles with sinistral spiraculum; and molecular phylogenetic relationships. Of similar size as P. punctatus; mean SVL 35.3 mm (maximum 39.0 mm) in males, 39.8 mm (43.3 mm) in females. Morphologically similar to the other three western Pelodytes species, but distinguished from all of them by no repetitions of B notes in advertisement calls according to the recordings analyzed herein (Fig. 7), from P. ibericus by longer limbs and larger body size, and from P. atlanticus sp. nov. by larger body size (Fig. 6; Table 3). Furthermore, it is distinguished from all other Pelodytes species by concordant differences in mtDNA and nDNA sequences. Holotype. EBD 34505 (JDR 2012-1), adult male collected by Jesús Díaz-Rodríguez on 9 April 2012 at Artunido (Casas de Carrasco), Sierra de Segura, Jaén province, Spain. Geo. coord: 38.119349, -2.691951. Paratypes. EBD 34506 (JDR 2012-2), adult male, and EBD 34507 (JDR 2012-3), adult female, collected on 8 April 2012 by same collector and at same locality as holotype. Individuals deposited in Doñana Biological Station with numbers EBD 28169, 28170, 28171, 28172, 28173, 28174, 28175 collected from Soria; EBD 20440 collected from Huesca, EBD 33728 and EBD 26828 collected from Almería; and specimens deposited in National Museum of Natural Sciences with the numbers MNCN 667, MNCN 669, MNCN 670, MNCN 672, MNCN 673, MNCN 684, MNCN 686, MNCN 2720, MNCN 2721, MNCN 2722, MNCN 2723, MNCN 2724, MNCN 2725 collected from Valencia; MNCN 689 collected from Ciempozuelos (Madrid); MNCN 25023, MNCN 25024, MNCN 25025, MNCN 25026, MNCN 25027, MNCN 25028, MNCN 25029, MNCN 25032, MNCN 25033, MNCN 25034, MNCN 25035, MNCN 25036 collected from La Rioja; MNCN 41414, MNCN 41416 collected from Masa (Burgos) and MNCN 24489, MNCN 24490 from Merindad del Río Ubierna (Burgos); MNCN 25020 from Teruel, MNCN 25021, MNCN 25037 from Zaragoza, MNCN 41697 from Brea de Tajo (Madrid), MNCN 41698 from Corpa (Madrid), MNCN 377 from Almodóvar del Campo (Ciudad Real); MNCN 25022 collected from Palencia and MNCN 20783 collected from Castellón. Etymology. The specific name is a latinized adjective derived from the name Hesperides (Ἑσπερίδες), Nymphs of the fading sun and daughters of the Titan Atlas, in allusion to an Iberian mythological garden. The Hesperian Massif also takes its name from the same etymology, meaning “western” in ancient greek, referring to its position in the Mediterranean sea. Description of the holotype. Adult male in excellent state of preservation. Some tissue removed ventrally from right thigh for molecular analysis. Measurements: SVL, 36.6; HW, 12.7; HL, 13.3; TD, 2.7; ED, 4.0; END, 4.3; NSD, 3.0; NND, 3.6; HAL, 10.6; FORL, 23.2; HIL, 61.6; FOL, 18.4; FOTL, 28.5; TIL, 18.4. Body relatively slender; head dorsally flattened, slightly longer than wide, wider than body; snout slightly pointed in dorsal view, rounded in lateral view; nostrils directed dorsolaterally, not protuberant, nearer to tip of snout than to eye; canthus rostralis indistinct, straight; loreal region very slightly concave, almost flat; eye moderately large, notably protuberant dorsally; interocular distance slightly smaller than horizontal eye diameter, larger than internarial and slightly smaller than eye-nostril distance; pupil vertical; tympanum externally visible, distinct, oval, wider vertically than horizontally, its horizontal diameter is 68% of maximum eye diameter, positioned near corner of mouth; supratympanic fold moderately distinct in its anterodorsal part running from posterior corner of eye to posterior corner of tympanum, indistinct in its posteroventral part running from posterior corner of tympanum to insertion of forelimb; tongue free posteriorly, of truncate shape, vomerine teeth distinct, in two patches between and at the same level as choanae; choanae moderately sized, ovoid. Premaxillary and maxillary teeth present. Arms slender, webbing almost absent, very small rudiments recognizable; comparative finger length I Colour of the holotype. The skin colour is between dark green to olive drab with light olive bands on the back and legs. The back is dotted with a pattern of small oval and linear prominences of olive green. On the lateral side, a clear band is more or less visible, from the tympanum to the insertion of posterior limbs. The ventral area, which is smooth or slightly rough, has a light colour from pale green to yellow tones. Colour in life is similar, with light greenish spots on the back, instead of drab colors, and a design of two crossed bands clear without dots (Fig. 12). Variation. In the paratypes EBD 34506 and EBD 34507, the foot webbing is much less distinctly expressed. Distribution. Pelodytes hespericus is distributed in the Iberian Peninsula and as far as known, restricted to Spain. Its range extends mainly in areas of low and mid-altitude mountains, from the sea level to 1,900 m in La Rioja (Zaldívar 2004), almost 2,000 m in Sierra Seca (Granada) and maybe higher at the locality of Castril, Granada (Fernández-Cardenete et al. 2000), but additional confirmation is required. It occupies most of the eastern and central parts of Spain (Lacomba & Sancho 1999; Barbadillo 2002b). The western range limits are in the southeast of Madrid (Martínez-Solano & García-París 2001; Martínez-Solano 2006), in the provinces of León and Zamora, and in the Montes de Toledo in Toledo and Ciudad Real. Southwards the species reaches the northern parts of Jaén and Granada provinces (Barbadillo 2002b; Ceacero et al. 2007) and coastal parts of Almería (González- Miras & Nevado 2008; Díaz-Rodríguez et al. 2015). Natural history. Lizana et al. (1986) and Esteban et al. (2004) provided notes on the natural history of this species. Pelodytes hespericus is a generalist that tolerates a wide range of climatic conditions and diverse substrates. However, it seems to avoid siliceous and some types of acid soils (Sánchez-Herráiz 2004). It can be found in semi-arid steppes in the Ebro basin and the southeast, as well as in temperate forests of beech and oak, but usually avoids the densest forest formations. It is common in abandoned quarries and cattle ponds and tolerates high levels of salinity. Specimens can reach eight years (males) and 10 years (females) of age (Esteban et al. 2004; Sánchez-Herráiz 2004). Advertisement call and reproductive behavior. As in the other western Pelodytes lineages, the call consists of two types of notes (A and B), nonetheless emitted just once time. In the recordings previously studied by Sánchez-Herráiz (2004) analyzed by us, and in numerous other individuals heard in the field, one call typically consists of one A note followed by a single B note (Fig. 7). Reciprocal calls of females have been reported (Sánchez-Herráiz 2004). Calling specimens are found at the edge or within water, with body temperatures between 10 °C –17.5°C (Sánchez-Herráiz 2004; Esteban et al. 2002). Call intensity and spectral frequency were not found to be related to the size or age of the male (Esteban et al. 2002).

Pelodytes atlanticus sp. nov. (Fig. 11) Identity. Populations of this species have been considered as P. punctatus until now (for instance, Sanchiz et al. 2002). The species was referred to as Pelodytes sp. by Crespo et al. (2008) and van de Vliet et al. (2012) and as lineage A in Díaz-Rodríguez et al. (2015). Diagnosis. Assigned to the genus Pelodytes based on high morphological similarity to P. punctatus, the type species of the genus; vertical pupil; network of dark lines in the skin of tadpoles; tadpoles with sinistral spiraculum; and molecular phylogenetic relationships. Of similar size as P. ibericus; mean SVL 31.4 mm (maximum 39.2 mm) in males, 34.7 mm (40.4 mm) in females. Morphologically similar to the other three western Pelodytes species, but distinguished from P. hespericus sp. nov. and P. punctatus by smaller body size (Table 3; Fig. 6), from P. ibericus by longer limbs and fewer B notes in advertisement calls (Figs. 6–7), and from P. hespericus sp. nov. by more B notes in advertisement calls (Fig. 7). Furthermore, distinguished from all other Pelodytes species by concordant differences in mtDNA and nDNA sequences. Holotype. EBD 34645 (GLA 03 -JDR2015), adult male collected by Jesús Díaz-Rodríguez on 29 January 2015 at Mindelo Ornithological Reserve, Mindelo, Porto, Portugal. Geo. coord: 41.318539, -8.737131. Paratypes. EBD 34646 (GLA 01 -JDR 2015) and EBD 34647 (GLA 02 -JDR 2015), adult males collected on 29 January 2015 by same collector and at same locality as holotype. ZSM 1 /2011 collected from Nives, north of Portimão; ZSM 1201/2012, collected from Barão de São Miguel; ZSM 194 /2016, male specimen collected near Vila do Bispo, Portugal; and vouchers from Lisbon Natural Museum, collected on 1983 by E.G. Crespo: MB 110 collected on 1983 by E.G. Crespo from Cacém; twelve specimens from the series MB 112, collected on 21 October 1982 by E.G. Crespo from Cascais; and ten specimens from the series MB 113 collected on 27 October 1987 by E.G. Crespo at Benfica. Etymology. The specific epithet atlanticus is a genitive adjective derived from the name of Titan Atlas who, according to the ancient greek mythology, lived beyond the strait of Gibraltar and refers to the species’ exclusive distribution along the Atlantic coast of Portugal. Description of the holotype. Adult male in excellent state of preservation. Some tissue removed ventrally from right thigh for molecular analysis. Measurements: SVL, 33.0; HW, 11.6; HL, 11.0; TD, 1.9; ED, 3.6; END, 2.7; NSD, 2.6; NND, 2.3; HAL, 9.0; FORL, 20.0; HIL, 52.6; FOL, 16.6; FOTL, 25.4; TIL, 16.5. Body relatively slender; head dorsally flattened, slightly wider than longer, wider than body; snout slightly pointed in dorsal view, rounded and flattened in lateral view; nostrils directed dorsolaterally, slightly protuberant, nearer to tip of snout than to eye; canthus rostralis indistinct, straight; loreal region very slightly concave, almost flat; eye moderately large, notably protuberant dorsally; interocular distance slightly larger than horizontal eye diameter, larger than internarial and eye-nostril distance; pupil vertical; tympanum externally visible, distinct, oval, wider vertically than horizontally, its horizontal diameter being 68% of vertical eye diameter, positioned near corner of mouth; supratympanic fold distinct in its anterodorsal part running from posterior corner of eye to level of forelimb insertion; tongue free posteriorly, of truncated shape, vomerine teeth distinct, in two patches between and at the same level as choanae; choanae moderately sized, ovoid. Premaxillary and maxillary teeth present. Arms slender, webbing almost absent, very small rudiments recognizable; comparative finger length I Colour of the holotype: The skin colour is between dark gray to olive brown with drab green bands on the back and legs. The back is dotted with small round or oval prominences of olive colors. The ventral area, which is smooth or slightly rough, has a light colour from pale green to gray tones. Nuptial pads are grass green. Colour in life (Fig. 11) is more clear with light greenish spots on the back and a design of a drab olive line from the eye to the insertion of anterior legs. Distribution. Pelodytes atlanticus is restricted to Portugal, comprising the central-eastern part of the country (Alto Alentejo), and coastal areas from Faro to Sagres in the Algarve, and from Cape of São Vicente to Cavado River, Esposende (Crespo et al. 2008; Matos et al. 2010). The ecological range is similar to that of P. ibericus with hybrids occurring in a long and narrow contact zone between this species and P. atlanticus (van de Vliet et al. 2012; Díaz-Rodríguez et al. 2015). Natural history. Poorly known. Reproductive phenology is similar to P. ibericus and takes place early in the season, starting with the first rains at mid-autumn and extending into the winter. More detailed fieldwork is needed to determine the conservation status of populations in some regions (van de Vliet et al. 2012; Nunes et al. 2014). Given that suitable breeding habitats (i.e., temporary ponds in traditional Mediterranean farmland) are disappearing at an alarming rate in Portugal (Beja & Alcazar 2003) affecting overall Mediterranean biodiversity (Stoate et al. 2009), it is urgent to assess the conservation status of this species and possibly implement conservation measures. In the Natural Reserve of Paul do Boquilobo (central Portugal), the introduction of the American crayfish (Procambarus clarkii) resulted in the extinction of local populations of P. atlanticus eight years later (Cruz et al. 2008). Advertisement call and reproductive behavior. The call is similar to that of other western Pelodytes lineages, consisting of two types of notes (A and B). The number of B notes in a call is on average lower than in P. ibericus, but higher than in P. hespericus sp. nov. (Fig. 7). Pargana (1998) found differences in call variables between Portuguese populations of P. atlanticus sp. nov.. from Lisboa, Alto Alentejo and Algarve, versus populations of P. ibericus from Baixo Alentejo and east Algarve. As with P. hespericus sp. nov., call parameters were not related to the size or age of the male. Tadpole. We assessed morphological data in one tadpole in developmental stage 37 (field number ZCMV 14042 from Vila do Bispo, BL 25.9 mm, TL 49.4 mm). The external morphology of this tadpole and one other specimen has a close similarity to those of P. punctatus, except the LTRF 4(2–4)/4(1–2) and 4(3–4)/5(1–3) (Fig. 9c).

Pelodytes punctatus (Daudin, 1802) Common Parsley frog (Fig. 8) Identity. Described as Rana punctata Daudin, 1802, from a locality in France ("aux environs de Beauvais", Department Oise, northern France) (Frost 2015). One synonym, Rana plicata Daudin, 1802, from near Montpellier, France (Frost 2015). We could not locate the type specimens in the Paris museum (see also Frost 2015), but a more intensive search of that and other French collections might be warranted to ascertain the fate of the types of these two nomina. However, because the localities of both taxa are deeply in the range of the genetic lineage D, there is no doubt about the identity of this lineage with the name Rana punctata, and the status of Rana plicata as its junior synonym. Morphology. A medium sized frog with an average SVL of 35.5 mm (maximum SVL 41.6 mm) in males and 38.8 mm (51.0 mm) in females (Table 3). As all congeners its dorsal skin has tubercles and folds forming an irregular pattern of discontinuous dorsolateral folds, with considerable variation among individuals. Ventral side whitish, dorsally with irregular greenish dots and patches which typically are centered on the tubercles and folds, and sometimes extending around these. No webbing between fingers and weakly expressed between toes, except in males (where foot webbing are clearly expressed). Males with distinct nuptial pads on fingers and arms. Further morphometric values are summarized in Table 3, and original measurements in Supplementary Table S2. Distribution. Pelodytes punctatus is widely distributed in France, extending into northwestern Italy (Liguria and Piedmont) and northeastern Spain (Catalonia) (Le Garff 1989; Salvidio & Bologna 2007). The northern limits are currently in the French department of Nord-Pas-de-Calais, where its presence is rare and limited (Duguet & Melki 2003). Fossil evidence indicates its existence further to the north in the past (Blain et al. 2014). In Italy, it is distributed in coastal Liguria (Salvidio et al. 2004), and in Piedmont (Boulenger, 1897) where it has recently been rediscovered (Andreone & Sindaco 1998; Sindaco et al. 2002). In the Iberian Peninsula, it is restricted to the northeast, including eastern Catalonia (Mart��nez-Rica 1983; Barbadillo 2002b), where it reaches its maximum altitude in the Pyrenees (2,000 m; Borr��s & Polls 1987; Guix et al. 2009). In France and Italy, P. punctatus inhabits plains (Guy��tant et al. 1999; Salvidio et al. 2004; Pottier 2008), is absent from most of the French Pyrenees (Boulenger, 1897; Guy��tant & Geniez 2013) and only exceeds elevations of 1,000 m in the Massif Central and in the Alpes Maritimes (Salvidio & Bologna 2007). The fossil record suggests an ancient presence of P. punctatus at the east of its present distribution (Blain & Bail��n 2003; Delfino 2004). The current distribution is roughly within the region of subtropical climate with dry summers in the Koppen-Geiger classification (Peel et al. 2007), and extends into the oceanic temperate region in western France and the northeast of the Iberian Peninsula. Natural history. Numerous studies on Pelodytes biology have been performed in France, Italy and northeastern Spain and therefore the respective data can be reliably assigned to P. punctatus. This encompasses general accounts on the biology of the species (e.g., Elzen 1975, 1976), diet analysis of adults from Catalonia (Bea et al. 1994), and diet preferences of tadpoles (Richter-Boix et al. 2007). Reproduction is related to an increase in seasonal rainfall and occurs early in the year, usually before that of other anuran species, with two breeding periods: one at mid-fall and another in late winter and spring (Sindaco & Andreone 1988; Toxopeus et al. 1993; Guy��tant et al. 1999; Boix et al. 2004; Egea-Serrano et al. 2005; Richter-Boix et al. 2006; Petitot et al. 2014). Reproduction in autumn is often minor and irregular (Cayuela et al. 2012), however, in southern populations the breeding season extends all year round (A. Montori pers. com.). This reproductive strategy may reduce interspecific competition (Jakob et al. 2003; Richter-Boix et al. 2006) and does not result in an inhibition of gene flow between early and late reproduction groups (Jourdan-Pineau et al. 2012). Breeding occurs in seasonal water bodies, mainly slow-flowing streams and ephemeral ponds located in cultivated areas, grasslands, shrubs and meadows (although the ponds often are more permanent than those used by P. atlanticus sp. nov. and P. ibericus). The species is frequently found at the entrance of caves and karstic sites (Elzen 1976; Olague & Lagares 2000; S��nchez-Herr��iz 2004; Blain et al. 2008b; Vento & P��rez 2011). Pelodytes punctatus tolerates high levels of salinity and shows a preference for clayeye soils as P. hespericus sp. nov. (but unlike P. ibericus, see however Reques and Tejedo 2014). Eggs are black and small (diameter 1.5���2 mm), arranged in strings of 70���80 mm length and containing 40 ��� 700 eggs. Overall clutch size can be up to 1,600 eggs (Boulenger, 1897; Lanza 1983; Toxopeus et al. 1993). Hatching success might be related to water oxygen content (Guy��tant et al. 1999). The incubation period is 8���9 days at 14���20��C (N��llert & N��llert 1992). Larvae reach metamorphosis in roughly two months but this period can be delayed to up to four months (Sindaco & Andreone 1988; Guy��tant et al. 1999), and up to eight months in hibernating larvae (Lataste, 1876). Phenotypic plasticity in morphology and larval development has been documented (Johansson and Richter-Boix 2013). Males reach sexual maturity at one year of age and females at two years and longevity can reach 8 years (in males) and 10 years (in females), similarly to other species of the genus (Esteban et al. 2004, Erişmiş et al. 2011). The sex ratio within a population is close to 1:1 (Toxopeus et al. 1993). The IUCN threat status of P. punctatus is Least Concern (Deno��l et al. 2009); however, in the edge of its range (north of France, Italy and east of Spain), the species is generally scarce and highly threatened, due to the loss of breeding sites, close to local extinction in some areas (Parent 1989; Salvidio et al. 2004; Escoriza 2015). Advertisement call and reproductive behavior. The advertisement call of P. punctatus has been described by several authors (Hotz 1971; Elzen 1975; Paillete et al. 1992). Several call parameters (e.g., call duration) were found to depend on the temperature (Boulenger, 1897; Paillete et al. 1992). As with all western Pelodytes, the call consists of two notes (A and B) and has been reported as typically containing only a single B note following A (Paillette et al. 1992). Our analysis shows that also sequences of several B notes can be emitted following an A (Fig. 7), and it remains to be assessed by a more detailed study whether populational variation exists in the frequency of B series. Males call from positions partially or even completely submerged in the water, and occasional episodes of mortality due to sudden drops in temperature have been observed (Montori et al. 2011). Males remain for a longer period of time than females at the breeding sites and show a marked philopatry (Toxopeus et al. 1993). Female release calls have been mentioned by Elzen (1976). The amplexus is inguinal and usually lasts a few hours (Guy��tant 1986). Tadpole. We assessed morphology in one tadpole in developmental stage 34 (field number ZCMV 14045, BL 22.2 mm, TL 44.5 mm, from Palam��s, Catalonia). The examined tadpole has an elliptical body, a narrowly rounded snout in dorsal view, protruding eyes from the surface of the body and a very short tail with a typical braided line pattern; and spiraculum sinistral. The distance between eyes is moderately wide and nares are very large, round, positioned high dorsally, and situated nearer to snout than to eye and below the eye level. Moderately wide oral disk (Fig. 9b), emarginated, upper labium is wider than lower labium, LTRF is 5(2���5)/5(1���3). The upper jaw sheath is partially keratinized with pointed serrations, moderately wide without medial convexity. The lower sheath is wide V-shaped, partially keratinized and partially hidden by the upper one. Both jaw sheaths have a smooth surface. This tadpole is characterized by the presence of a network of dark lines, as it is also typical for larvae of the anuran genera Bombina and Discoglossus, covering the whole body, in addition to the presence of sparse dark spots. The venter is transparent with irregularly spiral shape intestinal coils., Published as part of D��az-Rodr��guez, Jes��s, Gehara, Marcelo, M��rquez, Rafael, Vences, Miguel, Gon��alves, Helena, Sequeira, Fernando, Mart��nez-Solano, I��igo & Tejedo, Miguel, 2017, Integration of molecular, bioacoustical and morphological data reveals two new cryptic species of Pelodytes (Anura, Pelodytidae) from the Iberian Peninsula, pp. 1-41 in Zootaxa 4243 (1) on pages 21-23, DOI: 10.11646/zootaxa.4243.1.1, http://zenodo.org/record/398686, {"references":["Daudin, F. - M. (1802) Histoire Naturelle des Rainettes, des Grenouilles et des Crapauds. Quarto version. Levrault, Paris, 108 pp.","Frost, D. R. (2015) Amphibian Species of the World: an Online Reference. Version 6.0. American Museum of Natural History, New York. Electronic Database accessible. Available from: http: // research. amnh. org / herpetology / amphibia / index. html (accessed 1 November 2015)","Le Garff, B. (1989) Pelodytes punctatus (Daudin, 1802). In: Castanet, J. & Guyetant, R. (Eds.), Atlas de Repartition des Amphibiens et Reptiles de France. Societe Herpetologique de France, Paris, pp. 71.","Salvidio, S. & Bologna, M. A. (2007) Pelodytes punctatus. In: Lanza, B., Andreone, F., Bologna, M. A., Corti, C. & Razzetti, E. (Eds.), Fauna d'Italia. Fol. XLII. Amphibia. Calderini de Il Sole, Bologna, pp. 362 - 372.","Duguet, R. & Melki, F. (Eds.) (2003) Les Amphibiens de France, Belgique et Luxembourg. Biotope Editions, Meze, 480 pp.","Salvidio, S., Lamagni, L., Bombi, P. & Bologna, M. A. (2004) Distribution, ecology and conservation of the parsley frog (Pelodytes punctatus) in Italy (Amphibia, Pelodytidae). Bollettino di Zoologia, 71, 73 - 81.","Boulenger, G. A. (1897) The tailless batrachians of Europe. Part I. Ray Society, London, 376 pp.","Andreone, F. & Sindaco, R. (1998) Erpetologia del Piemonte e della Falle d'Aosta. Atlante degli Anfibi e dei Rettili. Museo Regionale di Scienze Naturali, Torino, 283 pp.","Sindaco, R., Biggi, E., Boano, G. & Delmastro, G. B. (2002) Novitates herpetologicae pedemontanae. Rivista Piemontese di Storia Naturale, 23, 195 - 206.","Martinez-Rica, J. P. (1983) Atlas herpetologico del Pirineo. Munibe Antropologia-Arkeologia, 35, 51 - 80.","Barbadillo, L. J. (2002 b) Pelodytes punctatus. In: Pleguezuelos, J. M., Marquez, R. & Lizana, M. (Eds.), Atlas y Libro Rojo de los Anfibios y Reptiles de Espana. Direccion General de Conservacion de la Naturaleza, Madrid, pp. 100 - 102.","Borras, S. & Polls, M. (1987) Los reptiles del prepirineo oriental (Cadi-Moixero y La Cerdana) y biogeografia de la herpetofauna de la zona. Miscellania Zoologica, 11, 309 - 318.","Guix, D., Maluquer, J. & Camprodon, J. (2009) Record altitudinal de Pelodytes punctatus (Daudin, 1802) a Catalunya i a la peninsula Iberica. Butlleti de la Societat Catalana d'Herpetologia, 18, 71 - 73.","Guyetant, R., Temmermans, W. & Avrillier, J. N. (1999) Phenologie de la reproduction chez Pelodytes punctatus Daudin, 1802 (Amphibia, Anura). Amphibia-Reptilia, 20, 149 ‾ 160.","Pottier, G. (2008) Atlas de repartition des reptiles et amphibiens de Midi-Pyrenees. Nature Midi-Pyrenees, Toulouse, 126 pp.","Guyetant, R. & Geniez, P. (2013) Pelodytes punctatus. In: Lescure, J. & de Massary, J. C. (Eds.), Atlas des Amphibiens et Reptiles de France. Biotope Editions-MNHN, Meze, pp. 104 - 105.","Blain, H. A. & Bailon, S. (2003) Les amphibiens et les reptiles des couches du Pleistocene superieur ancien du gisement d'Artenac (Charente, France). Quaternaire, 14, 85 - 95.","Delfino, M. (2004) The Middle Pleistocene herpetofauna of Valdemino cave (Liguria, North-Western Italy). Herpetological Journal, 14, 113 - 128.","Peel, M. C., Finlayson, B. L. & McMahon, T. A. (2007) Updated world map of the Koppen-Geiger climate classification. Hydrology and Earth System Sciences Discussions, 4, 439 - 473.","Elzen, P. V. (1975) Contribution a la conaissance de Pelodytes punctatus Daudin (1802) (Amphibia, Pelobatidae), etudie en Camargue. Bulletin de la Societe Zoologique de France, 100, 691 - 692.","Elzen, P. V. (1976) Remarques sur la biologie de Pelodytes punctatus Daudin 1802 (Amphibia, Pelobatidae) en Camargue. Revue Francaise d'Aquariologie, 3, 99 - 104.","Bea, A., Montori, A. & Pascual, X. (1994) Herpetofauna dels Aiguamolls de l'Emporda. In: Gosalbez, J. (Ed.), Els sistemes naturals dels Aiguamolls de l'Emporda. Treballs de la Institucio Catalana d'Historia Natural, 13, pp. 359 - 407.","Richter-Boix, A., Llorente, G. A., Montori, A. & Garcia, J. (2007) Tadpole diet selection varies with the ecological context in predictable ways. Basic and Applied Ecology, 8, 464 - 474.","Sindaco, R. & Andreone, F. (1988) Considerazioni sulla distribuzione di Pelodytes punctatus (Daudin, 1802) in territorio italiano. Atti del Museo Civico di Scienze Naturali di Trieste, 41, 161 - 167.","Toxopeus, A. G., Ohm, M. & Arntzen, J. W. (1993) Reproductive biology of the parsley frog, Pelodytes punctatus, at the northernmost part of its range. Amphibia-Reptilia, 14, 131 - 147.","Boix, D., Sala, J., Quintana, X. D. & Moreno-Amich, R. (2004) Succession of the animal community in a Mediterranean temporary pond. Journal of the North American Benthological Society, 23, 29 - 49. https: // doi. org / 10.1899 / 0887 - 3593 (2004) 023 % 3 C 0029: SOTACI % 3 E 2.0. CO; 2","Egea-Serrano, A., Oliva-Paterna, F. J. & Torralva, M. (2005) Fenologia reproductiva de la comunidad de anfibios del Noroeste de la Region de Murcia (SE Peninsula Iberica). Zoologia Baetica, 16, 59 - 72.","Richter-Boix, A., Llorente, G. A. & Montori, A. (2006) Breeding phenology of an amphibian community in a Mediterranean area. Amphibia-Reptilia, 27, 549 - 559.","Petitot, M., Manceau, N., Geniez, P. & Besnard, A. (2014) Optimizing occupancy surveys by maximizing detection probability: application to amphibian monitoring in the Mediterranean region. Ecology and Evolution, 4, 3538 - 3549. https: // doi. org / 10.1002 / ece 3.1207","Cayuela, H., Besnard, A., Bechet, A., Devictor, V. & Olivier, A. (2012) Reproductive dynamics of three amphibian species in Mediterranean wetlands: the role of local precipitation and hydrological regimes. Freshwater Biology, 57, 2629 - 2640. https: // doi. org / 10.1111 / fwb. 12034","Jakob, C., Poizat, G., Veith, M., Seitz, A. & Crivelli, A. J. (2003) Breeding phenology and larval distribution of amphibians in a Mediterranean pond network with unpredictable hydrology. Hydrobiologia, 499, 51 - 61. https: // doi. org / 10.1023 / A: 1026343618150","Jourdan-Pineau, H., David, P. & Crochet, P. A. (2012) Phenotypic plasticity allows the Mediterranean parsley frog Pelodytes punctatus to exploit two temporal niches under continuous gene flow. Molecular Ecology, 21, 876 - 886. https: // doi. org / 10.1111 / j. 1365 - 294 X. 2011.05420. x","Olague, M. C. & Lagares, J. L. (2000) Cavidades naturales del termino municipal de Ejulve (Somontano turolense): datos espeleometricos y bioespeleologicos. Revista del Instituto de Estudios Turolenses, 88, 75 - 108.","Blain, H. A., Bailon, S. & Cuenca-Bescos, G. (2008 b) The Early-Middle Pleistocene palaeoenvironmental change based on the squamate reptile and amphibian proxies at the Gran Dolina site, Atapuerca, Spain. Palaeogeography, Palaeoclimatology, Palaeoecology, 261, 177 - 192.","Vento, D. & Perez, C. (2011) Datos sobre la herpetofauna del termino municipal de Biar (Alicante, este de Espana). Boletin de la Asociacion Herpetologica Espanola, 22, 130 - 134.","Reques, R. & Tejedo, M. (2014) Los Anfibios de los Humedales del Sur de Cordoba. In: De la Cruz, J. (Coord.), Humedales cordobeses: 30 anos de proteccion. Consejeria de Medio Ambiente y Ordenacion del Territorio, Junta de Andalucia, Sevilla, pp. 159 - 164","Lanza, B. (1983) Guide per il riconoscimento celle specie animali delle acque interne italiane. 27. Amphibia, Reptilia. Consiglio Nazionale delle Richerche, Verona, 196 pp.","Nollert, A. & Nollert, C. (1992) Die Amphibien Europas. Kosmos, Stuttgart, 382 pp.","Lataste, F. (1876) Essai d'une faune herpetologique de la Gironde. Actes Societe Linneenne de Bordeaux, 30, 193 - 544.","Johansson, F. & Richter-Boix, A. (2013) Within-population developmental and morphological plasticity is mirrored in between-population differences: linking plasticity and diversity. Evolutionary Biology, 40, 494 - 503. https: // doi. org / 10.1007 / s 11692 - 013 - 9225 - 8","Esteban, M., Sanchez-Herraiz, M. J., Barbadillo, L. J. & Castanet, J. (2004) Age structure and growth in an isolated population of Pelodytes punctatus in northern Spain. Journal of Natural History, 38, 2789 ‾ 2801.","Erismis, U. C., Arikan, H., Konuk, M. & Guarino, F. M. (2011) Age structure and growth in Caucasian parsley frog Pelodytes caucasicus (Boulenger, 1896) from Turkey. Russian Journal of Herpetology, 16, 19 - 26.","Denoel, M., Beja, P., Andreone, F., Bosch, J., Miaud, C., Tejedo, M., Lizana, M., Martinez-Solano, I., Salvador, A., Garcia- Paris, M., Recuero Gil, E., Marquez, R., Cheylan, M., Diaz-Paniagua, C. & Perez-Mellado, V. (2009) Pelodytes punctatus. In: IUCN 2013. IUCN Red List of Threatened Species. Fersion 2015 - 3. Available from: http: // www. iucnredlist. org (accessed 1 November 2015)","Parent, G. H. (1989) Essai de delimitation de territoires biogeographiques pour l'herpetofaune du Benelux. Bulletin Societe Naturelle de Luxembourg, 89, 81 - 103.","Escoriza, D. (2015) Sapillo moteado - Pelodytes punctatus. In: Salvador, A., Martinez-Solano, I. (Eds.), Enciclopedia Firtual de los Fertebrados Espanoles. Museo Nacional de Ciencias Naturales, Madrid. Available from: http: // www. vertebradosibericos. org / (accessed 1 Noviembre 2015)","Hotz, H. (1971) Analyse des Paarungsrufes von Pelodytes punctatus (Daudin 1803) (Amphibia, Pelobatidae). Experientia, 27, 851 - 852.","Paillette, M., Oliveira, M. E., Rosa, H. D. & Crespo, E. G. (1992) Is there a dialect in Pelodytes punctatus from southern Portugal? Amphibia-Reptilia, 13, 97 - 108.","Montori, A., Giner, G., Bejar, X. & Alvarez, D. (2011) Descenso brusco de temperaturas y nevadas tardias como causas de mortalidad de anfibios durante el periodo reproductor. Boletin de la Asociacion Herpetologica Espanola, 22, 72 - 74.","Guyetant, R. (1986) Les amphibiens de France. Revue Francaise d'Aquariologie et Herpetologie, 1975, 24 - 28."]}