Abstract
Parsley frogs (Pelodytes) comprise the only genus in the family Pelodytidae, an ancient anuran lineage that split from their closest relatives over 140 million years ago. Pelodytes is a Palearctic group restricted to Western Eurasia including three extant species: the eastern species P. caucasicus, endemic to the Caucasus area, and two closely related species inhabiting Western Europe: the Iberian endemic P. ibericus and the more widespread P. punctatus. Previous studies based on mitochondrial and nuclear DNA markers have revealed the existence of two additional lineages of Parsley frogs in the Iberian Peninsula, which have been flagged as candidate species. Here, we integrate novel molecular, morphological and bioacoustical data to assess the differentiation of the four western Parsley frog lineages. Species trees and Bayesian population assignment analyses based on nuclear markers confirm previous studies and concordantly delineate four parapatric lineages with narrow hybrid zones. Mitochondrial divergence is low (< 2% pairwise distances in the 16S rRNA gene), in line with previously reported low mitochondrial substitution rates in non-neobatrachian frogs. Based on concordance between mitochondrial and nuclear markers, we conclude that four species of Parsley frogs occur in Western Europe: Pelodytes punctatus, distributed from northern Italy to northeastern Spain; Pelodytes ibericus, inhabiting southern Spain and southern Portugal; Pelodytes atlanticus sp. nov., from the Portuguese Atlantic coast; and Pelodytes hespericus sp. nov., occurring in central and eastern Spain. However, bioacoustical and morphological differentiation of these species is low, with no obvious and qualitative diagnostic characters allowing full species discrimination. Differences in the relative size of metacarpal tubercles exist but this character is variable. Pelodytes ibericus and Pelodytes atlanticus are smaller than the other two species, and P. ibericus has shorter limbs and various distinctive osteological characters. Bioacoustically, the pattern by which two different note types are combined in advertisement calls separates P. hespericus from the remaining species. Despite these differences, we emphasize that the taxonomic status of all four western Parsley frogs requires additional investigation, especially the patterns of genetic admixture across contact zones. While a status of separate species best conforms to the currently available data, alternative hypotheses are also discussed.
References
Abellán, P. & Svenning, J.C. (2014) Refugia within refugia–patterns in endemism and genetic divergence are linked to Late Quaternary climate stability in the Iberian Peninsula. Biological Journal of the Linnean Society, 113, 13–28.
https://doi.org/10.1111/bij.12309Andreone, F. & Sindaco, R. (1998) Erpetologia del Piemonte e della Valle d'Aosta. Atlante degli Anfibi e dei Rettili. Museo Regionale di Scienze Naturali, Torino, 283 pp.
Arévalo, E., Davis, S.K. & Sites, J.W. (1994) Mitochondrial DNA sequence divergence and phylogenetic relationships among eight chromosome races of the Sceloporus grammicus complex (Phrynosomatidae) in central Mexico. Systematic Biology, 43, 387–418.
https://doi.org/10.1093/sysbio/43.3.387Arnold, N., Burton, J.A. & Ovenden, D.W. (1978) A Field Guide to Reptiles and Amphibians of Britain and Europe. Harper Collins, London, 272 pp.
Arntzen, J.W., Jehle, R., Bardakci, F., Burke, T. & Wallis, G.P. (2009) Asymmetric viability of reciprocal-cross hybrids between crested and marbled newts (Triturus cristatus and T. marmoratus). Evolution, 63, 1191–1202.
https://doi.org/10.1111/j.1558-5646.2009.00611.xAvilés, J.M., Parejo, D. & Lavado, F. (1999) Análisis mediante escuchas de la distribución de Pelodytes punctatus en la provincia de Badajoz. Boletín de la Asociación Herpetológica Española, 10, 14–16.
Ayllón, E., Bustamante, P., Cabrera, F., Flox, L., Galindo, A.J., Gosálvez, R.U., Hernández, J.M., Morales, M., Torralvo, C. & Zamora, F. (2003) Preliminary distribution atlas of the amphibians and reptiles of Ciudad Real province (Castilla-La Mancha, Spain). Zoologica Baetica, 13/14, 155–202.
Babik, W., Szymura, J.M. & Rafinski, J. (2003) Nuclear markers, mitochondrial DNA, and male secondary sexual traits variation in a newt hybrid zone (Triturus vulgaris x T. montandoni). Molecular Ecology, 12, 1913–1930.
https://doi.org/10.1046/j.1365-294X.2003.01880.xBallard, J.W.O. & Whitlock, M.C. (2004) The incomplete natural history of the mitochondria. Molecular Ecology, 13, 729–744.
https://doi.org/10.1046/j.1365-294X.2003.02063.xBarbadillo, L.J., Lacomba, J.I., Pérez-Mellado, V., Sancho, V. & López-Jurado, L.F. (1999) Anfibios y reptiles de la Península Ibérica, Baleares y Canarias. Editorial GeoPlaneta, Barcelona, 419 pp.
Barbadillo, L.J. (2002a) Pelodytes ibericus. In: Pleguezuelos, J.M., Márquez, R. & Lizana, M. (Eds.), Atlas y Libro Rojo de los Anfibios y Reptiles de España. Dirección General de Conservación de la Naturaleza, Madrid, pp. 97–99.
Barbadillo, L.J. (2002b) Pelodytes punctatus. In: Pleguezuelos, J.M., Márquez, R. & Lizana, M. (Eds.), Atlas y Libro Rojo de los Anfibios y Reptiles de España. Dirección General de Conservación de la Naturaleza, Madrid, pp. 100–102.
Barley, A.J., White, J., Diesmos, A.C. & Brown, R.M. (2013) The challenge of species delimitation at the extremes: diversification without morphological change in Philippine sun skinks. Evolution, 67, 3556–3572.
https://doi.org/10.1111/evo.12219Barrow, L.N., Ralicki, H.F., Emme, S.A. & Lemmon, E.M. (2014) Species tree estimation of North American chorus frogs (Hylidae: Pseudacris) with parallel tagged amplicon sequencing. Molecular Phylogenetics and Evolution, 75, 78–90.
https://doi.org/10.1016/j.ympev.2014.02.007Bea, A., Montori, A. & Pascual, X. (1994) Herpetofauna dels Aiguamolls de l’Empordá. In: Gosálbez, J. (Ed.), Els sistemes naturals dels Aiguamolls de l’Empordá. Treballs de la Institució Catalana d’Història Natural, 13, pp. 359–407.
Beja, P. & Alcazar, R. (2003) Conservation of Mediterranean temporary ponds under agricultural intensification: an evaluation using amphibians. Biological Conservation, 114, 317–326.
https://doi.org/10.1016/S0006-3207(03)00051-XBickford, D., Lohman, D., Sodhi, N.S., Ng, P.K.L., Meier, R., Winker, K., Ingram, K. & Das, I. (2007) Cryptic species as a window on diversity and conservation. Trends in Ecology and Evolution, 22, 148–155.
https://doi.org/10.1016/j.tree.2006.11.004Blain, H.A. (2009) Contribution de la paléoherpétofaune (Amphibia & Squamata) à la connaissance de l´évolution du climat et du paysage du Pliocène supérieur au Pléistocène moyen d´Espagne. Treballs del Museu de Geologia de Barcelona, 16, 39–170.
Blain, H.A. & Bailón, S. (2003) Les amphibiens et les reptiles des couches du Pléistocène supérieur ancien du gisement d'Artenac (Charente, France). Quaternaire, 14, 85‒95.
https://doi.org/10.3406/quate.2003.1732Blain, H.A., Bailón, S. & Agust, J. (2008a) Amphibians and squamate reptiles from the latest early Pleistocene of Cueva Victoria (Murcia, southeastern Spain, SW Mediterranean): paleobiogeographic and paleoclimatic implications. Geologica Acta, 6, 345–361.
Blain, H.A., Bailón, S. & Cuenca-Bescós, G. (2008b) 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.
https://doi.org/10.1016/j.palaeo.2008.01.015Blain, H.A., Bailón, S., Cuenca-Bescós, G., Arsuaga, J.L., Bermúdez de Castro, J.M. & Carbonell, E. (2009) Long-term climate record inferred from early-middle Pleistocene amphibian and squamate reptile assemblages at the Gran Dolina Cave, Atapuerca, Spain. Journal of Human Evolution, 56, 55–65.
https://doi.org/10.1016/j.jhevol.2008.08.020Blain, H.A., López-García, J.M., Cordy, J.M., Pirson, S., Abrams, G., Di Modica, K.M. & Bonjean, D. (2014) New contribution to the study of the small-vertebrate assemblages of Scladina cave (late Pleistocene, province of Namur, Belgium). In: Middle Palaeolithic in North-West Europe: Multidisciplinary Approaches. Middle Paleolithic Conference, Moulins de Beez, Namur, pp. 50. [poster presentation, also published in the next reference]
Blain, H.A., López-García, J.M., Cordy, J.M., Pirson, S., Abrams, G., Di Modica, K. & Bonjean, D. (2014) Middle to Late Pleistocene herpetofauna from Scladina and Sous-Saint-Paul caves (Namur, Belgium). Comptes Rendus Palevol, 13, 681–690.
Böhme, M. & Ilg, A.(2003) fosFARbase. Available from: http://www.wahre-staerke.com (accessed 1 November 2015)
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%3C0029:SOTACI%3E2.0.CO;2Bonaparte, C.L.J.L. (1838) Iconographia della Fauna Italica per le Quattro Classi degli Animali Vertebrati. Tomo II. Amphibi. Fascicolo 23. Salviucci, Rome, 158 pp.
Borrás, S. & Polls, M. (1987) Los reptiles del prepirineo oriental (Cadí-Moixeró y La Cerdaña) y biogeografia de la herpetofauna de la zona. Miscellània Zoològica, 11, 309–318.
Bosch, J., Tejedo, M., Lizana, M., Beja, P., Martínez-Solano, I., Salvador, A., García-París, M., Recuero Gil, E., Márquez, R., Díaz-Paniagua, C. & Pérez-Mellado, V. (2009) Pelodytes ibericus. In: The IUCN Red List of Threatened Species e.T58055A11724130. Version 2015-3. Available from: http://www.iucnredlist.org (accessed 1 November 2015)
Boulenger, G.A. (1896) Descriptions of new batrachians in the British Museum. Annals and Magazine of Natural History, Series 6, 17, 401–406.
https://doi.org/10.1080/00222939608680389Boulenger, G.A. (1897) The tailless batrachians of Europe. Part I. Ray Society, London, 376 pp.
Busack, S.D. (1977) Zoogeography of amphibians and reptiles in Cadiz province, Spain. Annals of the Carnegie Museum, 46, 285–316.
Busack, S.D. & Jaksic, F.M. (1982) Ecological and historical correlates of Iberian herpetofaunal diversity: an analysis at regional and local levels. Journal of Biogeography, 9, 289–302.
https://doi.org/10.2307/2844716Camargo, A. & Sites Jr., J. (2013) Species delimitation: A decade after the renaissance. In: Pavlinov, I. (Ed.), The Species Problem - Ongoing Issues. InTech. [published online]
https://doi.org/10.5772/52664Cardellini, P. & Sala, M. (1987) Hemoglobin transition in the anuran Pelodytes punctatus. Comparative Biochemistry and Physiology Part A: Physiology, 86, 85–89.
https://doi.org/10.1016/0300-9629(87)90281-7Carstens, B.C., Pelletier, T.A., Reid, N.M. & Satler, J.D. (2013) How to fail at species delimitation. Molecular Ecology, 22, 4369–4383.
https://doi.org/10.1111/mec.12413Cayuela, 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.12034Ceacero, F., García-Muñoz, E., Pedrajas, L., Hidalgo, A. & Guerrero, F. (2007) Actualización herpetológica de la provincia de Jaén. In: Gos, A., Egaña-Callejo, A. & Rubio, X. (Eds.), Herpetologia iberiarraren egoera = Estado actual da Herpetologia Ibérica = Estado actual de la Herpetología Ibérica: Lehen Herpetologia Kongressua Euskal Herrian, IX Congresso Luso-Espanhol, XIII Congreso Español de Herpetología. Munibe. Suplemento. Vol. 25. Aranzadi Zientzi Elkartea, Donostia, pp. 130–139.
Colliard, C., Sicilia, A., Turrisi, G.F., Arculeo, M., Perrin, N. & Stöck, M. (2010) Strong reproductive barriers in a narrow hybrid zone of West-Mediterranean green toads (Bufo viridis subgroup) with Plio-Pleistocene divergence. BMC Evolutionary Biology 10, 232.
https://doi.org/10.1186/1471-2148-10-232Crespo, E.G., Márquez, R., Pargana, J. & Tejedo, M. (2008) Pelodytes spp. Bonaparte, 1838. In: Loureiro, A., Ferrand de Almeida, N., Carretero, M.A. & Paulo, O.S. (Eds.), Atlas dos Anfíbios e Répteis de Portugal. Instituto da Conservaçao da Natureza e da Biodiversidade, Lisboa, pp. 112–115.
Crespo, E. G. & Oliveira, M.E. (1989) Atlas da Distribução dos Anfibios e Répteis de Portugal Continental. Serviço Nacional de Parques Reservas e Conservação da Natureza, Lisboa, 98 pp.
Cruz, M.J., Segurado, P., Sousa, M. & Rebelo, R. (2008) Collapse of the amphibian community of the Paul do Boquilobo Natural Reserve (central Portugal) after the arrival of the exotic American crayfish Procambarus clarkii. Herpetological Journal, 18, 197–204.
Darevsky, I.S., Drozdov, N.N. & Orlova, V.F. (1971) The new data about geographic distribution of the Caucasian parsley frog—Pelodytes caucasicus Boulenger (Amphibia, Salientia). Vestnik Zoologie, 3, 77–78.
Darriba, D., Taboada, G.L., Doallo, R. & Posada, D. (2012) jModelTest 2: more models, new heuristics and parallel computing. Nature Methods, 9, 772.
https://doi.org/10.1038/nmeth.2109Da Silva, E. (1994) Contribución al Atlas herpetológico de la provincia de Badajoz I. Anfibios. Revista Española de Herpetología, 8, 87–94.
Daudin, F.-M. (1802) Histoire Naturelle des Rainettes, des Grenouilles et des Crapauds. Quarto version. Levrault, Paris, 108 pp.
Dayrat, B. (2005) Toward integrative taxonomy. Biological Journal of the Linnean Society, 85, 407–415.
https://doi.org/10.1111/j.1095-8312.2005.00503.xDegnan, J.H. & Rosenberg, N.A. (2009) Gene tree discordance, phylogenetic inference and the multispecies coalescent. Trends in Ecology and Evolution, 24, 332–340.
https://doi.org/10.1016/j.tree.2009.01.009Delfino, M. (2002) Erpetofaune italiane del Neogene e del Cuaternario. Tesi di Dottorato, Universitá degli Studi di Modena e Regio Emilia, Modena, 382 pp.
Delfino, M. (2004) The Middle Pleistocene herpetofauna of Valdemino cave (Liguria, North-Western Italy). Herpetological Journal, 14, 113–128.
Denoël, M., Beja, P., Andreone, F., Bosch, J., Miaud, C., Tejedo, M., Lizana, M., Martínez-Solano, I., Salvador, A., García-París, M., Recuero Gil, E., Márquez, R., Cheylan, M., Díaz-Paniagua, C. & Pérez-Mellado, V. (2009) Pelodytes punctatus. In: IUCN 2013. IUCN Red List of Threatened Species. Version 2015-3. Available from: http://www.iucnredlist.org (accessed 1 November 2015)
de Queiroz, K. (1998) The general lineage concept of species, species criteria, and the process of speciation: A conceptual unification and terminological recommendations. In: Howard, D.J. & Berlocher, S.H. (Eds.), Endless Forms: Species and Speciation. Oxford University Press, New York, pp 57–75.
de Queiroz, K. (2007) Species concepts and species delimitation. Systematic Biology, 56, 879–886.
https://doi.org/10.1080/10635150701701083Devitt, T.J., Baird, S.J. & Moritz, C. (2011) Asymmetric reproductive isolation between terminal forms of the salamander ring species Ensatina eschscholtzii revealed by fine-scale genetic analysis of a hybrid zone. BMC Evolutionary Biology, 11 (1), 245.
https://doi.org/10.1186/1471-2148-11-245Díaz-Paniagua, C. (1983) Influencia de las características del medio acuático sobre las poblaciones de larvas de anfibios en la Reserva Biológica de Doñana (Huelva, España). Doñana, Acta Vertebrata, 10, 41–53.
Díaz-Paniagua, C. (1986) Reproductive period of amphibians in the Biological Reserve of Doñana (SW Spain). In: Rocek, Z. (Ed.), Studies in herpetology. Charles University, Prague, pp. 429–432.
Díaz-Paniagua, C. (1988) Temporal segregation in larval amphibian communities in temporal ponds at a locality in SW Spain. Amphibia-Reptilia, 9, 15–26.
https://doi.org/10.1163/156853888X00161Díaz-Paniagua, C. (1989) Larval diets of two anuran species, Pelodytes punctatus and Bufo bufo, in SW Spain. Amphibia-Reptilia, 10, 71–75.
https://doi.org/10.1163/156853889X00304Díaz-Paniagua, C. (1990) Temporary ponds as breeding sites of amphibians at a locality in Southwestern Spain. Herpetological Journal, 11, 447–453.
Díaz-Paniagua, C. & Rivas C.R. (1987) Datos sobre actividad de anfibios y pequeños reptiles de Doñana (Huelva, España). Mediterránea. Serie de Estudios Biológicos, 9, 15–27.
https://doi.org/10.14198/MDTRRA1987.9.02Díaz-Paniagua, C., Gómez-Rodríguez, C., Portheault, A. & de Vries, W. (2005) Los anfibios de Doñana. Organismo Autónomo Parques Nacionales, Madrid, 181 pp.
Díaz-Rodríguez, J., Gonçalves, H., Sequeira, F., Sousa-Neves, T., Tejedo, M., Ferrand, N. & Martínez-Solano, I. (2015) Molecular evidence for cryptic candidate species in Iberian Pelodytes (Anura, Pelodytidae). Molecular Phylogenetics and Evolution, 83, 224–241.
https://doi.org/10.1016/j.ympev.2014.12.007Díaz-Rodríguez, J. (2016) Historia evolutiva de los sapillos moteados (Pelodytes spp.) en la Península Ibérica. Filogenia, filogeografía y patrones de genética de poblaciones inferidos a partir de la variación en el ADN mitocondrial, nuclear y microsatélites. Tesis doctoral, Universidad de Sevilla, Sevilla, 195 pp.
Duellman, W.E. & Trueb, L. (1986) Biology of Amphibians. McGraw-Hill, New York, 670 pp.
Duguet, R. & Melki, F. (Eds.) (2003) Les Amphibiens de France, Belgique et Luxembourg. Biotope Éditions, Mèze, 480 pp.
Earl, D.A. & von Holdt, B.M. (2012) STRUCTURE HARVESTER: a website and program for visualizing STRUCTURE output and implementing the Evanno method. Conservation Genetics Resources, 4, 359–361.
https://doi.org/10.1007/s12686-011-9548-7Egea-Serrano, A., Oliva-Paterna, F.J. & Torralva, M. (2005) Fenología reproductiva de la comunidad de anfibios del Noroeste de la Región de Murcia (SE Península Ibérica). Zoologia Baetica, 16, 59–72.
Elzen, P.V. (1975) Contribution à la conaissance de Pelodytes punctatus Daudin (1802) (Amphibia, Pelobatidae), étudié en Camargue. Bulletin de la Société Zoologique de France, 100, 691–692.
Elzen, P.V. (1976) Remarques sur la biologie de Pelodytes punctatus Daudin 1802 (Amphibia, Pelobatidae) en Camargue. Revue Française d’Aquariologie, 3, 99–104.
Erişmiş, 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.
Escoriza, D. & Ruhí, A. (2014) Macroecological patterns of amphibian assemblages in the Western Palearctic: Implications for conservation. Biological Conservation, 176, 252–261.
https://doi.org/10.1016/j.biocon.2014.05.030Escoriza, D. (2015) Sapillo moteado – Pelodytes punctatus. In: Salvador, A., Martínez-Solano, I. (Eds.), Enciclopedia Virtual de los Vertebrados Españoles. Museo Nacional de Ciencias Naturales, Madrid. Available from: http://www.vertebradosibericos.org/ (accessed 1 Noviembre 2015)
Esteban, M., Sánchez-Herráiz, M.J., Barbadillo, L.J., Castanet, J. & Márquez, R. (2002) Effects of age, size and temperature on the advertisement calls of two Spanish populations of Pelodytes punctatus. Amphibia-Reptilia, 23, 249–258.
https://doi.org/10.1163/15685380260449135Esteban, M., Sánchez-Herráiz, 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.
https://doi.org/10.1080/00222930310001618859Evanno, G., Regnaut, S. & Goudet, J. (2005) Detecting the number of clusters of individuals using the software STRUCTURE: a simulation study. Molecular Ecology, 14, 2611–2620.
https://doi.org/10.1111/j.1365-294X.2005.02553.xFalush, D., Stephens, M. & Pritchard, J.K. (2003) Inference of population structure using multilocus genotype data: linked loci and correlated allele frequencies. Genetics, 164, 1567–1587.
Fernández-Cardenete, J.R., Luzón-Ortega, J.M., Pérez-Contreras, J. & Tierno de Figueroa, J.M. (2000) Revisión de la distribución y conservación de los anfibios y reptiles en la provincia de Granada (España). Zoologica Baetica, 11, 77–104.
Folmer, O.M., Black, W., Hoeh, Lutz, R. & Vrijenhoek, R. (1994) DNA primers for amplification of mitochondrial cytochrome c oxidase subunit I from diverse metazoan invertebrates. Molecular Marine Biotechnology, 3, 294–299.
Ford, L.S. & Cannatella, D.C. (1993) The major clades of frogs. Herpetological Monographs, 7, 94–117.
https://doi.org/10.2307/1466954Fouquet, A., Gilles, A., Vences, M., Marty, C., Blanc, M. & Gemmell, N.J. (2007) Underestimation of species richness in Neotropical frogs revealed by mtDNA analyses. PLoS ONE, 2, e1109.
https://doi.org/10.1371/journal.pone.0001109Franco, A. & Rodríguez de los Santos, M. (Eds.) (2001) Libro Rojo de los Vertebrados Amenazados de Andalucía. Consejería de Medio Ambiente, Junta de Andalucía, Sevilla, 336 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)
Funk, W.C., Caminer, M. & Ron, S.R. (2012) High levels of cryptic species diversity uncovered in Amazonian frogs. Proceedings of the Royal Society of London B: Biological Sciences, 279, 1806–1814.
https://doi.org/10.1098/rspb.2011.1653García-Muñoz, E., Guerrero, F., Bicho, R.C. & Parra, G. (2011) Effects of ammonium nitrate on larval survival and growth of four Iberian amphibians. Bulletin of Environmental Contamination and Toxicology, 87, 16–20.
https://doi.org/10.1007/s00128-011-0289-9García-París, M., Buchholz, D.R. & Parra-Olea, G. (2003) Phylogenetic relationships of Pelobatoidea re-examined using mtDNA. Molecular Phylogenetics and Evolution, 28, 12–23.
https://doi.org/10.1016/S1055-7903(03)00036-8Garriga, N. & Llorente, G.A. (2011) Chondrocranial ontogeny of Pelodytes punctatus (Anura: Pelodytidae). Response to competition: geometric morphometric and allometric change analysis. Acta Zoologica, 93, 453–464.
https://doi.org/10.1111/j.1463-6395.2011.00520.xGerhardt, H. C. (1991) Female mate choice in treefrogs: static and dynamic acoustic criteria. Animal Behaviour, 42, 615–635.
Gómez, A. & Lunt, D. (2007) Refugia within refugia: patterns of phylogeographic concordance in the Iberian Peninsula. In: Weiss, S. & Ferrand, N. (Eds.), Phylogeography in Southern European Refugia: Evolutionary Perspectives on the Origins and Conservation of European Biodiversity. Springer, Dordrecht, pp. 155–188.
https://doi.org/10.1016/s0003-3472(05)80245-3González de la Vega, J.P. (1988) Anfibios y reptiles de la provincia de Huelva. Ertisa, Huelva, 121 pp.
González-Miras, E. & Nevado, J.C. (2008) Atlas de distribución de los anfibios de la provincia de Almería (sudeste ibérico, España) Boletín de la Asociación Herpetológica Española, 19, 85‒90.
Grazziotin, F.G., Monzel, M., Echeverrigaray, S. & Bonatto, S.L. (2006) Phylogeography of the Bothrops jararaca complex (Serpentes: Viperidae): past fragmentation and island colonization in the Brazilian Atlantic Forest. Molecular Ecology, 15, 3969–3982.
https://doi.org/10.1111/j.1365-294X.2006.03057.xGuix, D., Maluquer, J. & Camprodon, J. (2009) Rècord altitudinal de Pelodytes punctatus (Daudin, 1802) a Catalunya i a la península Ibèrica. Butlletí de la Societat Catalana d’Herpetologia, 18, 71–73.
Guyétant, R. (1986) Les amphibiens de France. Revue Française d’Aquariologie et Herpetologie, 1975, 24–28.
Guyétant, R. & Geniez, P. (2013) Pelodytes punctatus. In: Lescure, J. & de Massary, J.C. (Eds.), Atlas des Amphibiens et Reptiles de France. Biotope Éditions-MNHN, Mèze, pp. 104–105.
Guyétant, R., Temmermans, W. & Avrillier, J.N. (1999) Phénologie de la reproduction chez Pelodytes punctatus Daudin, 1802 (Amphibia, Anura). Amphibia-Reptilia, 20, 149−160.
Hall, T.A. (1999) BioEdit: a user-friendly biological sequence alignment editor and analysis program for Windows 95/98/NT. Nucleic Acids Symposium Series, 41, 95–98.
Hauswaldt, J.S., Ludewig, A.-K., Vences, M. & Pröhl, H. (2011) Widespread co-occurrence of divergent mitochondrial haplotype lineages in a Central American species of poison frog (Oophaga pumilio). Journal of Biogeography, 38, 711–726.
https://doi.org/10.1111/j.1365-2699.2010.02438.xHawlitschek, O., Nagy, Z.T. & Glaw, F. (2012) Island evolution and systematic revision of Comoran snakes: why and when subspecies still make sense. PLoS ONE, 7, e42970.
https://doi.org/10.1371/journal.pone.0042970Heled, J. & Drummond, A.J. (2010) Bayesian inference of species trees from multilocus data. Molecular Biology and Evolution, 27, 570–580.
https://doi.org/10.1093/molbev/msp274Hoegg, S., Vences, M., Brinkmann, H. & Meyer, A. (2004) Phylogeny and comparative substitution rates of frogs inferred from three nuclear genes. Molecular Biology and Evolution, 21, 1188–1200.
https://doi.org/10.1093/molbev/msh081Holman, J.A. (1998) Pleistocene Amphibians and Reptiles in Britain and Europe. Oxford Monographs on Geology and Geophysics 38. Oxford University Press, New York, 254 pp.
Hotz, H. (1971) Analyse des Paarungsrufes von Pelodytes punctatus (Daudin 1803) (Amphibia, Pelobatidae). Experientia, 27, 851–852.
https://doi.org/10.1007/BF02136906Huson, D.H. & Bryant, D. (2006) Application of phylogenetic networks in evolutionary studies. Molecular Biology and Evolution, 23, 254–267.
https://doi.org/10.1093/molbev/msj030Iğci , N., Akman, B., Göçmen, B., Adakul, A. & Oğuz, M.A. (2013) A new locality record of Caucasian parsley frog, Pelodytes caucasicus Boulenger, 1896 (Amphibia: Anura: Pelodytidae) in the eastern Black Sea region of Anatolia. Biharean Biologist, 7, 54–56.
Irisarri, I., San Mauro, D., Abascal, F., Ohler, A., Vences, M. & Zardoya, R. (2012) The origin of modern frogs (Neobatrachia) was accompanied by acceleration in mitochondrial and nuclear substitution rates. BMC Genomics, 13, e626.
https://doi.org/10.1186/1471-2164-13-626Irwin, D.M., Kocher, T.D. & Wilson, A.C. (1991) Evolution of the cytochrome b gene of mammals. Journal of Molecular and Evolution, 32, 128–144.
https://doi.org/10.1007/BF02515385Iskanderov, T. (2009) Current Status of the Caucasus Toad (Bufo verrucosissimus Pall., 1814) and Caucasus Parsley Frog (Pelodytes caucasicus Boul., 1896) in Azerbaijan. Status and Protection of Globally Threatened Species in the Caucasus. World Wildlife Fund, Tblisi, pp. 151–156.
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:1026343618150Jakobsson, M. & Rosenberg, N.A. (2007) CLUMPP: a cluster matching and permutation program for dealing with label switching and multimodality in analysis of population structure. Bioinformatics, 23, 1801–1806.
https://doi.org/10.1093/bioinformatics/btm233Jezkova, T., Leal, M. & Rodríguez-Robles, J.A. (2013) Genetic drift or natural selection? Hybridization and asymmetric mitochondrial introgression in two Caribbean lizards (Anolis pulchellus and Anolis krugi). Journal of Evolutionary Biology, 26, 1458–1471.
https://doi.org/10.1111/jeb.12149Johansson, 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/s11692-013-9225-8Jourdan-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.1365-294X.2011.05420.xKomaki, S., Kurabayashi, A., Islam, M.M., Tojo, K. & Sumida, M. (2012) Distributional change and epidemic introgression in overlapping areas of Japanese pond frog species over 30 years. Zoological Science, 29, 351–358.
https://doi.org/10.2108/zsj.29.351Kopelman, N.M., Mayzel, J., Jakobsson, M., Rosenberg, N.A. & Mayrose, I. (2015) CLUMPAK: a program for identifying clustering modes and packaging population structure inferences across K. Molecular Ecology Resources, 15, 1179–1191.
https://doi.org/10.1111/1755-0998.12387Kuchta, S.R., Parks, D.S., Mueller, R.L. & Wake, D.B. (2009) Closing the ring: historical biogeography of the salamander ring species Ensatina eschscholtzii. Journal of Biogeography, 36, 982–995.
https://doi.org/10.1111/j.1365-2699.2008.02052.xKuzmin, S. L. (1997) Pelodytes caucasicus (Boulenger, 1896). In: Gasc, J.P., Cabela, A., Crnobrnja-Isailovic, Dolmen, D., Grossenbacher, K., Haffner, P., Lescure, J., Martens, H., Martínez Rica, J.P., Maurin, H., Oliveira, M.E., Sofianidou, T.S., Veith, M. & Zuiderwijk, A. (Eds.), Atlas of amphibians and reptiles in Europe. Publications scientifiques du Museum national d'Histoire Naturelle, Paris, pp. 114−115.
Lacomba, V. & Sancho, V. (1999) Atlas de anfibios y reptiles de la Comunidad Valenciana. Boletín de la Asociación Herpetológica Española, 10, 2−10.
Lanza, B. (1983) Guide per il riconoscimento celle specie animali delle acque interne italiane. 27. Amphibia, Reptilia. Consiglio Nazionale delle Richerche, Verona, 196 pp.
Lataste, F. (1876) Essai d’une faune herpétologique de la Gironde. Actes Société Linnéenne de Bordeaux, 30, 193–544.
Le Garff, B. (1989) Pelodytes punctatus (Daudin, 1802). In: Castanet, J. & Guyétant, R. (Eds.), Atlas de Répartition des Amphibiens et Reptiles de France. Société Herpétologique de France, Paris, pp. 71.
Librado, P. & Rozas, J. (2009) DNAsp v5: a software for comprehensive analysis of polymorphism DNA data. Bioinformatics, 25, 1451 –1452.
https://doi.org/10.1093/bioinformatics/btp187Liu, K., Wang, F., Chen, W., Tu, L., Min, M.S., Bi, K. & Fu, J. (2010) Rampant historical mitochondrial genome introgression between two species of green pond frogs, Pelophylax nigromaculatus and P. plancyi. BMC Evolutionary Biology, 10, 201.
https://doi.org/10.1186/1471-2148-10-201Lizana. M., Ciudad Pizarro, M.J. & Pérez-Mellado, V. (1986) Uso de los recursos tróficos de una comunidad ibérica de anfibios. Revista Española de Herpetología, 1, 207–272.
Lobo, J.M., Martínez-Solano, I. & Sanchiz, B. (2016) A review of the palaeoclimatic inference potential of Iberian Quaternary fossil batrachians. Palaeobiodiversity and Palaeoenvironments, 96, 125–148.
https://doi.org/10.1007/s12549-015-0224-xMacLeod, A., Rodríguez, A., Vences, M., Orozco-terWengel, P., García, C., Trillmich, F., Gentile, G., Caccone, A., Quezada, G. & Steinfartz, S. (2015) Hybridization masks speciation in the evolutionary history of the Galápagos marine iguana. Proceedings of the Royal Society B, 282, 20150425.
https://doi.org/10.1098/rspb.2015.0425Malone, J.H. & Fontenot, B.E. (2008) Patterns of reproductive isolation in toads. PLOS ONE, 3, e3900.
https://doi.org/10.1371/journal.pone.0003900Márquez, R., Pargana, J.M. & Crespo, E.G. (2001) Acoustic competition in male Pelodytes ibericus. Interactive playback tests. Copeia, 2001, 1142–1150.
https://doi.org/10.1643/0045-8511(2001)001[1142:ACIMPI]2.0.CO;2Martínez-Rica, J. P. (1983) Atlas herpetológico del Pirineo. Munibe Antropologia-Arkeologia, 35, 51 –80.
Martínez-Solano, I. (2006) Atlas de distribución y estado de conservación de los anfibios de la Comunidad de Madrid. Graellsia, 62, 253–291.
https://doi.org/10.3989/graellsia.2006.v62.iExtra.121Martínez-Solano, I. & García-París, M. (2001) Distribución y estado de conservación de Alytes obstetricans y Pelodytes punctatus en el SE de Madrid. Boletín de la Asociación Herpetológica Española, 12, 37–41.
Martínez-Solano, I. & Sanchiz, B. (2005) Anfibios y Reptiles del Pleistoceno medio de Ambrona. In: Santonja, M. & Pérez, González, A. (Eds.), Los yacimientos paleolíticos de Ambrona y Torralba (Soria). Un siglo de investigaciones arqueológicas. Museo Arqueológico Regional, Alcal de Henares, pp. 232–239.
Mateo, J.A., Pleguezuelos, J.M., Fahd, S., Geniez, P. & Martínez-Medina, F.J. (2003) Los anfibios, los reptiles y el Estrecho de Gibraltar. Un ensayo sobre la herpetofauna de Ceuta y su entorno. Instituto de Estudios Ceutíes, Ceuta, 388 pp.
Matos, C., Sillero, N. & Soares, A. (2010) New records of Pelodytes spp. and Pleurodeles waltl outside their distribution range in northern Portugal. Herpetology Notes, 3, 293–294.
Milne, I., Wright, F., Rowe, G., Marshall, D.F., Husmeier, D. & McGuire, G. (2004) TOPALi: software for automatic identification of recombinant sequences within DNA multiple alignments. Bioinformatics, 20, 1806–1807.
https://doi.org/10.1093/bioinformatics/bth155Miralles, A. & Vences, M. (2013) New metrics for comparison of taxonomies reveal striking discrepancies among species delimitation methods in Madascincus lizards. PLoS ONE, 8, 68242.
https://doi.org/10.1371/journal.pone.0068242Mo, Y., Zhang, W., Zhou, S., Chen, T., Tang, H., Meng, Y. & Chen, W. (2013) A new species of Kaloula (Amphibia: Anura: Microhylidae) from southern Guangxi, China. Zootaxa, 3710 (2), 165–178.
https://doi.org/10.11646/zootaxa.3710.2.3Monahan, W.B., Pereira, R.J. & Wake, D.B. (2012) Ring distributions leading to species formation: a global topographic analysis of geographic barriers associated with ring species. BMC Biology, 10, 20.
https://doi.org/10.1186/1741-7007-10-20Montori, A., Giner, G., Béjar, X. & Álvarez, D. (2011) Descenso brusco de temperaturas y nevadas tardías como causas de mortalidad de anfibios durante el período reproductor. Boletín de la Asociación Herpetológica Española, 22, 72–74.
Morescalchi, A., Olmo, E. & Stingo, V. (1977) Trends of karyological evolution in pelobatoid frogs. Experientia, 33, 1577−1578.
https://doi.org/10.1007/bf01934007Moritz, C., Schneider, C.J. & Wake, D.B. (1992) Evolutionary relationships within the Ensatina eschscholtzii complex confirm the ring species interpretation. Systematic Biology, 41, 273–291.
https://doi.org/10.1093/sysbio/41.3.273Muñoz del Viejo, A., Pérez, J.L., Rodríguez, S.P. & García, J.M. (2005) Atlas de los anfibios y reptiles de la provincia de Cáceres y zonas importantes para la herpetofauna en la provincia de Badajoz, Informe Final. UNEX- Dirección General de Medio Ambiente, Consejería de Agricultura y Medio Ambiente de Extremadura, Badajoz, 68 pp.
Narins, P.M. & Capranica, R.R. (1978) Communicative significance of the two-note call of the treefrog Eleutherodactylus coqui. Journal of Comparative Physiology, 127, 1–9.
https://doi.org/10.1007/BF00611921Nöllert, A. & Nöllert, C. (1992) Die Amphibien Europas. Kosmos, Stuttgart, 382 pp.
Nunes, A.L., Orizaola, G., Laurila, A. & Rebelo, R. (2014) Morphological and life-history responses of anurans to predation by an invasive crayfish: an integrative approach. Ecology and Evolution, 4, 1491–1503.
https://doi.org/10.1002/ece3.979Olague, M.C. & Lagares, J.L. (2000) Cavidades naturales del término municipal de Ejulve (Somontano turolense): datos espeleométricos y bioespeleológicos. Revista del Instituto de Estudios Turolenses, 88, 75–108.
Oliveira, M.E., Paillette, M., Rosa, H.D. & Crespo, E.G. (1991) A natural hybrid between Hyla arborea and Hyla meridionalis detected by mating calls. Amphibia-Reptilia, 12, 15–20.
https://doi.org/10.1163/156853891X00284Padial, J.M., Miralles, A., De la Riva, I. & Vences, M. (2010) The integrative future of taxonomy. Frontiers in Zoology, 7, 16.
https://doi.org/10.1186/1742-9994-7-16Paillette, 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.
https://doi.org/10.1163/156853892X00292Palomo, J.A. (1993) Atlas provisional de los anfibios y reptiles de Extremadura. Aegypius, 11, 7–20.
Palumbi, S.R., Martin, A., Romano, S., McMillan, W.O., Stice, L. & Grabowski, G. (1991) The Simple Fool’s Guide to PCR. Version 2.0. Univ. Hawaii, Honolulu.
Parent, G.H. (1989) Essai de délimitation de territoires biogéographiques pour l'herpétofaune du Benelux. Bulletin Société Naturelle de Luxembourg, 89, 81–103.
Pargana, J.M. (1998) Características espectrais e temporais e correlações genéticas do canto de acasalamento de Pelodytes punctatus (Amphibia, Anura). Masters Thesis, Departamento de Biofísica, Universidade de Lisboa, Lisbon, 163 pp.
Pargana, J.M., Márquez, R., Reques, R., Sánchez-Herráiz, M.J., Tejedo, M. & Crespo, E.G., (2003) The mating call of Pelodytes ibericus (Anura, Pelodytidae). Herpetological Journal, 13, 199–204.
Peel, M.C., Finlayson, B.L. & McMahon, T.A. (2007) Updated world map of the Köppen-Geiger climate classification. Hydrology and Earth System Sciences Discussions, 4, 439–473.
https://doi.org/10.5194/hessd-4-439-2007Pereira, R.J., Monahan, W.B. & Wake, D.B. (2011) Predictors for reproductive isolation in a ring species complex following genetic and ecological divergence. BMC Evolutionary Biology, 11, 194.
https://doi.org/10.1186/1471-2148-11-194Petitot, 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/ece3.1207Pinho, C., Rocha, S., Carvalho, B.M., Lopes, S., Mourão, S., Vallinoto, M., Brunes, T.O., Haddad, C.F.B., Gonçalves, H. & Sequeira, F. (2010) New primers for the amplification and sequencing of nuclear loci in a taxonomically wide set of reptiles and amphibians. Conservation Genetics Resources, 2, 181–185.
https://doi.org/10.1007/s12686-009-9126-4Pleguezuelos, J.M., Márquez, R. & Lizana, M. (Eds.) (2002) Atlas y Libro Rojo de los Anfibios y Reptiles de España. Dirección General de Conservación de la Naturaleza- Asociación Herpetológica Española, Madrid, 587 pp.
Pottier, G. (2008) Atlas de répartition des reptiles et amphibiens de Midi-Pyrénées. Nature Midi-Pyrénées, Toulouse, 126 pp.
Pritchard, J.K., Stephens, M. & Donnelly, P. (2000) Inference of population structure using multilocus genotype data. Genetics, 155, 945–959.
Rakotoarison, A., Crottini, A., Müller, J., Rödel, M. O, Glaw, F. & Vences, M. (2015) Revision and phylogeny of narrow-mouthed treefrogs (Cophyla) from northern Madagascar: integration of molecular, osteological, and bioacoustic data reveals three new species. Zootaxa, 3937 (1), 61–89.
https://doi.org/10.11646/zootaxa.3937.1.3Rambaut, A., Suchard, M.A., Xie, D. & Drummond, A.J. (2014) Tracer v1.6. Available from: http://beast.bio.ed.ac.uk/Tracer (accessed 1 November 2015)
Renoult, J.P., Geniez, P., Bacquet, P., Benoit, L. & Crochet, P.A. (2009) Morphology and nuclear markers reveal extensive mitochondrial introgressions in the Iberian Wall Lizard species complex. Molecular Ecology, 18, 4298–4315.
https://doi.org/10.1111/j.1365-294X.2009.04351.xReques, R. (2000) Anfibios, Ecología y Conservación. Serie Recursos Naturales de Córdoba. Diputación de Córdoba, Delegación de Medio Ambiente y Protección Civil, Córdoba, 140 pp.
Reques, R. (2014) Sapillo moteado ibérico - Pelodytes ibericus. In: Salvador, A. & Martínez-Solano, I. (Eds.), Enciclopedia Virtual de los Vertebrados Españoles. Museo Nacional de Ciencias Naturales, Madrid. Available from: http://www.vertebradosibericos.org (accessed 1 November 2015)
Reques, R., Caro, J. & Pleguezuelos, J.M. (2006) Parajes importantes para la conservación de anfibios y reptiles en España. Volumen I y II. Consejería de Medio Ambiente. Junta de Andalucía, Sevilla, 572 pp.
Reques, R. & Tejedo, M, (1991) Dinámica y hábitats reproductivos de una comunidad de Anfibios en la Sierra de Cabra. Revista Española de Herpetología, 6, 49–54.
Reques, R. & Tejedo, M. (2014) Los Anfibios de los Humedales del Sur de Córdoba. In: De la Cruz, J. (Coord.), Humedales cordobeses: 30 años de protección. Consejería de Medio Ambiente y Ordenación del Territorio, Junta de Andalucía, Sevilla, pp. 159–164
Richter-Boix, A., Llorente, G.A. & Montori, A. (2006) Breeding phenology of an amphibian community in a Mediterranean area. Amphibia-Reptilia, 27, 549–559.
https://doi.org/10.1163/156853806778877149Richter-Boix, A., Llorente, G.A., Montori, A. & García, J. (2007) Tadpole diet selection varies with the ecological context in predictable ways. Basic and Applied Ecology, 8, 464–474.
https://doi.org/10.1016/j.baae.2006.09.009Roček, Z. (2005) Late Miocene Amphibia from Rudabánya. Palaeontographia Italica, 90, 11–29.
Roček, Z. & Rage, J.C. (2000) Tertiary Anura of Europe, Asia, Africa, Asia, North America, and Australia. In: Heatwole, H., Carroll, R.L. (Eds.), Amphibian Biology. Surrey Beatty, Chipping Norton, pp. 1332–1387.
Roelants, K., Gower, D.J., Wilkinson, M., Loader, S.P., Biju, S.D., Guillaume, K., Moriau, L. & Bossuyt, F. (2007) Global patterns of diversification in the history of modern amphibians. Proceedings of the National Academy of Sciences of the U.S.A., 104, 887–892.
Salvador, A. & García-París, M. (2001) Anfibios Españoles. Identificación, Historia natural y Distribución. Canseco Editores, Talavera de la Reina, 269 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.
Salvidio, S. & Bologna, M.A. (2007) Pelodytes punctatus. In: Lanza, B., Andreone, F., Bologna, M.A., Corti, C. & Razzetti, E. (Eds.), Fauna d’Italia. Vol. XLII. Amphibia. Calderini de Il Sole, Bologna, pp. 362–372.
Salzburger, W., Ewing, G.B. & Von Haeseler, A. (2011) The performance of phylogenetic algorithms in estimating haplotype genealogies with migration. Molecular Ecology, 20, 1952–1963.
https://doi.org/10.1111/j.1365-294X.2011.05066.xSambrook, J., Fritsch, E.F. & Maniatis, T. (1989) Molecular Cloning: A Laboratory Manual, 2nd Edition. Cold Spring Harbor Press, New York, 1626 pp.
Sánchez-Herráiz, M.J. (2004) Análisis de la diferenciación genética, morfológica y ecológica asociadas a la especiación en el género Pelodytes (Anura, Pelodytidae). Tesis doctoral, Universidad Autónoma de Madrid, Madrid, 374 pp.
Sánchez-Herráiz, M.J., Barbadillo, L., Machordom, A. & Sanchiz, B. (2000) A new species of pelodytid frog from the Iberian Peninsula. Herpetologica, 56, 105–118.
Sanchiz, B. (1978) Nuevos restos fósiles de la familia Pelodytidae (Amphibia, Anura). Estudios Geológicos, 34, 9–27.
Sanchiz, B. (1998a) Salientia. Handbuch der Paläoherpetologie Pars 4. Dr. F. Pfeil, Munich, 275 pp.
Sanchiz, B. (1998b) Vertebrates from the early Miocene lignite deposits of the opencast mine Oberdorf (Western Styrian Basin, Austria): 2. Amphibia. Annalen des Natuhistorischen Museums Wien A, 99, 13–29.
Sanchiz, B., Tejedo, M. & Sánchez-Herráiz, M.J. (2002) Osteological differentiation among Iberian Pelodytes (Anura, Pelodytidae). Graellsia, 58, 35–68.
https://doi.org/10.3989/graellsia.2002.v58.i2.277Sequeira, F., Ferrand, N. & Harris, D. (2006) Assessing the phylogenetic signal of the nuclear beta-fibrinogen intron 7 in salamandrids (Amphibia: Salamandridae). Amphibia-Reptilia, 27, 409–418.
https://doi.org/10.1163/156853806778190114Shen, X.X., Liang, D. & Zhang, P. (2012) The development of three long universal nuclear protein-coding locus markers and their application to osteichthyan phylogenetics with nested PCR. PLoS ONE, 7, e39256.
https://doi.org/10.1371/journal.pone.0039256Silvestro, D. & Michalak, I. (2010) RAxML GUI: a graphical front-end for RAML. Available from: http://sourceforge.net/projects/raxmlgui/ (accessed 1 November 2015)
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.
Sindaco, R., Biggi, E., Boano, G. & Delmastro, G.B. (2002) Novitates herpetologicae pedemontanae. Rivista Piemontese di Storia Naturale, 23, 195–206.
Singhal, S. & Moritz, C. (2012) Testing hypotheses for genealogical discordance in a rainforest lizard. Molecular Ecology, 21, 5059–5072.
https://doi.org/10.1111/j.1365-294X.2012.05747.xSites, J.W. Jr. & Marshall, J.C. (2003) Delimiting species: a Renaissance issue in systematic biology. Trends in Ecology and Evolution, 18, 462–420.
https://doi.org/10.1016/S0169-5347(03)00184-8Stamatakis, A. (2006) RAxML-VI-HPC: maximum likelihood-based phylogenetic analyses with thousands of taxa and mixed models. Bioinformatics, 22, 2688–2690.
https://doi.org/10.1093/bioinformatics/btl446StatSoft Inc. (2007) STATISTICA (data analysis software system), version 7.1. Available from: http://www.statsoft.com (accessed 1 November 2015)
Steiner, H.M. (1968) Pelodytes caucasicus Boulenger, 1896 (Pelobatidae, Amphibia) in der Türkei. Annalen des Naturhistorischen Museums Wien, 72, 291–298.
Steinfartz, S., Veith, M. & Tautz, D. (2000) Mitochondrial sequence analysis of Salamandra taxa suggests old splits of major lineages and postglacial recolonizations of Central Europe from distinct source populations of Salamandra salamandra. Molecular Ecology, 9, 397–410.
https://doi.org/10.1046/j.1365-294x.2000.00870.xStephens, M., Smith, N.J. & Donnelly, P. (2001) A new statistical method for haplotype reconstruction from population data. The American Journal of Human Genetics, 68, 978–989.
https://doi.org/10.1086/319501Stephens, M. & Donnelly, P. (2003) A comparison of Bayesian methods for haplotype reconstruction from population genotype data. The American Journal of Human Genetics, 73, 1162–1169.
https://doi.org/10.1086/379378Stoate, C., Báldi, A., Beja, P., Boatman, N.D., Herzon, I., Van Doorn, A., de Snoo, G.R, Lakosy, L. & Ramwell, C. (2009) Ecological impacts of early 21st century agricultural change in Europe–a review. Journal of Environmental Management, 91, 22–46.
https://doi.org/10.1016/j.jenvman.2009.07.005Tamura, K., Stecher, G., Peterson, D., Filipski, A. & Kumar, S. (2013) MEGA6: molecular evolutionary genetics analysis version 6.0. Molecular Biology and Evolution, 30, 2725–2729.
https://doi.org/10.1093/molbev/mst197Tejedo, M. (1991) Effect of predation by two species of sympatric tadpoles on embryo survival in natterjack toads (Bufo calamita). Herpetologica, 47, 322–327.
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.
https://doi.org/10.1163/156853893X00309Truett, G.E., Heeger, P., Mynatt, R.L., Truett, A.A., Walker, J.A. & Warman, M.L. (2000) Preparation of PCR quality mouse genomic DNA with hot sodium hydroxide and Tris (HotSHOT). BioTechniques, 29, 52–54.
van de Vliet, M.S., Beebee, T.J. & Diekmann, O.E. (2012) Genetic evidence for a distinct Pelodytes lineage in southwest Portugal: implications for the use of pre-developed microsatellite markers. Conservation Genetics, 13, 605–611.
https://doi.org/10.1007/s10592-011-0299-5Veith, M., Steinfartz, S., Zardoya, R., Seitz, A. & Meyer, A. (1998) A molecular phylogeny of ‘true’ salamanders (family Salamandridae) and the evolution of terrestriality of reproductive modes. Journal of Zoological Systematics and Evolutionary Research, 36, 7–16.
https://doi.org/10.1111/j.1439-0469.1998.tb00774.xVeith, M., Fromhage, L., Kosuch, J. & Vences, M. (2006) Historical biogeography of Western Palaearctic pelobatid and pelodytid frogs: a molecular phylogenetic perspective. Contributions to Zoology, 75, 109–120.
Vences, M., Thomas, M., Bonett, R.M. & Vieites, D.R. (2005) Deciphering amphibian diversity through DNA barcoding: chances and challenges. Philosophical Transactions of the Royal Society of London B: Biological Sciences, 360, 1859–1868.
https://doi.org/10.1098/rstb.2005.1717Vences, M., Gehara, M., Köhler, J. & Glaw, F. (2012) Description of a new Malagasy treefrog (Boophis) occurring syntopically with its sister species, and a plea for studies on non-allopatric speciation in tropical amphibians. Amphibia-Reptilia, 33, 503–520.
https://doi.org/10.1163/15685381-00002856Vences, M., Glaw, F., Köhler, J. & Wollenberg, K.C. (2010) Molecular phylogeny, morphology and bioacoustics reveal five additional species of arboreal microhylids of the genus Anodonthyla from Madagascar. Contributions to Zoology, 79, 1–32.
Vences, M., Hauswaldt, J.S., Steinfartz, S., Rupp, O., Goesmann, A., Künzel, S., Orozco-terWengel, P., Vieites, D.R., Nieto-Roman, S., Haas, S., Laugsch, C., Gehara, M., Bruchmann, S., Pabijan, M., Ludewig, A.K., Rudert, D., Angelini, C., Borkin, L.J., Crochet, P.A., Crottini, A., Dubois, A., Ficetola, G.F., Galán, P., Geniez, P., Hachtel, M., Jovanovic, O., Litvinchuk, S.N., Lymberakis, P., Ohler, A. & Smirnov, N.A. (2013) Radically different phylogeographies and patterns of genetic variation in two European brown frogs, genus Rana. Molecular Phylogenetics and Evolution, 68, 657–670.
https://doi.org/10.1016/j.ympev.2013.04.014Vento, D. & Pérez, C. (2011) Datos sobre la herpetofauna del término municipal de Biar (Alicante, este de España). Boletín de la Asociación Herpetológica Española, 22, 130–134.
Vieites, D.R., Wollenberg, K.C., Andreone, F., Köhler, J., Glaw, F. & Vences, M. (2009) Vast underestimation of Madagascar's biodiversity evidenced by an integrative amphibian inventory. Proceedings of the National Academy of Sciences of the U.S.A., 106, 8267–8272.
https://doi.org/10.1073/pnas.0810821106Wake, D.B. (1997) Incipient species formation in salamanders of the Ensatina complex. Proceedings of the National Academy of Sciences of the U.S.A., 94, 7761–7767.
https://doi.org/10.1073/pnas.94.15.7761Watterson, G.A. (1975), On the number of segregating sites in genetical models without recombination. Theoretical Population Biology, 7, 256–276.
https://doi:10.1016/0040-5809(75)90020-9, PMID 1145509
Weisrock, D.W., Rasoloarison, R.M., Fiorentino, I., Ralison, J.M., Goodman, S.M., Kappeler, P.M. & Yoder, A.D. (2010) Delimiting species without nuclear monophyly in Madagascar's mouse lemurs. PLoS ONE, 5, e9883.
https://doi.org/10.1371/journal.pone.0009883Wielstra, B. & Arntzen, J.W. (2012) Postglacial species displacement in Triturus newts deduced from asymmetrically introgressed mitochondrial DNA and ecological niche models. BMC Evolutionary Biology, 12, 161.
https://doi.org/10.1186/1471-2148-12-161Zaldívar, C. (2004) Los anfibios de La Rioja. Páginas de Información Ambiental Gobierno de la Rioja, 16, 24–28.
Zamudio, K.R. & Savage, W.K. (2003) Historical isolation, range expansion, and secondary contact of two highly divergent mitochondrial lineages in spotted salamanders (Ambystoma maculatum). Evolution, 57, 1631–1652.
https://doi.org/10.1111/j.0014-3820.2003.tb00370.xZhang, H., Yan, J., Zhang, G. & Zhou, K. (2008) Phylogeography and demographic history of Chinese black-spotted frog populations (Pelophylax nigromaculata): evidence for independent refugia expansion and secondary contact. BMC Evolutionary Biology, 8, e21.
https://doi.org/10.1186/1471-2148-8-21Zieliński, P., Nadachowska-Brzyska, K., Wielstra, B., Szkotak, R., Covaciu-Marcov, S.D., Cogǎlniceanu, D. & Babik, W. (2013) No evidence for nuclear introgression despite complete mtDNA replacement in the Carpathian newt (Lissotriton montandoni). Molecular Ecology, 22, 1884–1903.
https://doi.org/10.1111/mec.12225