[en] Palearctic water frogs (genus Pelophylax) are an outstanding model in ecology and evolution, being widespread, speciose, either threatened or threatening to other species through biological invasions, and capable of siring hybrid offspring that escape the rules of sexual eproduction. Despite half a century of genetic research and hundreds of publications, the diversity, systematics and biogeography of Pelophylax still remain highly confusing, in no small part due to a lack of correspondence between studies. To provide a comprehensive overview, we gathered >13,000 sequences of barcoding genes from >1700 native and introduced localities and built multigene mitochondrial (~17 kb) and nuclear (~10 kb) phylogenies. We mapped all currently recognized taxa and their phylogeographic lineages (>40) to get a grasp on taxonomic issues, cyto-nuclear discordances, the genetic makeup of hybridogenetic hybrids, and the origins of introduced populations. Competing hypotheses for the molecular calibration were evaluated through plausibility tests, implementing a new approach relying on predictions from the anuran speciation continuum. Based on our timetree, we propose a new biogeographic paradigm for the Palearctic since the Paleogene, notably by attributing a prominent role to the dynamics of the Paratethys, a vast paleo-sea
that extended over most of Europe. Furthermore, our results show that distinct marsh frog
lineages from Eastern Europe, the Balkans, the Near East, and Central Asia (P. ridibundus ssp.) are naturally capable of inducing hybridogenesis with pool frogs (P. lessonae). We identified 14 alien lineages (mostly of P. ridibundus) over ~20 areas of invasions, especially in Western Europe, with genetic signatures disproportionally pointing to the Balkans and Anatolia as the regions of origins, in line with exporting records of the frog leg industry and the stocks of pet sellers. Pelophylax thus emerges as one of the most invasive amphibians worldwide, and deserves much higher conservation
Research Center/Unit :
FOCUS - Freshwater and OCeanic science Unit of reSearch - ULiège
Dufresnes, Christophe; Laboratory of Amphibian Systematics and Evolutionary Research (LASER), College of Biology and the Environment, Nanjing Forestry University, Nanjing, People's Republic of China ; Institut de Systématique, Muséum national d'Histoire naturelle, Evolution ; CNRS, Sorbonne Université, EPHE, Université des Antilles, Paris, France
Monod-Broca, Benjamin; UMR 5023 LEHNA, Université Claude Bernard Lyon 1, CNRS, ENTPE, Villeurbanne, France
Bellati, Adriana; Department of Ecological and Biological Sciences, University of Tuscia, Viterbo, Italy
Canestrelli, Daniele; Department of Ecological and Biological Sciences, University of Tuscia, Viterbo, Italy
Ambu, Johanna; Laboratory of Amphibian Systematics and Evolutionary Research (LASER), College of Biology and the Environment, Nanjing Forestry University, Nanjing, People's Republic of China
Wielstra, Ben; Institute of Biology Leiden, Leiden University, Leiden, The Netherlands ; Naturalis Biodiversity Center, Leiden, The Netherlands
Dubey, Sylvain; Department of Ecology and Evolution, University of Lausanne, Lausanne, Switzerland
Crochet, Pierre-André; CEFE, CNRS, Univ Montpellier, EPHE, Montpellier, France
Denoël, Mathieu ✱; Université de Liège - ULiège > Freshwater and OCeanic science Unit of reSearch (FOCUS) ; Université de Liège - ULiège > Département de Biologie, Ecologie et Evolution > Laboratoire d'Écologie et de Conservation des Amphibiens (LECA)
Jablonski, Daniel ✱; Department of Zoology, Comenius University in Bratislava, Bratislava, Slovakia
✱ These authors have contributed equally to this work.
Language :
English
Title :
Piecing the barcoding puzzle of Palearctic water frogs (Pelophylax) sheds light on amphibian biogeography and global invasions
Achalhi, M., Münch, P., Cornée, J.-J., Azdimousa, A., Melinte-Dobrinescu, M., Quillévéré, F., Drinia, H., Fauquette, S., Jiménez-Moreno, G., Merzeraud, G., Ben Moussa, A., El Kharim, Y., & Feddi, N. (2016). The late Miocene Mediterranean-Atlantic connections through the North Rifian Corridor: New insights from the Boudinar and Arbaa Taourirt basins (northeastern Rif, Morocco). Palaeogeography, Palaeoclimatology, Palaeoecology, 459, 131–152.
Akhmet'ev, M. A. (2011). Problems of Paleogene stratigraphy and paleogeography in the middle latitudes of Eurasia. Russian Geology and Geophysics, 52, 1075–1091.
Akın, Ç., Bilgin, C. C., Beerli, P., Westaway, R., Ohst, T., Litvinchuk, S. N., Uzzell, T., Bilgin, M., Hotz, H., Guex, G.-D., & Plötner, J. (2010). Phylogeographic patterns of genetic diversity in eastern Mediterranean water frogs were determined by geological processes and climate change in the Late Cenozoic. Journal of Biogeography, 37, 2111–2124.
Altaba, C. R. (1997). Phylogeny and biogeography of midwife toads (Alytes, Discoglossidae): A reappraisal. Contributions to Zoology, 66, 257–262.
Altaba, C. R. (1998). Testing vicariance: Melanopsid snails and Neogene tectonics in the Western Mediterranean. Journal of Biogeography, 25, 541–551.
Altherr, S., Auliya, M., & Nithart, C. (2022). Deadly dish – Role and responsibility of the European Union in the international frogs' legs trade. Report by Pro Wildlife & Robin des Bois. Munich, Germany, 28 pp.
Alves, R. R., Vieira, W. L., Santana, G. G., Vieira, K. S., & Montenegro, P. F. (2013). Herpetofauna used in traditional folk medicine: Conservation implications. In R. R. N. Alves & I. L. Rosa (Eds.), Animals in traditional folk medicine: Implications for conservation (pp. 109–133). Springer-Verlag Berlin Heidelberg.
Ambu, J., Martínez-Solano, Í., Suchan, T., Hernandez, A., Wielstra, B., Crochet, P.-A., & Dufresnes, C. (2023). Genomic phylogeography illuminates deep cyto-nuclear discordances in midwife toads (Alytes). Molecular Phylogenetics and Evolution, 183, 107783.
Anadón, P., Cabrera, L., & Roca, E. (1989). Contexto estructural y paleogeográfico de los sistemas lacustres cenozoicos de España. Acta Geologica Hispánica, 24, 167–184.
Andersen, B. G., & Borns, H. W. (1994). The ice age world. Scandinavia University Press.
Arano, B., Llorente, G., Garcia-Paris, M., & Herrero, P. (1995). Species translocation menaces Iberian waterfrogs. Conservation Biology, 9, 196–198.
Arntzen, J. W., Beukema, W., Galis, F., & Ivanović, A. (2015). Vertebral number is highly evolvable in salamanders and newts (family Salamandridae) and variably associated with climatic parameters. Contributions to Zoology, 84, 85–113.
Atamuradov, K. I. (1994). Paleogeography of Turkmenistan. In V. Fet & K. I. Atamuradov (Eds.), Biogeography and ecology of Turkmenistan (pp. 49–64). Kluwer Academic Publishers.
Auliya, M., Altherr, S., Nithart, C., Hughes, A., & Bickford, D. (2023). Numerous uncertainties in the multifaceted global trade in frogs' legs with the EU as the major consumer. Nature Conservation, 51, 71–135.
Avise, J. C. (2009). Phylogeography: Retrospect and prospect. Journal of Biogeography, 36, 3–15.
Baláž, V., Vojar, J., Civiš, P., Šandera, M., & Rozínek, R. (2014). Chytridiomycosis risk among Central European amphibians based on surveillance data. Diseases of Aquatic Organisms, 112, 1–8.
Balletto, E., Giacoma, C., Palestrini, C., Rolando, A., Ara, M., Barberis, A., Salvidio, S., Mensi, B., & Cassulo, L. (1990). On some aspects of the biography of northern Macaronesia. In A. Azzaroli (Ed.), Biogeographical aspects of insularity (pp. 167–199). Academia Nazionale dei Lincei.
Barnes, G. L. (2003). Origins of the Japanese islands: The new “big picture”. Japan Review, 15, 3–50.
Bartol, M., Mikuž, V., & Horvat, A. (2014). Palaeontological evidence of communication between the Central Paratethys and the Mediterranean in the late Badenian/early Serravalian. Palaeogeography, Palaeoclimatology, Palaeoecology, 394, 144–157.
Beddek, M., Zenboudji-Beddek, S., Geniez, P., Fathalla, R., Sourouille, P., Arnal, V., Dellaoui, B., Koudache, F., Telailia, S., Peyre, O., & Crochet, P.-A. (2018). Comparative phylogeography of amphibians and reptiles in Algeria suggests common causes for the east-west phylogeographic breaks in the Maghreb. PLoS One, 13, e0201218.
Beerli, P., Hotz, H., & Uzzell, T. (1996). Geologically dated sea barriers calibrate a protein clock for Aegean water frogs. Evolution, 50, 1676–1687.
Bellati, A., Bassu, L., Nulchis, V., & Corti, C. (2019). Detection of alien Pelophylax species in Sardinia (western Mediterranean, Italy). BioInvasions Records, 8, 8–25.
Bellati, A., Chiocchio, A., Razzetti, E., Bisconti, R., & Canestrelli, D. (2023). A hotchpotch of water frogs in northern Italy. Biological Invasions, 25, 2737–2745.
Berger, L. (1967). Embryonal and larval development of F1 generation of green frogs' different combinations. Acta Zoologica Cracoviensia, 12, 123–160.
Bisconti, R., Martino, G., Chiocchio, A., Siclari, A., & Canestrelli, D. (2019). Balkan marsh frogs Pelophylax kurtmuelleri (Gayda, 1940) introduced in the Aspromonte National Park, southern Italy. Bioinvasion Records, 8, 26–33.
Blackburn, D. C., Nielsen, S. V., & Jdeidi, T. B. (2022). Atlas of the frogs of Libya. Herpetological Review, 53, 211–219.
Blair, C. (2023). Organellar DNA continues to provide a rich source of information in the genomics era. Molecular Ecology, 32, 2144–2150.
Böhme, M., & Ilg, A. (2003). fosFARbase. https://www.fosfarbase.org/
Booth-Rea, G., Ranero, C. R., & Grevemeyer, I. (2018). The Alboran volcanic-arc modulated the Messinian faunal exchange and salinity crisis. Scientific Reports, 8, 13015.
Bouckaert, R., Vaughan, T. G., Barido-Sottani, J., Duchêne, S., Fourment, M., Gavryushkina, A., Heled, J., Jones, G., Kühnert, D., De Maio, N., Matschiner, M., Mendes, F. K., Müller, N. F., Ogilvie, H. A., du Plessis, L., Popinga, A., Rambaut, A., Rasmussen, D., Siveroni, I., … Stadler, T. (2019). BEAST 2.5: An advanced software platform for Bayesian evolutionary analysis. PLoS Computational Biology, 15, e1006650.
Bouckaert, R. R., & Heled, J. (2014). DensiTree 2: Seeing trees through the forest. https://doi.org/10.1101/012401
Bougeois, L., Dupont-Nivet, G., de Rafélis, M., Tindall, J. C., Proust, J.-N., Reichart, G.-J., de Nooijer, L. J., Guo, Z., & Ormukov, C. (2018). Asian monsoons and aridification response to Paleogene sea retreat and Neogene westerly shielding indicated by seasonality in Paratethys oysters. Earth and Planetary Science Letters, 485, 99–110.
Briggs, J. C. (1995). Global biogeography. Elsevier Science.
Bruni, G., Mirabella, I., Domeneghetti, D., Fasola, M., & Bellati, A. (2020). Will there be a second extinction? Molecular identification of multiple alien water frogs (Pelophylax ridibundus sensu lato) in Tuscany, Central Italy, reveals genetic pollution within a unique hybridogenetic system. Herpetological Journal, 30, 147–158.
Bulakhova, N., Alfimov, A., & Berman, D. (2020). The eastern boundary of the geographic range of the Pallas' spadefoot Pelobates vespertinus (Anura, Amphibia) is limited by overwintering temperatures. Herpetozoa, 33, 171–175.
Bülbül, U., Matsui, M., Kutrup, B., & Eto, K. (2011). Taxonomic relationships among Turkish water frogs as revealed by phylogenetic analyses using mtDNA gene sequences. Zoological Science, 28, 930–936.
Busack, S. (1986). Biogeographic analysis of the herpetofauna separated by the formation of the Strait of Gibraltar. National Geographic Research, 2, 17–36.
Capula, M. (2006). Rana bergeri Günther, 1985/Rana klepton hispanica Bonaparte, 1839. In R. Sindaco, G. Doria, E. Razzetti, & F. Bernini (Eds.), Atlante degli Anfini e dei Rettili d'Italia (pp. 236–239). Societa Herpetologica Italiana.
Chapman, M. A., & Burke, J. M. (2007). Genetic divergence and hybrid speciation. Evolution, 61, 1773–1780.
Çiçek, K., Ayaz, D., Afsar, M., Bayrakcı, Y., Akın Pekşen, Ç., Cumhuriyet, O., Bayram İsmail, İ., Yenmiş, M., Üstündağ, E., Varol Tok, C., Bilgin, C. C., & Akçakaya, H. R. (2020). Unsustainable harvest of water frogs in southern Turkey for the European market. Oryx, 55, 364–372.
Cuevas, A., Patrelle, C., Ciavatti, F., Gendre, T., Sourrouille, P., Geniez, P., Doniol-Valcroze, P., & Crochet, P.-A. (2022). A new PCR-RFLP method for the identification of parental and hybridogenetic western European water frogs, including the Pelophylax perezi-grafi system. Salamandra, 58, 218–230.
De Busschere, C., Courant, J., Herrel, A., Rebelo, R., Rödder, D., Measy, G. J., & Backeljau, T. (2016). Unequal contribution of native South African phylogeographic lineages to the invasion of the African clawed frog, Xenopus laevis, in Europe. PeerJ, 4, e1659.
De Leeuw, A., Bukowski, K., Krijgsman, W., & Kuiper, K. F. (2010). Age of the Badenian salinity crisis; impact of Miocene climate variability on the circum-Mediterranean region. Geology, 38, 715–718.
Denoël, M., Duret, C., Lorrain-Soligon, L., Padilla, P., Pavis, J., Pille, F., Tendron, P., Ficetola, G. F., & Falaschi, M. (2022). High habitat invasibility unveils the invasiveness potential of water frogs. Biological Invasions, 24, 3447–3459.
Dermitzakis, M. D., & Papanikolaou, D. J. (1981). Paleogeography and geodynamics of the Aegean region during the Neogene. Annales Géologiques des Pays Helléniques, 4, 245–289.
Doležálková-Kaštánková, M., Mazepa, G., Jeffries, D. L., Perrin, N., Plötner, M., Plötner, J., Guex, G.-D., Mikulíček, P., Poustka, A. J., Grau, J., & Choleva, L. (2021). Capture and return of sexual genomes by hybridogenetic frogs provides clonal genome enrichment in a sexual species. Scientific Reports, 11, 1633.
Domeneghetti, D., Bruni, G., Fasola, M., & Bellati, A. (2013). Discovery of alien water frogs (gen. Pelophylax) in Umbria, with first report of P. shqipericus for Italy. Acta Herpetologica, 8, 171–176.
Doniol-Valcroze, P., Mazepa, G., Grimal, F., Sourrouille, P., Perrin, N., Litvinchuk, S. N., & Crochet, P.-A. (2021). Discovery of a Pelophylax saharicus (Anura, Ranidae) population in Southern France: A new potentially invasive species of water frogs in Europe. Amphibia-Reptilia, 42, 427–442.
Drillon, O., Dufresnes, G., Perrin, N., Crochet, P. A., & Dufresnes, C. (2019). Reaching the edge of the speciation continuum: Hybridization between three sympatric species of tree frogs (Hyla). Biological Journal of the Linnean Society, 126, 743–750.
Dubey, S., & Dufresnes, C. (2017). An extinct vertebrate preserved by its living hybridogenetic descendant. Scientific Reports, 7, 12768.
Dubey, S., Lavanchy, G., Thiébaud, J., & Dufresnes, C. (2019). Herps without borders: A new newt case and a review of transalpine alien introductions in Western Europe. Amphibia-Reptilia, 40, 13–27.
Dubey, S., Leuenberger, J., & Perrin, N. (2014). Multiple origins of invasive and ‘native’ water frogs (Pelophylax spp.) in Switzerland. Biological Journal of the Linnean Society, 112, 442–449.
Dubey, S., Maddalena, T., Bonny, L., Jeffries, D. L., & Dufresnes, C. (2019). Population genomics of an exceptional hybridogenetic system of Pelophylax water frogs. BMC Evolutionary Biology, 19, 164.
Dubois, A. (1983). A propos de cuisses de grenouilles. Alytes, 2, 69–111.
Duellman, W. E., & Trueb, L. (1994). Biology of amphibians. Johns Hopkins University Press.
Dufresnes, C. (2024). Data from: Piecing the barcoding puzzle of Palearctic water frogs (Pelophylax) sheds light on amphibian biogeography and global invasions. https://doi.org/10.5281/zenodo.10423702
Dufresnes, C., Ambu, J., Kumar Prasad, V., Borzée, A., & Litvinchuk, S. N. (2022). A phylogeographic framework for Zhangixalus gliding frogs, with insights on their plasticity of nesting behavior. Biological Journal of the Linnean Society, 135, 40–51.
Dufresnes, C., Brelsford, A., Jeffries, D. L., Mazepa, G., Suchan, T., Canestrelli, D., Nicieza, A., Fumagalli, L., Dubey, S., Martínez-Solano, I., Litvinchuk, S. N., Vences, M., Perrin, N., & Crochet, P.-A. (2021). Mass of genes rather than master genes underlie the genomic architecture of amphibian speciation. Proceedings of the National Academy of Sciences of the United States of America, 118, e2103963118.
Dufresnes, C., & Crochet, P.-A. (2022). Sex chromosomes as supergenes of speciation: Why amphibians defy the rules? Philosophical Transactions of the Royal Society, B: Biological Sciences, 377, 1856.
Dufresnes, C., Denoël, M., Di Santo, L., & Dubey, S. (2017). Multiple uprising invasions of Pelophylax water frog, potentially inducing a new hybridogenetic complex. Scientific Reports, 7, 6506.
Dufresnes, C., Di Santo, L., Leuenberger, J., Schuerch, J., Mazepa, G., Grandjean, N., Canestrelli, D., Perrin, N., & Dubey, S. (2017). Cryptic invasion of Italian pool frogs (Pelophylax bergeri) across Western Europe unraveled by multilocus phylogeography. Biological Invasions, 19, 1407–1420.
Dufresnes, C., & Dubey, S. (2020). Invasion genomics supports an old hybrid swarm of pool frogs in Western Europe. Biological Invasions, 22, 205–210.
Dufresnes, C., & Hernandez, A. (2023). Towards completing the crocodile newts puzzle with all-inclusive phylogeographic resources. Zoological Journal of the Linnean Society, 197, 620–640.
Dufresnes, C., Leuenberger, J., Amrhein, V., Bühler, C., Thiébaud, J., Bohnenstengel, T., & Dubey, S. (2018). Invasion genetics of marsh frogs (Pelophylax ridibundus sensu lato) in Switzerland. Biological Journal of the Linnean Society, 123, 402–410.
Dufresnes, C., & Litvinchuk, S. N. (2022). Diversity, distribution and molecular species delimitation in frogs and toads from the Eastern Palearctic. Zoological Journal of the Linnean Society, 195, 695–760.
Dufresnes, C., & Mazepa, G. (2020). Hybridogenesis in water frogs. eLS, 1, 718–726.
Dufresnes, C., Mazepa, G., Jablonski, D., Caliari Oliveira, R., Wenseleers, T., Shabanov, D. A., Auer, M., Ernst, R., Ramírez-Chaves, H. E., Mulder, K. P., Simonov, E., Tiutenko, A., Kryvokhyzha, D., Wennekes, P. L., Zinenko, O. I., Korshunov, O. V., Al-Johany, A. M., Peregontsev, E. A., Betto-Colliard, C., … Litvinchuk, S. N. (2019). Fifteen shades of green: The evolution of Bufotes toads revisited. Molecular Phylogenetics and Evolution, 141, 106615.
Dufresnes, C., Poyarkov, N. A., & Jablonski, D. (2023). Acknowledging more biodiversity without more species. Proceedings of the National Academy of Sciences of the United States of America, 120, e2302424120.
Dufresnes, C., Strachinis, I., Suriadna, N. N., Mykytynets, G., Cogalniceanu, D., Székely, P., Vukov, T., Arntzen, J. W., Wielstra, B., Lymberakis, P., Geffen, E., Gafny, S., Kumlutas, Y., Ilgaz, C., Candan, K., Mizsei, E., Szabolcs, M., Kolenda, K., Smirnov, N. A., … Denoël, M. (2019). Phylogeography of a cryptic speciation continuum in Eurasian spadefoot toads (Pelobates). Molecular Ecology, 28, 3257–3270.
Ehl, S., Vences, M., & Veith, M. (2019). Reconstructing evolution at the community level: A case study on Mediterranean amphibians. Molecular Phylogenetics and Evolution, 134, 211–225.
Fromhage, L., Vences, M., & Veith, M. (2004). Testing alternative vicariance scenarios in Western Mediterranean discoglossid frogs. Molecular Phylogenetics and Evolution, 31, 308–322.
Frost, D. (2023). Amphibian species of the world 6.1, an online reference. American Museum of Natural History. https://amphibiansoftheworld.amnh.org/
Furman, B. L. S., Bewick, A. J., Harrison, T. L., Greenbaum, E., Gvoždík, V., Kusamba, C., & Evans, B. J. (2015). Pan-African phylogeography of a model organism, the African clawed frog ‘Xenopus laevis’. Molecular Ecology, 24, 909–925.
Garcia-Porta, J., Litvinchuk, S. N., Crochet, P. A., Romano, A., Geniez, P. H., Lo-Valvo, M., Lymberakis, P., & Carranza, S. (2012). Molecular phylogentics and historical biogeography of the west-palearctic common toads (Bufo bufo species complex). Molecular Phylogenetics and Evolution, 63, 113–130.
Garrick, R. C., Banusiewicz, J. D., Burgess, S., Hyseni, C., & Symula, R. E. (2019). Extending phylogeography to account for lineage fusion. Journal of Biogeography, 46, 268–278.
Gervais, M. (1836). Reptiles des Iles Canaries. In P. Webb & S. Berthelot (Eds.), Histoire Naturelle des Iles Canaries (pp. 15–24). Tome Deuxième, Deuxième Partie.
Gibert, L., Scott, G. R., Montoya, P., Ruiz-Sánchez, F. J., Morales, J., Luque, L., Abella, J., & Lería, M. (2013). Evidence for an African-Iberian mammal dispersal during the pre-evaporitic Messinian. Geology, 41, 691–694.
Göğüş, O., Pysklywec, R. N., Şengör, A. M. C., & Gün, E. (2017). Drip tectonics and the enigmatic uplift of the Central Anatolian Plateau. Nature Communications, 8, 1538.
Gouy, M., Tannier, E., Comte, N., & Parsons, D. P. (2021). Seaview v.5: A multiplatform software for multiple sequence alignment, molecular phylogenetic analyses, and tree reconciliation. Molecular Biology and Evolution, 2231, 241–260.
Graf, J.-D., & Polls, P. M. (1989). Evolutionary genetics of the Rana esculenta complex. In R. M. Dawley & J. P. Bogart (Eds.), Evolution and ecology of unisexual vertebrates. New York State Museum Bulletin 466 (pp. 289–302). New York State Museum.
Guerra-Merchán, A., & Serrano, F. (1993). Tectosedimentary setting and chronostratigraphy of the Neogene reefs in the Almazora Corridor (Betic Cordillera, Spain). Geobios, 26, 57–67.
Guindon, S., Dufayard, J.-F., Lefort, V., Anisimova, M., Hordijk, W., & Gascuel, O. (2010). New algorithms and methods to estimate maximum-likelihood phylogenies: Assessing the performance of PhyML 3.0. Systematic Biology, 59, 307–321.
Gvoždík, V., Nečas, T., Jablonski, D., Lemmon, E. M., Lemmon, A. R., Jandzik, D., & Moravec, J. (2023). Phylogenomics of Anguis and Pseudopus (Squamata, Anguidae) indicates Balkan-Apennine mitochondrial capture associated with the Messinian event. Molecular Phylogenetics and Evolution, 180, 107674.
He, J., Gao, Z., Su, Y., Lin, S., & Jiang, H. (2019). Geographical and temporal origins of terrestrial vertebrates endemic to Taiwan. Journal of Biogeography, 45, 2458–2470.
Heled, J., & Drummond, A. J. (2010). Bayesian inference of species trees from multilocus data. Molecular Biology and Evolution, 27, 570–580.
Herczeg, D., Vörös, J., Christiansen, D. G., Benovics, M., & Mikulíček, P. (2016). Taxonomic composition and ploidy level among European water frogs (Anura: Ranidae: Pelophylax) in eastern Hungary. Journal of Zoological Systematics and Evolutionary Research, 55, 129–137.
Hipsley, C. A., & Müller, J. (2014). Beyond fossil calibrations: Realities of molecular clock practices in evolutionary biology. Frontiers in Genetics, 5, 138.
Hoffmann, A., Plötner, J., Pruvost, N. B. M., Christiansen, D. G., Röthlisberger, S., Choleva, L., Mikulíček, P., Cogălniceanu, D., Sas-Kovács, I., Shabanov, D., Morozov-Leonov, S., & Reyer, H.-U. (2015). Genetic diversity and distribution patterns of diploid and polyploid hybrid water frog populations (Pelophylax esculentus complex) across Europe. Molecular Ecology, 24, 4371–4391.
Höglund, J., Bolender, L., Cortazar-Chinarro, M., Meurling, S., Laurila, A., Hermaniuk, A., & Dufresnes, C. (2022). Low neutral and immunogenetic diversity in northern fringe populations of the green toad Bufotes viridis: Implications for conservation. Conservation Genetics, 23, 139–149.
Holsbeek, G., & Jooris, R. (2010). Potential impact of genome exclusion by alien species in the hybridogenetic water frogs (Pelophylax esculentus complex). Biological Invasions, 12, 1–13.
Holsbeek, G., Mergeay, J., Hotz, H., Plötner, J., Volckaert, F. A. M., & De Meester, L. (2008). A cryptic invasion within an invasion and widespread introgression in the European water frog complex: Consequences of uncontrolled commercial trade and weak international legislation. Molecular Ecology, 17, 5023–5035.
Holsbeek, G., Mergeay, J., Volckaert, F. A. M., & De Meester, L. (2010). Genetic detection of multiple exotic water frog species in Belgium illustrates the need for monitoring and immediate action. Biological Invasions, 12, 1459–1463.
Hotz, H., Beerli, P., Uzzell, T., Guex, G.-D., Pruvost, N. B. M., Schreiber, R., & Plötner, J. (2013). Balancing a cline by influx of migrants: A genetic transition in water frogs of Eastern Greece. Journal of Heredity, 104, 57–71.
Hotz, H., Mancino, G., Bucci-Innocenti, S., Ragghianti, M., Berger, L., & Uzzell, T. (1985). Rana ridibunda varies geographically in inducing clonal gametogenesis in interspecies hybrids. Journal of Experimental Zoology, 236, 199–210.
Hotz, H., & Uzzell, T. (1983). Interspecific hybrids of Rana ridibunda without germ line exclusion of a parental genome. Experientia, 39, 538–540.
Huson, D. H., & Bryant, D. (2006). Application of phylogenetic networks in evolutionary studies. Molecular Biology and Evolution, 23, 254–267.
Igawa, T., Kurabayashi, A., Nishioka, M., & Sumida, M. (2006). Molecular phylogenetic relationship of toads distributed in the Far East and Europe inferred from the nucleotide sequences of mitochondrial DNA genes. Molecular Phylogenetics and Evolution, 38, 250–260.
Isozaki, Y., Maruyama, S., Nakama, T., Yamamoto, S., & Yanai, S. (2011). Growth and shrinkage of an active continental margin: Updated geotectonic history of the Japanese islands. Journal of Geography, 120, 65–99.
IUCN SSC Amphibian Specialist Group. (2020). Pelophylax plancyi. The IUCN Red List of Threatened Species 2020. e.T58697A63867236.
Jelić, M., Franjević, D., Đikić, D., Korlević, P., Vucić, M., Jelić, D., Becking, T., Grandjean, F., & Klobučar, G. (2022). Hybrid swarm as a result of hybridization between two alien and two native water frog species (genus Pelophylax) in Central Croatia: Crna Mlaka fishpond acting as a species melting pot? Biological Invasions, 24, 3291–3304.
Kamata, H., & Kodama, K. (1999). Volcanic history and tectonics of the southwest Japan Arc. The Island Arc, 8, 393–403.
Kamath, P. L., Sepulveda, A. J., & Layhee, M. (2016). Genetic reconstruction of a bullfrog invasion to elucidate vectors of introduction and secondary spread. Ecology and Evolution, 6, 5221–5233.
Kapp, P., Pullen, A., Pelletier, J. D., Russell, J., Goodman, P., & Cai, F. (2015). From dust to dust: Quaternary wind erosion of the Mu Us Desert and Loess Plateau, China. Geology, 43, 835–838.
Kieren, S., Sparreboom, M., Hochkirch, A., & Veith, M. (2018). A biogeographic and ecological perspective to the evolution of reproductive behaviour in the family Salamandridae. Molecular Phylogenetics and Evolution, 121, 98–109.
Kok, P., Jooris, R., Percsy, C., & Lenglet, G. (2002). Dangers pour la faune indigène de l'introduction d' espèces animales à des fins ornementales: Rana bedriagae (Amphibia: Anura: Ranidae), un futur cas d'école? Bulletin de l'Institut Royal des Sciences naturelles de Belgique, 7(Suppl), 219–221.
Kolenda, K., Pietras-Lebioda, A., Hofman, S., Ogielska, M., & Pabijan, M. (2017). Preliminary genetic data suggest the occurrence of the Balkan water frog, Pelophylax kurtmuelleri, in southwestern Poland. Amphibia-Reptilia, 38, 187–196.
Komaki, S., Igawa, T., Lin, S.-M., Tojo, K., Min, M.-S., & Sumida, M. (2015). Robust molecular phylogeny and palaeodistribution modelling resolve a complex evolutionary history: Glacial cycling drove recurrent mtDNA introgression among Pelophylax frogs in East Asia. Journal of Biogeography, 42, 2159–2171.
Koster, S., Prins, N., Dufresnes, C., France, J., de Visser, M. C., Struijk, R. P. J. H., & Wielstra, B. (2023). The conservation paradox of an introduced population of a threatened species: Spadefoot toads in the coastal dunes of The Netherlands. Amphibia-Reptilia, 44, 11–18.
Kouwenhoven, T. J., & van der Zwaan, G. J. (2006). A reconstruction of late Miocene Mediterranean circulation patterns using benthic foraminifera. Palaeogeography Palaeoclimatology Palaeoecology, 238, 373–385.
Krijgsman, W., Hilgen, F. J., Raffi, I., Sierro, F. J., & Wilson, D. S. (1999). Chronology, causes and progression of the Messinian Salinity Crisis. Nature, 400, 652–655.
Kuijt, M., Oskam, L., den Boer, I., Dufresnes, C., France, J., Gilbert, M. J., de Visser, M. C., Struijk, R. P. J. H., & Wielstra, B. (2023). The introduction of three cryptic tree frog species in the Dutch coastal dunes challenges conservation paradigms. Amphibia-Reptilia, 44, 1–10.
Kumar, S., Suleski, M., Craig, J. M., Kasprowicz, A. E., Sanderford, M., Li, M., Stecher, G., & Hedges, S. B. (2022). TimeTree 5: An expanded resource for species divergence times. Molecular Biology and Evolution, 39, msac174.
Kuzmin, S. L. (1999). The amphibians of the Former Soviet Union. Pensoft Publisher.
Lavanchy, G., & Schwander, T. (2019). Hybridogenesis. Current Biology, 29, R9–R11.
Lee, B.-R., Yoo, D.-G., & Lee, G.-S. (2022). High-resolution sequence stratigraphy and evolution of the Jeju Strait shelf, Korea, since the Last Glacial Maximum. Marine and Petroleum Geology, 135, 105389.
Lefort, V., Longueville, J.-E., & Gascuel, O. (2017). SMS: Smart model selection in PhyML. Molecular Biology and Evolution, 34, 2422–2424.
Lemierre, A., Gendry, D., Poirier, M.-M., Gillet, V., & Vullo, R. (2023). The oldest articulated ranid from Europe: A Pelophylax specimen from the lowest Oligocene of Chartres-de-Bretagne (N.W. France). Journal of Vertebrate Paleontology, 42, e2191663.
Leroy, S., & Dupont, L. (1994). Development of vegetation and continental aridity in northwestern Africa during the Late Pliocene: The pollen record of ODP Site 658. Palaeogeography, Palaeoclimatology, Palaeoecology, 109, 295–2316.
Li, J.-T., Wang, J.-S., Nian, H.-H., Litvinchuk, S. N., Wang, J., Li, Y., Rao, D.-Q., & Klaus, S. (2015). Amphibians crossing the Bering Land Bridge: Evidence from holarctic treefrogs (Hyla, Hylidae, Anura). Molecular Phylogenetics and Evolution, 87, 80–90.
Litvinchuk, S. N., Ivanov, A. Y., Lukonina, S. A., & Ermakov, O. A. (2020). A record of alien Pelophylax species and widespread mitochondrial DNA transfer in Kaliningradskaya Oblast' (the Baltic coast, Russia). BioInvasions Records, 9, 599–617.
Liu, 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.
Lonergan, L., & White, N. (1997). Origin of the Betic-Rif mountain belt. Tectonics, 16, 504–522.
Lyapkov, S. M., Ermakov, O. A., & Titov, S. V. (2018). Distribution and origin of two forms of the marsh frog Pelophylax ridibundus complex (Anura, Ranidae) from Kamchatka based on mitochondrial and nuclear DNA data. Biology Bulletin, 45, 699–705.
Lymberakis, P., Poulakakis, N., Manthalou, G., Tsigenopoulos, C. S., Magoulas, A., & Mylonas, M. (2007). Mitochondrial phylogeography of Rana (Pelophylax) populations in the Eastern Mediterranean region. Molecular Phylogenetics and Evolution, 44, 115–125.
Ma, D., Liu, H.-T., Ji, Y.-J., Shi, C.-M., Yang, Y.-H., & Zhang, D.-X. (2015). Black-spotted pond frog (Pelophylax nigromaculatus) on the Chinese Loess Plateau represents a cryptic species: Evidence from molecular phylogeny and ecological niche modeling. Journal of Systematics and Evolution, 53, 339–350.
Ma, X., Wang, Z., Tian, B., & Sun, H. (2019). Phylogeographic analyses of the East Asian endemic genus Prinsepia and the role of the east Asian monsoon system in shaping a north-south divergence pattern in China. Frontiers in Genetics, 10, 128.
Macaluso, L., Bertini, A., Carnevale, G., Eronen, J. T., Martinetto, E., Saarinen, J., Villa, A., Capasso, F., & Delfino, M. (2023). A combined palaeomodelling approach reveals the role as selective refugia of the Mediterranean peninsulas. Palaeogeography, Palaeoclimatology, Palaeoecology, 625, 111699.
Malacuso, L., Mannion, P. D., Evans, S. E., Carnevale, G., Monti, S., Marchitelli, D., & Delfino, M. (2022). Biogeographic history of Palearctic caudates revealed by a critical appraisal of their fossil record quality and spatio-temporal distribution. Royal Society Open Science, 9, 220935.
Malkmus, R. (2004). Amphibians and reptiles of Portugal, Madeira and the Azores-Archipelago. Ganter Verlag K.G.
Martín, J. M., Puga-Bernabéu, Á., Aguirre, J., & Braga, J. C. (2014). Miocene Atlantic-Mediterranean seaways in the Betic Cordillera (Southern Spain). Revista de la Sociedad Geológica de España, 27, 175–186.
Measey, J., Wagener, C., Mohanty, N. P., Baxter-Gilbert, J., & Pienaar, E. F. (2020). The cost and complexity of assessing impact. NeoBiota, 62, 279–299.
Meijer, N., Dupont-Nivet, G., Abels, H. A., Kaya, M. Y., Licht, A., Xiao, M., Zhang, Y., Roperch, P., Poujol, M., Lai, Z., & Guo, Z. (2019). Central Asian moisture modulated by proto-Paratethys Sea incursions since the early Eocene. Earth and Planetary Science Letters, 510, 73–84.
Meulenkamp, J. E., & Sissingh, W. (2003). Tertiary palaeogeography and tectonostratigraphic evolution of the Northern and Southern Peri-Tethys platforms and the intermediate domains of the African–Eurasian convergent plate boundary zone. Palaeogeography, Palaeoclimatology, Palaeoecology, 196, 209–228.
Min, M.-S., Park, S. K., Che, J., Park, D. S., & Lee, H. (2008). Genetic diversity among local populations of the gold-spotted pond frog, Rana plancyi chosenica (Amphibia: Ranidae), assessed by mitochondrial cytochrome b gene and control region sequences. Korean Journal of Systematic Zoology, 24, 25–32.
Mittan-Moreau, C. S., Kelehear, C., Toledo, L. F., Bacon, J., Guayasamin, J. M., Snyder, A., & Zamudio, K. R. (2022). Cryptic lineages and standing genetic variation across independent cane toad introductions. Molecular Ecology, 31, 6440–6456.
Miura, I., Vershinin, V., Vershinina, S., Lebedinskii, A., Trofimov, A., Sitnikov, I., & Ito, M. (2021). Hybridogenesis in the water frogs from western Russian territory: Intrapopulation variation in genome elimination. Genes, 12, 244.
Nagai, Y., Doi, T., Ito, K., Yuasa, Y., Fujitani, T., Naito, J.-I., Ogata, M., & Miura, I. (2018). The distributions and boundary of two distinct, local forms of Japanese pond frog, Pelophylax porosus brevipodus, inferred from sequences of mitochondrial DNA. Frontiers in Genetics, 9, 79.
Neveu, A. (2004). La raniculture est-elle une alternative à la récolte? Etat actuel en France. Productions Animales, 17, 167–175.
Neveu, A. (2009). Suitability of European green frogs for intensive culture: Comparison between different phenotypes of the esculenta hybridogenetic complex. Aquaculture, 295, 30–37.
Nicolas, V., Matame, A., Crochet, P.-A., Geniez, P., & Ohler, A. (2015). Phylogeographic patterns in North African water frog Pelophylax saharicus (Anura: Ranidae). Journal of Zoological Systematics and Evolutionary Research, 53, 239–248.
Oosterbroek, P., & Arntzen, J. W. (1992). Area-cladograms of circum-Mediterranean taxa in relation to Mediterranean palaeogeography. Journal of Biogeography, 19, 3–20.
Pabijan, M., Crottini, A., Reckwell, D., Irisarri, I., Hauswaldt, J. S., & Vences, M. (2012). A multigene species tree for Western Mediterranean painted frogs (Discoglossus). Molecular Phylogenetics and Evolution, 64, 690–696.
Pabijan, M., Wandycz, A., Hofman, S., Węcek, K., Piwczyński, M., & Szymura, J. M. (2013). Complete mitochondrial genomes resolve phylogenetic relationships within Bombina (Anura: Bombinatoridae). Molecular Phylogenetics and Evolution, 69, 63–74.
Padilla, P., Herrel, A., & Denoël, M. (2023). May future climate change promote the invasion of the marsh frog? An integrative thermo-physiological study. Oecologia, 202, 227–238.
Pagano, A., Lodé, T., & Crochet, P.-A. (2001). New contact zone and assemblages among water frogs of Southern France. Journal of Zoological Systematics and Evolutionary Research, 39, 63–67.
Palcu, D. V., & Krijgsman, W. (2023). The dire straits of Paratethys: Gateways to the anoxic giant of Eurasia. Geological Society, London, Special Publications, 523, 111–139.
Palcu, D. V., Patina, I. S., Șandric, I., Lazarev, S., Vasiliev, I., Stoica, M., & Krijgsman, W. (2021). Late Miocene megalake regressions in Eurasia. Scientific Reports, 11, 11471.
Palcu, D. V., Popov, S. V., Golovina, L., Kuiper, K. F., Liu, S., & Krijgsman, W. (2019). The shutdown of an anoxic giant: Magnetostratigraphic dating of the end of the Maikop Sea. Gondwana Research, 67, 82–100.
Papežík, P., Kubala, M., Jablonski, D., Doležálková-Kaštánková, M., Choleva, L., Benovics, M., & Mikulíček, P. (2021). Morphological differentiation of endemic water frogs (Ranidae: Pelophylax) from the southwestern Balkans. Salamandra, 57, 105–123.
Papežík, P., Mikulíček, P., Benovics, M., Balogová, M., Choleva, L., Doležálková-Kaštánková, M., Lymberakis, P., Mizsei, E., Papežíková, S., Poulakakis, N., Saçdanaku, E., Szabolcs, M., Šanda, R., Uhrin, M., Vukić, J., & Jablonski, D. (2023). Comparative mitochondrial phylogeography of water frogs (Ranidae: Pelophylax spp.) from the southwestern Balkans. Vertebrate Zoology, 73, 525–544.
Papežík, P., Sciberras, A., Benovics, M., Sciberras, J., Deidun, A., & Mikulíček, P. (2024). Far from home: Tracing the non-native origin of water frogs (genus Pelophylax) in Malta by molecular markers. Biological Invasions. https://doi.org/10.1007/s10530-023-03228-8
Percsy, C., & Percsy, N. (2002). Evolution des populations indigènes et introduites de grenouilles “vertes” en Brabant wallon. Bulletin de l'Institut Royal des Sciences naturelles de Belgique, 72(Suppl), 213–217.
Pille, F., Pinto, L., & Denoël, M. (2021). Predation pressure of invasive marsh frogs: A threat to native amphibians? Diversity, 13, 595.
Pille, F., Pinto, L., & Denoël, M. (2023). Functional and temporal facets of predation by marsh frogs across the aquatic-terrestrial ecotone of ponds and implications in the context of biological invasions. Freshwater Biology, 68, 2184–2196.
Plaziat, J.-C. (1981). Late Cretaceous to Late Eocene palaeogeographic evolution of southwest Europe. Palaeogeography, Palaeoclimatology, Palaeoecology, 36, 263–320.
Plötner, J., Akın Pekşen, Ç., Baier, F., Uzzell, T., & Can, B. C. (2015). Genetic evidence for human-mediated introduction of Anatolian water frogs (Pelophylax cf. bedriagae) to Cyprus (Amphibia: Ranidae). Zoology in the Middle East, 61, 125–132.
Plötner, J., Baier, F., Akın, Ç., Mazepa, G., Schreiber, R., Beerli, P., Litvinchuk, S. N., Bilgin, C. C., Borkin, L., & Uzzell, T. (2012). Genetic data reveal that water frogs of Cyprus (genus Pelophylax) are an endemic species of Messinian origin. Zoosystematics and Evolution, 88, 261–283.
Plötner, J., Köhler, F., Uzzell, T., Beerli, P., Schreiber, R., Guex, G. D., & Hotz, H. (2009). Evolution of serum albumin intron-1 is shaped by a 5′ truncated non-long terminal repeat retrotransposon in Western Palearctic water frogs (Neobatrachia). Molecular Phylogenetics and Evolution, 53, 784–791.
Plötner, J., Uzzell, T., Beerli, P., Spolsky, C., Ohst, T., Litvinchuk, S. N., Guey, G.-D., Reyer, H.-U., & Hotz, H. (2008). Widespread unidirectional transfer of mitochondrial DNA: A case in western Palaearctic water frogs. Journal of Evolutionary Biology, 21, 668–681.
Popov, S. V., Shcherba, I. G., Ilyina, L. B., Nevesskaya, L. A., Paramonova, N. P., Khondkarian, S. O., & Magyar, I. (2006). Late Miocene to Pliocene palaeogeography of the Paratethys and its relation to the Mediterranean. Palaeogeography, Palaeoclimatology, Palaeoecology, 238, 91–106.
Pramuk, J. B., Robertson, T., Sites, J. W., Jr., & Noonan, B. P. (2008). Around the world in 10 million years: Biogeography of the nearly cosmopolitan true toads (Anura: Bufonidae). Global Ecology and Biogeography, 17, 72–83.
Quilodrán, C. S., Montoya-Burgos, J. I., & Currat, M. (2015). Modelling interspecific hybridization with genome exclusion to identify conservation actions: The case of native and invasive Pelophylax waterfrogs. Evolutionary Applications, 8, 199–210.
Rage, J.-C., & Roček, Z. (2003). Evolution of anuran assemblages in the Tertiary and Quaternary of Europe, in the context of palaeoclimate and palaeogeography. Amphibia-Reptilia, 24, 133–167.
Ramstein, G., Fluteau, F., Besse, J., & Joussaume, S. (1997). Effect of orogeny, plate motion and land-sea distribution on Eurasian climate change over the past 30 million years. Nature, 386, 788–795.
Reilly, S. B., Wogan, G. O. U., Stubbs, A. L., Arida, E., Iskandar, D. T., & McGuire, J. A. (2017). Toxic toad invasion of Wallacea: A biodiversity hotspot characterized by extraordinary endemism. Global Change Biology, 23, 5029–5031.
Renault, D., Manfrini, E., Leroy, B., Diagne, C., Ballesteros-Mejia, L., Angulo, E., & Courchamp, F. (2021). Biological invasions in France: Alarming costs and even more alarming knowledge gaps. NeoBiota, 67, 191–224.
Rivera, D., Prates, I., Firneno, T. J., Jr., Trefaut Rodrigues, M., Caldwell, J. P., & Fujita, M. K. (2022). Phylogenomics, introgression, and demographic history of South American true toads (Rhinella). Molecular Ecology, 31, 978–992.
Robbemont, J., van Veldhuijzen, S., Allain, S. J. R., Ambu, J., Boyle, R., Canestrelli, D., Cathasaigh, É. Ó., Cathrine, C., Chiocchio, A., Cogalniceanu, D., Cvijanović, M., Dufresnes, C., Ennis, C., Gandola, D. R., Jablonski, D., Julian, A., Kranželić, D., Lukanov, S., Martínez-Solano, I., … Wielstra, B. (2023). An extended mtDNA phylogeography for the alpine newt illuminates the provenance of introduced populations. Amphibia-Reptilia, 44, 347–361.
Roure, B., Baurain, D., & Philippe, H. (2013). Impact of missing data on phylogenies inferred from empirical phylogenomic data sets. Molecular Biology and Evolution, 30, 197–214.
Rubinoff, D., Cameron, S., & Will, K. (2006). A genomic perspective on the shortcomings of mitochondrial DNA for “barcoding” identification. Journal of Heredity, 97, 581–594.
Sagonas, K., Karameta, E., Kotsakiozi, P., & Poulakakis, N. (2020). Cross-species testing of nuclear markers in Pelophylax water frogs in Greece and examination of their power to detect genetic admixture. Amphibia-Reptilia, 41, 253–259.
Santucci, F., Nascetti, G., & Bullini, L. (1996). Hybrid zones between two genetically differentiated forms of the pond frog Rana lessonae in southern Italy. Journal of Evolutionary Biology, 9, 429–450.
Scotese, C. R. (2016). PALEOMAP PaleoAtlas for GPlates and the PaleoData plotter program. PALEOMAP project. http://www.earthbyte.org/paleomap-paleoatlas¬for¬gplates/
Sibuet, J.-C., & Hsu, S.-K. (2004). How was Taiwan created? Tectonophysics, 379, 159–181.
Silva-Rocha, I., Montes, E., Salvi, D., Sillero, N., Mateo, J. A., Ayllón, E., Pleguezuelos, J. M., & Carretero, M. A. (2018). Herpetological history of the Balearic Islands: When aliens conquered these islands and what to do next. In A. I. Queiroz & S. Pooley (Eds.), Histories of bioinvasions in the Mediterranean (pp. 105–131). Springer.
Sofianidou, T. S. (1996). Electrophoretic studies of hybrids of water frogs (Rana epeirotica, R. balcanica) in the Ionian zone of Greece. Israel Journal of Zoology, 42, 149–157.
Soto, I., Cuthbert, R. N., Kouba, A., Capinha, C., Turbelin, A., Hudgins, E. J., Diagne, C., Courchamp, F., & Haubrock, P. J. (2022). Global economic costs of herpetofauna invasions. Scientific Reports, 12, 10829.
Speybroeck, J., Beukema, W., Dufresnes, C., Fritz, U., Jablonski, D., Lymberakis, P., Martínez-Solano, I., Razzetti, E., Vamberger, M., Vences, M., Vörös, J., & Crochet, P.-A. (2020). Species list of the European herpetofauna – 2020 Update by the Taxonomic Committee of the Societas Europaea Herpetologica. Amphibia-Reptilia, 41, 139–189.
Spigonardi, M. P., Resendes, R., & Cunha, R. (2012). Preliminary morphometric data of Pelophylax perezi (Amphibia: Ranidae) in São Jorge Island (Azores). Relatórios e Comunicações do Departamento de Biologia, 40, 79–87.
Stephens, M., Smith, N., & Donnelly, P. (2001). A new statistical method for haplotype reconstruction from population data. American Journal of Human Genetics, 68, 978–989.
Sumida, M., Ogata, M., & Nishioka, M. (2000). Molecular phylogenetic relationship of pond frogs distributed in the Palearctic region inferred from DNA sequences of mitochondrial 12S ribosomal DNA and cytochrome b genes. Molecular Phylogenetics and Evolution, 16, 278–285.
Sun, J., Gong, Z., Tian, Z., Jia, Y., & Windley, B. (2015). Late Miocene stepwise aridification in the Asian interior and the interplay between tectonics and climate. Palaeogeography, Palaeoclimatology, Palaeoecology, 421, 48–59.
Sun, J., Zhang, Z., Cao, M., Windley, B. F., Tian, S., Sha, J., Abdulov, S., Gadoev, M., & Oimahmadov, I. (2020). Timing of seawater retreat from proto-Paratethys, sedimentary provenance, and tectonic rotations in the late Eocene-early Oligocene in the Tajik Basin, Central Asia. Palaeogeography, Palaeoclimatology, Palaeoecology, 545, 109657.
Sun, Y., Ma, L., Bloemendal, J., Clemens, S., Qiang, X., & An, Z. (2015). Miocene climate change on the Chinese Loess Plateau: Possible links to the growth of the northern Tibetan Plateau and global cooling. Geochemistry, Geophysics, Geosystems, 16, 2097–2108.
Svinin, A. O., Dedukh, D. V., Borkin, L. J., Ermakov, O. A., Ivanov, A. Y., Litvinchuk, J. S., Zamaletdinov, R. I., Mikhaylova, R. I., Trubyanov, A. B., Skorinov, D. V., Rosanov, Y. M., & Litvinchuk, S. N. (2021). Genetic structure, morphological variation, and gametogenic peculiarities in water frogs (Pelophylax) from northeastern European Russia. Journal of Zoological Systematics and Evolutionary Research, 59, 646–662.
Taberlet, P., Fumagalli, L., Wust-Saucy, A. G., & Cosson, J. F. (1998). Comparative phylogeography and postglacial colonization routes in Europe. Molecular Ecology, 7, 453–464.
Teng, L. S. (1990). Geotectonic evolution of late Cenozoic arc continent collision in Taiwan. Tectonophysics, 183, 57–76.
Toli, E.-A., Bounas, A., Christopoulos, A., Pafilis, P., & Sotiropoulos, K. (2023). Phylogenetic analysis of the critically endangered Karpathos water frog (Anura, Amphibia): Conservation insights from complete mitochondrial genome sequencing. Amphibia-Reptilia, 44, 277–287.
Tunner, H. G. (1974). Die klonale Struktur einer Wasserfroschpopulation. Zeitschrift fur Zoologische Systematik und Evolutionsforschung, 12, 309–314.
Ualiyeva, D., Ermakov, O. A., Litvinchuk, S. N., Guo, X., Ivanov, A. Y., Xu, R., Li, J., Xu, F., Arifulova, I. I., Kaptyonkina, A. G., Khromov, V. A., Krainyuk, V. N., Sarzhanov, F., & Dujsebayeva, T. N. (2022). Diversity, phylogenetic relationships and distribution of marsh frogs (the Pelophylax ridibundus complex) from Kazakhstan and northwest China. Diversity, 14, 869.
Velo-Antón, G., Chambers, E. A., Poyarkov, N. A., Jr., Canestrelli, D., Bisconti, R., Naumov, B., Fernández Benéitez, M. J., Borisenko, A., & Martínez-Solano, I. (2023). COI barcoding provides reliable species identification and pinpoints cryptic diversity in Western Palearctic amphibians. Amphibia-Reptilia, 44, 399–413.
Vences, M., Brown, J. L., Lathrop, A., Rosa, G. M., Cameron, A., Crottini, A., Dolch, R., Edmonds, D., Freeman, K. L. M., Glaw, F., Grismer, L. L., Litvinchuk, S., Milne, M. G., Moore, M., Solofo, J. F., Noël, J., Nguyen, T. Q., Ohler, A., Randriaantoandro, C., … Murphy, R. W. (2017). Tracing a toad invasion: Lack of mitochondrial DNA variation, haplotype origins, and potential distribution of introduced Duttaphrynus melanostictus in Madagascar. Amphibia-Reptilia, 38, 197–207.
Vences, M., Vieites, D. R., Glaw, F., Brinkmann, H., Kosuch, J., Veith, M., & Meyer, A. (2003). Multiple overseas dispersal in amphibians. Proceedings of the Royal Society B: Biological Sciences, 270, 2435–2442.
Vucić, M., Jelić, D., Klobučar, G. I. V., Prkljačić, B., & Jelić, M. (2018). Molecular identification of species and hybrids of water frogs (genus Pelophylax) from Lake Skadar, Southeast Adriatic drainages (Amphibia: Ranidae). Salamandra, 54, 147–157.
Wang, S., Fan, L., Liu, C., Li, J., Gao, X., Zhu, W., & Li, Y. (2017). The origin of invasion of an alien frog species in Tibet, China. Current Zoology, 63, 615–621.
Wang, S., Liu, C., Zhu, W., Gao, X., & Li, Y. (2016). Tracing the origin of the black-spotted frog, Pelophylax nigromaculatus, in the Xinjiang Uyghur autonomous region. Asian Herpetological Research, 7, 69–74.
Weijermars, R. (1988). Neogene tectonics in the Western Mediterranean may have caused the Messinian Salinity Crisis and an associated glacial event. Tectonophysics, 148, 211–219.
Wicherley, J., Doran, S., & Beebee, T. J. C. (2003). Tracing aliens: Identification of introduced water frogs in Britain by male advertisement call characteristics. Herpetological Journal, 13, 43–50.
Wielstra, B., Burke, T., Butlin, R. K., & Arntzen, J. W. (2017). A signature of dynamic biogeography: Enclaves indicate past species replacement. Proceedings of the Royal Society B: Biological Sciences, 284, 20172014.
Wielstra, B., McCartney-Melstad, E., Arntzen, J. W., Butlin, R. K., & Shaffer, H. B. (2019). Phylogenomics of the adaptive radiation of Triturus newts supports gradual ecological niche expansion towards an incrementally aquatic lifestyle. Molecular Phylogenetics and Evolution, 133, 120–127.
Wielstra, B., Salvi, D., & Canestrelli, D. (2021). Genetic divergence across glacial refugia despite interglacial gene flow in a crested newt. Evolutionary Biology, 48, 17–26.
Wiens, J. J., & Tiu, J. (2012). Highly incomplete taxa can rescue phylogenetic analyses from the negative impacts of limited taxon sampling. PLoS One, 7, e42925.
Wilson, R. C. L., Drury, S. A., & Chapman, J. L. (2000). The great ice age: climate change and life. Routledge The Open University.
Wogan, G. O. U., Stuart, B. L., Iskandar, D. T., & McGuire, J. A. (2016). Deep genetic structure and ecological divergence in a widespread human commensal toad. Biology Letters, 12, 20150807.
Yan, F., Lü, J., Zhang, B., Yuan, Z., Zhao, H., Huang, S., Wei, G., Mi, X., Zou, D., Xu, W., Chen, S., Wang, J., Xie, F., Wu, M., Xiao, H., Liang, Z., Jin, J., Wu, S., Xu, C., … Che, J. (2018). The Chinese giant salamander exemplifies the hidden extinction of cryptic species. Current Biology, 28, R590–R592.
Yuan, Z.-Y., Zhou, W.-W., Chen, X., Poyarkov, N. A., Jr., Chen, H.-M., Jang-Liaw, N.-H., Chou, W.-H., Matzke, N. J., Iizuka, K., Min, M.-S., Kuzmin, S. L., Zhang, Y.-P., Cannatella, D. C., Hillis, D. M., & Che, J. (2016). Spatiotemporal diversification of the true frogs (genus Rana): A historical framework for a widely studied group of model organisms. Systematic Biology, 65, 824–842.
Zeisset, I., & Beebee, T. J. C. (2003). Population genetics of a successful invader: The marsh frog Rana ridibunda in Britain. Molecular Ecology, 12, 639–646.
Zhang, R., Jiang, D., Zhang, Z., & Zhang, C. (2021). Effects of Tibetan Plateau growth, Paratethys Sea retreat and global cooling on the East Asian climate by the early Miocene. Geochemistry, Geophysics, Geosystems, 22, e2021GC009655.
Zhang, Z., Wang, H. J., Guo, Z. T., & Jiang, D. (2007). What triggers the transition of palaeoenvironmental patterns in China, the Tibetan Plateau uplift or the Paratethys Sea retreat? Palaeogeography, Palaeoclimatology, Palaeoecology, 245, 317–331.
Zhou, S.-B., Zhang, Q.-Y., Hu, Z.-Q., Xia, Z.-Y., Miao, Q., Guan, Q., Guan, P., & Shi, J.-S. (2023). The validity of Pelophylax chosenicus (Okada, 1931) and P. hubeiensis (Fei & Ye, 1982) (Amphibia, Ranidae). Herpetozoa, 36, 143–152.
Zieliński, P., Nadachowska-Brzyska, K., Wielstra, B., Szkotak, R., Covaciu-Marcov, S. D., Cogălniceanu, D., & Babik, W. (2013). No evidence for nuclear introgression despite complete mtDNA replacement in the Carpathian newt (Lissotriton montandoni). Molecular Ecology, 22, 1884–1903.
