the second half of the 20th century (Dubois, 1983). Accordingly, genetic analyses of the stocks bred by farms and sellers revealed a mix of Eastern Mediterranean lineages (Bellati et al., 2023; Holsbeek et al., 2008, 2010).

In China, the frog trade, both for consumption and traditional medicine (Alves et al., 2013) is also a potential vector of translocations. There are thousands of farms distributing live frogs to an even higher number of food sellers across the whole country, opening countless opportunities for escapes and invasions. Although the exploitation of *Hoplobatrachus* has been particularly highlighted (Alves et al., 2013), *Pelophylax* are also harvested (IUCN SSC Amphibian Specialist Group, 2020). It would thus not be surprising if, like in Europe, future prospections would find that water frog diversity and distribution patterns in Asia were also human-mediated. For instance, recent research on Giant salamanders showed that farming was the cause of multiple unsuspected introductions (Yan et al., 2018).

These observations unequivocally incriminate commercial imports as the primary source of introductions (see also Dubois, 1983). This is nicely illustrated by the spread of *P. ridibundus* in Switzerland during the second half of the 20th century, which started by independent clusters located near airports and breeding facilities (Dufresnes et al., 2018). In W-Europe, a few frog farms continue to develop, notably by producing their own breeds that are advertised as local delicacy. For instance, some companies use the race "Rivan 92", which was engineered through selective breeding so the frogs can be fed fish granulates rather than live insects (Neveu, 2009), and which initially descent from individuals caught in Britany (France) but are of uncertain taxonomic nature (Neveu, 2004).

Besides farming and trading activities, it is likely that a large number of introductions also originate for the purpose of ornamentation in garden ponds. Water frogs were largely sold as ornamental species by local sellers or provided with aquatic vegetation (Kok et al., 2002; Percsy & Percsy, 2002). Given the widespread popularity of stocking garden ponds with ornamental species, this may represent a cause of multiple invasions, often likely cryptic (P.-A.C., M.D., personal observation). In addition, water frogs are also popular educational and research models (alongside *Xenopus*), and escapes from university and schools are common, for example, in France and Switzerland (P.-A.C., C.D., personal observation).

Although the situation seems beyond repair in several countries, the issue of *Pelophylax* invasions remains largely ignored by conservation authorities—water frogs are typically overlooked in bioinvasion reports (e.g., Renault et al., 2021). We see several reasons for this lack of awareness. Exotic water frogs resemble native ones and have been present for decades in some areas (Dubois, 1983), hence they do not attract much attention compared to more spectacular species (e.g., *X. laevis* and *L. catesbeianus* in France and Italy). In addition, many introduced populations are protected by national decree because they belong to species that supposedly occur naturally in other parts of the same country (e.g., *P. ridibundus* in France). Such a "conservation paradox" (sensu Koster et al., 2023) does not only safeguards invasive species, it also prevents conservation biologists from studying invasions by making it more difficult to obtain research permits.

The unstable taxonomy of *Pelophylax* is also certainly not favorable to the implementation of long-term conservation policies, which rely on scientific (taxonomic) names. Taxonomic uncertainties even benefits frog exporting companies, which abusively use generic names (e.g., "*P. esculentus*") to escape the legislation (Altherr et al., 2022). Furthermore, several exotic lineages await a formal description more than 15 years after their discovery, being instead collectively designated as "*P. cf. bedriagae*" or "*P. cf. ridibundus*". At the same time, taxonomic accounts disagree on the validity and species/subspecies rank of many described taxa, depending on the applied concepts and criteria (e.g., *P. ridibundus/kurtmuelleri*; *P. bedriagae/cypriensis/caralitanus/cerigensis*; *P. plancyi/hubeiensis/choenicus*; see Dufresnes & Litvinchuk, 2022; Frost, 2023; Speybroeck et al., 2020). Solving the taxonomy of water frogs thus appears urgent if distinct lineages are to be independently treated in national and international legislation, which will be necessary to ban invasive lineages from being imported, to eradicate them in invaded areas (when feasible), and to protect them in their native range, where unsustainable harvest is causing significant population declines (Çiçek et al., 2020). The ambition of reaching a stable taxonomy in *Pelophylax* will require a tremendous amount of dedicated taxonomic work, ideally based on next-generation phylogeography (i.e., implementing genomic data), to describe unnamed lineages, clarify nomenclatural ambiguities, and delimit species under standardized criteria (e.g., Dufresnes et al., 2023). For the time being, we tentatively list species and subspecies based on previous taxonomic discussions and our updated phylogeographic overview, which should offer a basis for coming updates (Table S1). In addition, our mitochondrial haplotype database, which spans six barcoding genes, is designed so that it can be easily reused and complemented by future invasion genetic studies.

While most water frog populations with a foreign genetic signature have a human-mediated origin, our results also call for cautious interpretations in a few cases. First, rather than multiple translocations, postglacial expansions could have also naturally carried *kurtmuelleri* mtDNA and nuclear haplotypes over Central and Northern Europe (Kolenda et al., 2017; Litvinchuk et al., 2020), as seen in some anurans of Balkan origin (e.g., *Bufotes viridis*, Höglund et al., 2022). This scenario is possible given that *P. ridibundus ridibundus* and *P. ridibundus kurtmuelleri* seem to represent shallow refugial lineages of the same biological species (Papežík et al., 2023) that remain highly similar at the genomic scale (Dubey, Lavanchy, et al., 2019) and may thus hybridize without reproductive barriers.

Likewise, the presence of the Italian pool frog lineage *bergeri* A in Switzerland and France was attributed to ancient translocations on the basis of the large distributional gap between the closest native (N-Apennines; Capula, 2006) and putatively introduced records (near Lyon; Dufresnes, Di Santo, et al., 2017), which made natural colonization an unlikely hypothesis (Dubey et al., 2014; Dufresnes & Dubey, 2020; Dufresnes, Di Santo, et al., 2017). We showed here that the *bergeri* mitogroup extends in at least parts of this gap along the Ligurian and Eastern French Mediterranean coastline, although at this stage, it remains undetermined whether these populations are indigenous and if they belong to pool frogs, marsh frogs or their
the second half of the 20th century (Dubois, 1983). Accordingly, genetic analyses of the stocks bred by farms and sellers revealed a mix of Eastern Mediterranean lineages (Bellati et al., 2023; Holsbeek et al., 2008, 2010).

In China, the frog trade, both for consumption and traditional medicine (Alves et al., 2013) is also a potential vector of translocations. There are thousands of farms distributing live frogs to an even higher number of food sellers across the whole country, opening countless opportunities for escapes and invasions. Although the exploitation of *Hoplobatrachus* has been particularly highlighted (Alves et al., 2013), *Pelophylax* are also harvested (IUCN SSC Amphibian Specialist Group, 2020). It would thus not be surprising if, like in Europe, future prospections would find that water frog diversity and distribution patterns in Asia were also human-mediated. For instance, recent research on Giant salamanders showed that farming was the cause of multiple unsuspected introductions (Yan et al., 2018).

These observations unequivocally incriminate commercial imports as the primary source of introductions (see also Dubois, 1983). This is nicely illustrated by the spread of *P. ridibundus* in Switzerland during the second half of the 20th century, which started by independent clusters located near airports and breeding facilities (Dufresnes et al., 2018). In W-Europe, a few frog farms continue to develop, notably by producing their own breeds that are advertised as local delicacy. For instance, some companies use the race "Rivan 92", which was engineered through selective breeding so the frogs can be fed fish granulates rather than live insects (Neveu, 2009), and which initially descent from individuals caught in Britany (France) but are of uncertain taxonomic nature (Neveu, 2004).

Besides farming and trading activities, it is likely that a large number of introductions also originate for the purpose of ornamentation in garden ponds. Water frogs were largely sold as ornamental species by local sellers or provided with aquatic vegetation (Kok et al., 2002; Percsy & Percsy, 2002). Given the widespread popularity of stocking garden ponds with ornamental species, this may represent a cause of multiple invasions, often likely cryptic (P.-A.C., M.D., personal observation). In addition, water frogs are also popular educational and research models (alongside *Xenopus*), and escapes from university and schools are common, for example, in France and Switzerland (P.-A.C., C.D., personal observation).

Although the situation seems beyond repair in several countries, the issue of *Pelophylax* invasions remains largely ignored by conservation authorities—water frogs are typically overlooked in bioinvasion reports (e.g., Renault et al., 2021). We see several reasons for this lack of awareness. Exotic water frogs resemble native ones and have been present for decades in some areas (Dubois, 1983), hence they do not attract much attention compared to more spectacular species (e.g., *X. laevis* and *L. catesbeianus* in France and Italy). In addition, many introduced populations are protected by national decree because they belong to species that supposedly occur naturally in other parts of the same country (e.g., *P. ridibundus* in France). Such a "conservation paradox" (sensu Koster et al., 2023) does not only safeguards invasive species, it also prevents conservation biologists from studying invasions by making it more difficult to obtain research permits.

The unstable taxonomy of *Pelophylax* is also certainly not favorable to the implementation of long-term conservation policies, which rely on scientific (taxonomic) names. Taxonomic uncertainties even benefits frog exporting companies, which abusively use generic names (e.g., "*P. esculentus*") to escape the legislation (Altherr et al., 2022). Furthermore, several exotic lineages await a formal description more than 15 years after their discovery, being instead collectively designated as "*P. cf. bedriagae*" or "*P. cf. ridibundus*". At the same time, taxonomic accounts disagree on the validity and species/subspecies rank of many described taxa, depending on the applied concepts and criteria (e.g., *P. ridibundus/kurtmuelleri*; *P. bedriagae/cypriensis/caralitanus/cerigensis*; *P. plancyi/hubeiensis/chosenicus*; see Dufresnes & Litvinchuk, 2022; Frost, 2023; Speybroeck et al., 2020). Solving the taxonomy of water frogs thus appears urgent if distinct lineages are to be independently treated in national and international legislation, which will be necessary to ban invasive lineages from being imported, to eradicate them in invaded areas (when feasible), and to protect them in their native range, where unsustainable harvest is causing significant population declines (Çiçek et al., 2020). The ambition of reaching a stable taxonomy in *Pelophylax* will require a tremendous amount of dedicated taxonomic work, ideally based on next-generation phylogeography (i.e., implementing genomic data), to describe unnamed lineages, clarify nomenclatural ambiguities, and delimit species under standardized criteria (e.g., Dufresnes et al., 2023). For the time being, we tentatively list species and subspecies based on previous taxonomic discussions and our updated phylogeographic overview, which should offer a basis for coming updates (Table S1). In addition, our mitochondrial haplotype database, which spans six barcoding genes, is designed so that it can be easily reused and complemented by future invasion genetic studies.

While most water frog populations with a foreign genetic signature have a human-mediated origin, our results also call for cautious interpretations in a few cases. First, rather than multiple translocations, postglacial expansions could have also naturally carried *kurtmuelleri* mtDNA and nuclear haplotypes over Central and Northern Europe (Kolenda et al., 2017; Litvinchuk et al., 2020), as seen in some anurans of Balkan origin (e.g., *Bufotes viridis*, Höglund et al., 2022). This scenario is possible given that *P. ridibundus ridibundus* and *P. ridibundus kurtmuelleri* seem to represent shallow refugial lineages of the same biological species (Papežík et al., 2023) that remain highly similar at the genomic scale (Dubey, Lavanchy, et al., 2019) and may thus hybridize without reproductive barriers.

Likewise, the presence of the Italian pool frog lineage *bergeri* A in Switzerland and France was attributed to ancient translocations on the basis of the large distributional gap between the closest native (N-Apennines; Capula, 2006) and putatively introduced records (near Lyon; Dufresnes, Di Santo, et al., 2017), which made natural colonization an unlikely hypothesis (Dubey et al., 2014; Dufresnes & Dubey, 2020; Dufresnes, Di Santo, et al., 2017). We showed here that the *bergeri* mitogroup extends in at least parts of this gap along the Ligurian and Eastern French Mediterranean coastline, although at this stage, it remains undetermined whether these populations are indigenous and if they belong to pool frogs, marsh frogs or their
<figure><img src="image_27.png" /></figure>

hybridogenetic hybrids—mitochondrial transfer among all three is common (Plötner et al., 2008). From a biogeographic perspective, it is conceivable that *P. lessonae bergeri* briefly expanded out of NW-Italy after the last glacial maximum, after which it was replaced by other *Pelophylax* species (e.g., *P. perezi* and now *P. ridibundus*), given that this Italo-French route is one of Europe's post-glacial recolonization's paradigms (Taberlet et al., 1998).

## 4.6 | Conclusions

Despite well-known limitations (Rubinoff et al., 2006), single gene barcoding shall continue to provide essential information, even in the era of genomics (Blair, 2023). By combining decades of barcoding efforts in a meta-analysis framework, we revisit the diversification of Palearctic water frogs in space and time to appraise its evolution, diversity and biological invasions. This overview should motivate future surveys to illuminate the evolutionary relevance of the documented phylogroups, their systematics, their interactions in secondary contact zones, and hopefully contributes to a better consideration of their emerging invasions by stakeholders. Our study also reaches the limits of molecular analyses based on a few genes, and further investigations should rely on genomic data, starting by a phylogenomic species tree and multilocus analyses of hybridization and gene flow in hotspots of phylogeographic diversity for the genus, namely the Balkans, Anatolia, Central Asia, Japan and mainland China. As its complexity unfolds after over half a century of frenetic research (Berger, 1967), *Pelophylax* hybridogenesis still holds many fascinating secrets that are bound to challenge our perception of biological notions as fundamental as sexual reproduction and speciation.

## AUTHOR CONTRIBUTIONS

**Christophe Dufresnes:** Conceptualization; data curation; formal analysis; investigation; methodology; writing – original draft.

**Benjamin Monod-Broca:** Data curation; investigation. **Adriana Bellati:** Resources; writing – review and editing. **Daniele Canestrelli:** Resources; writing – review and editing. **Johanna Ambu:** Investigation.

**Ben Wielstra:** Conceptualization; writing – review and editing. **Sylvain Dubey:** Resources; writing – review and editing. **Pierre-André Crochet:** Resources; writing – review and editing. **Mathieu Denoël:** Conceptualization; resources; writing – review and editing.

**Daniel Jablonski:** Conceptualization; investigation; resources; writing – review and editing.

**Sylvain Dubey:** Resources; writing – review and editing. **Pierre-André Crochet:** Resources; writing – review and editing. **Mathieu Denoël:** Conceptualization; resources; writing – review and editing.

**Daniel Jablonski:** Conceptualization; investigation; resources; writing – review and editing.

**Jablonski:** Conceptualization; investigation; resources; writing – review and editing.

**Daniel Jablonski:** Conceptualization; investigation; resources; writing – review and editing.

## ACKNOWLEDGEMENTS

We are thankful to G. Mazepa and S. N. Litvinchuk for assistance with some of the *Pelophylax* barcoding literature, as well as P. Geniez for curating new samples. M.D. is a Research Director at F.R.S.– FNRS (Fonds de la Recherche Scientifique). This study was partly funded by a PDR FNRS grant (T.0070.19 to M.D.) and a RFIS-NSFC grant (3211101356 to C.D.).

## CONFLICT OF INTEREST STATEMENT

The authors have no conflict of interest to declare.

## DATA AVAILABILITY STATEMENT

All datasets and metadata have been deposited on the Zenodo platform (Dufresnes, 2024).

## ORCID

Christophe Dufresnes https://orcid.org/0000-0002-8497-8908

Benjamin Monod-Broca https://orcid.org/0000-0002-9133-5548

Adriana Bellati https://orcid.org/0000-0002-7259-7569

Daniele Canestrelli https://orcid.org/0000-0001-9351-4972

Johanna Ambu https://orcid.org/0000-0002-8072-7677

Ben Wielstra https://orcid.org/0000-0002-7112-5965

Sylvain Dubey https://orcid.org/0000-0002-0422-3960

Pierre-André Crochet https://orcid.org/0000-0003-4342-9432

Mathieu Denoël https://orcid.org/0000-0002-3586-8323

Daniel Jablonski https://orcid.org/0000-0002-5394-0114

## REFERENCES

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 paleo-geography in the middle latitudes of Eurasia. *Russian Geology and Geophysics*, 52, 1075–1091.

Akin, 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 pakeogeográfico 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.
<figure><img src="image_30.png" /></figure>

hybridogenetic hybrids—mitochondrial transfer among all three is common (Plötner et al., 2008). From a biogeographic perspective, it is conceivable that *P. lessonae bergeri* briefly expanded out of NW-Italy after the last glacial maximum, after which it was replaced by other *Pelophylax* species (e.g., *P. perezi* and now *P. ridibundus*), given that this Italo-French route is one of Europe's post-glacial recolonization's paradigms (Taberlet et al., 1998).

## 4.6 | Conclusions

Despite well-known limitations (Rubinoff et al., 2006), single gene barcoding shall continue to provide essential information, even in the era of genomics (Blair, 2023). By combining decades of barcoding efforts in a meta-analysis framework, we revisit the diversification of Palearctic water frogs in space and time to appraise its evolution, diversity and biological invasions. This overview should motivate future surveys to illuminate the evolutionary relevance of the documented phylogroups, their systematics, their interactions in secondary contact zones, and hopefully contributes to a better consideration of their emerging invasions by stakeholders. Our study also reaches the limits of molecular analyses based on a few genes, and further investigations should rely on genomic data, starting by a phylogenomic species tree and multilocus analyses of hybridization and gene flow in hotspots of phylogeographic diversity for the genus, namely the Balkans, Anatolia, Central Asia, Japan and mainland China. As its complexity unfolds after over half a century of frenetic research (Berger, 1967), *Pelophylax* hybridogenesis still holds many fascinating secrets that are bound to challenge our perception of biological notions as fundamental as sexual reproduction and speciation.

## AUTHOR CONTRIBUTIONS

**Christophe Dufresnes:** Conceptualization; data curation; formal analysis; investigation; methodology; writing – original draft.

**Benjamin Monod-Broca:** Data curation; investigation. **Adriana Bellati:** Resources; writing – review and editing. **Daniele Canestrelli:** Resources; writing – review and editing. **Johanna Ambu:** Investigation.

**Ben Wielstra:** Conceptualization; writing – review and editing. **Sylvain Dubey:** Resources; writing – review and editing. **Pierre-André Crochet:** Resources; writing – review and editing. **Mathieu Denoël:** Conceptualization; resources; writing – review and editing.

**Daniel Jablonski:** Conceptualization; investigation; resources; writing – review and editing.

**Sylvain Dubey:** Resources; writing – review and editing. **Pierre-André Crochet:** Resources; writing – review and editing. **Mathieu Denoël:** Conceptualization; resources; writing – review and editing.

**Daniel Jablonski:** Conceptualization; investigation; resources; writing – review and editing.

**Jablonski:** Conceptualization; investigation; resources; writing – review and editing.

**Daniel Jablonski:** Conceptualization; investigation; resources; writing – review and editing.

## ACKNOWLEDGEMENTS

We are thankful to G. Mazepa and S. N. Litvinchuk for assistance with some of the *Pelophylax* barcoding literature, as well as P. Geniez for curating new samples. M.D. is a Research Director at F.R.S.–FNRS (Fonds de la Recherche Scientifique). This study was partly funded by a PDR FNRS grant (T.0070.19 to M.D.) and a RFIS-NSFC grant (3211101356 to C.D.).

## CONFLICT OF INTEREST STATEMENT

The authors have no conflict of interest to declare.

## DATA AVAILABILITY STATEMENT

All datasets and metadata have been deposited on the Zenodo platform (Dufresnes, 2024).

## ORCID

Christophe Dufresnes https://orcid.org/0000-0002-8497-8908

Benjamin Monod-Broca https://orcid.org/0000-0002-9133-5548

Adriana Bellati https://orcid.org/0000-0002-7259-7569

Daniele Canestrelli https://orcid.org/0000-0001-9351-4972

Johanna Ambu https://orcid.org/0000-0002-8072-7677

Ben Wielstra https://orcid.org/0000-0002-7112-5965

Sylvain Dubey https://orcid.org/0000-0002-0422-3960

Pierre-André Crochet https://orcid.org/0000-0003-4342-9432

Mathieu Denoël https://orcid.org/0000-0002-3586-8323

Daniel Jablonski https://orcid.org/0000-0002-5394-0114

## REFERENCES

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 paleo-geography in the middle latitudes of Eurasia. *Russian Geology and Geophysics*, 52, 1075–1091.

Akin, 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.
<figure><img src="image_28.png" /></figure>

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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). *Biolnvasions 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.

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<figure><img src="image_31.png" /></figure>

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## SUPPORTING INFORMATION

Additional supporting information can be found online in the Supporting Information section at the end of this article.

**How to cite this article:** Dufresnes, C., Monod-Broca, B., Bellati, A., Canestrelli, D., Ambu, J., Wielstra, B., Dubey, S., Crochet, P.-A., Denoël, M., & Jablonski, D. (2024). Piecing the barcoding puzzle of Palearctic water frogs (*Pelophylax*) sheds light on amphibian biogeography and global invasions. *Global Change Biology*, **30**, e17180. https://doi.org/10.1111/gcb.17180
<figure><img src="image_33.png" /></figure>

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.

## SUPPORTING INFORMATION

Additional supporting information can be found online in the Supporting Information section at the end of this article.

**How to cite this article:** Dufresnes, C., Monod-Broca, B., Bellati, A., Canestrelli, D., Ambu, J., Wielstra, B., Dubey, S., Crochet, P.-A., Denoël, M., & Jablonski, D. (2024). Piecing the barcoding puzzle of Palearctic water frogs (*Pelophylax*) sheds light on amphibian biogeography and global invasions. *Global Change Biology*, **30**, e17180. https://doi.org/10.1111/gcb.17180