Bombina variegata (Linnaeus, 1758)

Class: Amphibia > Order: Anura > Family: Bombinatoridae > Genus: Bombina > Species: Bombina variegata

Rana variegata Linnaeus, 1758, Syst. Nat., Ed. 10, 1: 211. Type(s): Not designated, presumed lost. Type locality: "Exteris regionibus"; restricted to "Schweiz" (= Switzerland) by Mertens and Müller, 1928, Abh. Senckenb. Naturforsch. Ges., 41: 16.

Bufo salsus Schrank In Schrank and Moll, 1785, Naturhist. Briefe Oest. Salzburg Passau Berchtesgaden, 1: 308. Type(s): Not designated and presumed lost. Type locality: "Berchtesgaden", Bavaria, Germany. Synonymy by Mertens and Müller, 1928, Abh. Senckenb. Naturforsch. Ges., 41: 16.

Rana sonans Lacépède, 1788, Hist. Nat. Quadrup. Ovip. Serpens, 16mo ed., 2: 293, 457. Lacépède, 1788, Hist. Nat. Quadrup. Ovip. Serpens, Quarto ed., 1: Table following page 618, and referencing account starting on page 535. Type(s): Not stated, but including animal figured in vol. 2, pl. 13 of the 16mo edition and plate facing page 534 of 4to edition. Type locality: "Allemagne" (= Germany). Intended as a collective replacement name for Rana campanisona Laurenti (now = Rana temporaria) , Rana bombina Linnaeus, and Rana variegata Wurffbain (= Rana variegata Linnaeus). Rejected as published in a nonbinominal work by Opinion 2104, Anonymous, 2005, Bull. Zool. Nomencl., 62: 55.

Rana salsaGmelin, 1789, Syst. Nat., Ed. 13, 1(3): 1049.

Bombinator pachypus Bonaparte, 1838, Iconograph. Fauna Ital., 2 (Fasc. 23): unnumbered (119). Type(s): Not designated or known to exist. Type localities: "Monti Italiani . . . . delle Alpi Apuane . . . . Monti Ascolani"; restricted to "Apuanische Alpen, Norditalien" by Mertens and Müller, 1928, Abh. Senckenb. Naturforsch. Ges., 41: 16. See comment for history of synonymy. 

Bombinator brevipes Blasius, 1839, Isis von Oken, 32: 667. Type(s): Not designated or known to exist. Type locality: "am Unterharze, namentlich bei Goslar, in einem Sumpfe am Fusse des Hartzensteins und bei Osterode vorfommend"; given as swamp at foot of Katzenstein at Goslar by Mertens and Wermuth, 1960, Amph. Rept. Europas: 40. Synonymy by Boulenger, 1896, Boll. Mus. Zool. Anat. Comp. Univ. Torino, 11 (261): 1–2.

Bombinator scaber Küster, 1843, Isis von Oken, 36: 656. Type(s): Not designated. Type locality: "Lache bei Cettigne [= Centinje]", Montenegro. Synonymy by Mertens and Müller, 1940, Abh. Senckenb. Naturforsch. Ges., 451: 7.

Bombinator appeninicus Gistel in Gistel and Bromme, 1850, Handb. Naturgesch.: 333. Type(s): Not designated or known to exist. Type locality: "Italien auf den Apenninen [Mountains]". Synonymy with Bombina variegatus by Gorham, 1974, Checklist World Amph.: 42, and with Bombina pachypus by Stejneger, 1907, Bull. U.S. Natl. Mus., 58: 51.

Bombina maculatus Gistel, 1868, Die Lurche Europas: 162. Type(s): Not designated or known to exist. Type locality: Regensburg [Bavaria, Germany]. Synonymy by Mertens, 1936, Senckenb. Biol., 18: 76.

Bombina igneus var. brevipesKoch, 1872, Ber. Senckenb. Naturforsch. Ges., 1871–72: 165.

Bombinator pachypus var. Kolombatovici Bedriaga, 1890 "1889", Bull. Soc. Imp. Nat. Moscou, N.S.,, 3: 568. Type(s): ZIM. Type locality: "Dalmatien . . . . Spalato". Synonymy by Mertens and Müller, 1928, Abh. Senckenb. Naturforsch. Ges., 41: 16.

Bombinator pachypus var. brevipesBoulenger, 1896, Boll. Mus. Zool. Anat. Comp. Univ. Torino, 11 (261): 1-2.

Bombinator pachypus var. nigriventris Dürigen, 1897, Deutschl. Amph. Rept.: 546. Type(s): Not stated and presumed lost. Type locality: "bosnischen Exemplaren . . . [und] Montenegro" (= Bosnia and Montenegro). Synonymy by Gorham, 1974, Checklist World Amph.: 42.

Bombinator variegataPoche, 1912, Verh. Zool. Bot. Ges. Wien, 61: 405.

Bombina salsaNikolskii, 1918, Fauna Rossii, Zemnovodnye: 174; Stejneger, 1905, Science, 22: 502; Mertens, 1923, Senckenb. Biol., 5: 214.

Bombina salsa var. csikii Fejérváry, 1922, Mag. Tudom. Akd. Balkan Kutat. Tudomán. Eredm., 1: 24. Syntype(s): MNH 2540.5 (4 specimens; Ipek) 2540.6 (1 specimen; Mt. Korab), 2561.3 (1 specimen; Kula Lums), by original designation. Type localities: "Ipek", "Mount Korab collected, at 1800 m.", and "Kula Lums (from the banks of the Luma)", Albania; restricted to "Ipek, Albanien" by Mertens and Müller, 1928, Abh. Senckenb. Naturforsch. Ges., 41: 16. Synonymy with Bombina variegata scabra by Mertens and Müller, 1940, Abh. Senckenb. Naturforsch. Ges., 451: 7.

Bombina variegataMertens, 1928, Z. Morphol. Ökol. Tiere, 11: 614.

Bombina variegata pachypus — Mertens and Müller, 1928, Abh. Senckenb. Naturforsch. Ges., 41: 16.

Bombina variegata variegataMertens and Müller, 1928, Abh. Senckenb. Naturforsch. Ges., 41: 16.

Bombina variegata csikiiMertens and Müller, 1928, Abh. Senckenb. Naturforsch. Ges., 41: 16. Considered a junior synonym of Bombina variegata scabra by Mertens and Müller, 1940, Abh. Senckenb. Naturforsch. Ges., 451: 7.

Bombina variegata kolombatoviciMertens and Müller, 1928, Abh. Senckenb. Naturforsch. Ges., 41: 16.

Bombina variegata gracilis Bolkay, 1929, Glasn. Zemaljskog Muz. Bosni Hercegov., 41: 2. Holotype: Mus. Bosn. Hercegov. (presumably ZMBH). Type locality: "Zabrgje [= Zabre] (Northeastern Bosnia)", .

Bombina variegata scabraMüller, 1940, Mitt. König. Naturwiss. Inst. Sofia, 13: 1; Mertens and Müller, 1940, Abh. Senckenb. Naturforsch. Ges., 451: 7.

Bombina (Bombina) variegataTian and Hu, 1985, Acta Herpetol. Sinica, Chengdu, N.S.,, 4 (3): 219–224; Dubois, 1987 "1986", Alytes, 5: 97.

Bombina pachypus — Lanza and Vanni, 1991 "1990", in Internatl. Symp. Biogeograph. Aspects of Insularity: 337.

English Names

Saline Frog (Rana salsa [no longer recognized]: Shaw, 1802, Gen. Zool., 3(1): 119).

Yellow-bellied Toad (Hellmich, 1962, Rept. Amph. Eur.: 60; Arnold and Burton, 1978, Field Guide Rept. Amph. Brit. Eur.: 60; Ananjeva, Borkin, Darevsky, and Orlov, 1988, Dict. Amph. Rept. Five Languages: 51; Stumpel-Rienks, 1992, Ergänzungsband Handbuch Rept. Amph. Eur., Trivialnamen der Herpetofauna Eur.: 44; Kuzmin, 1999, Amph. Former Soviet Union: 201).

Yellowbelly Toad (Frank and Ramus, 1995, Compl. Guide Scient. Common Names Amph. Rept. World: 38).

Mountain Toad (Hellmich, 1962, Rept. Amph. Eur.: 60).

Appenine Yellow-bellied Toad (Bombina pachypus: Di Cerbo and Ferri, 1996, Naturschutz Rep., 11: 91).

Distribution

Europe (excluding the Iberian Peninsula and peninsular Italy south of the Po River in Italy), east to southern Poland, southwestern Ukraine, Bulgaria and Greece to European Turkey; introduced into England. 

Comment

The taxonomy of the Bombina variegata complex is inherently unstable due to how hybrid zones are interpreted. Those who favor lineage-species will look at the existing data and see several species (e.g., Bombina pachypus, Bombina scabra) while those favoring a reproductive species concept will opt for subspecies. Currently the latter is in favor, although in DRF's opinion most are doing one while claiming to do the other. Regardless, the currently most favored taxonomy is accepted for purposes of this catalog. See Poche, 1912, Verh. Zool. Bot. Ges. Wien, 61: 403–405, for initial sorting of Bombina bombina and Bombina variegata. See Arntzen, 1978, J. Biogeograph., 5: 335–345, for detailed statement of distribution. Lanza, 1983, Guide Reconoscimento Spec. Animali, Anf. Rett.: 83–86, provided an account of Bombina bombina pachypus. Schneider, Hussein, and Mamdouh, 1986, Bonn. Zool. Beitr., 37: 49–67, documented strong interpopulational variation of call structures. Gollmann, 1987, Amphibia-Reptilia, 8: 213–224, reported on hybridization with Bombina bombina in Hungary. Lang, 1988, Brit. Herpetol. Soc. Bull., 26: 6–13, summarized the subspecies and their distribution. Bombina pachypus was removed from the synonymy of Bombina variegata on the basis of genetic distance by Lanza and Vanni, 1991 "1990", in Internatl. Symp. Biogeograph. Aspects of Insularity: 335–344, and on the basis of morphology by (Nascetti, Vanni, Bullini, and Lanza, 1982, Boll. Zool., Padua, 49 (Suppl.): 134–135, where it had been placed by Bonaparte, 1840, Mem. Accad. Sci. Torino, Ser. 2, 2: 446, and Duméril and Bibron, 1841, Erp. Gen., 8: 488 (and with Bombinator igneus by Bonaparte, 1840, Icon. Faun. Ital., 2 (Introd.): preface, unnumbered). Nöllert and Nöllert, 1992, Die Amph. Eur.: 257–261, provided an account and polygon map. Szymura and Gollmann, 1996, Naturschutz Rep., 11: 9–15, documented strong electrophoretically distinctions between Bombina variegata variegataBombina variegata pachypus, and Bombina variegata scabraGollmann, Szymura, Arntzen, and Pialek, 1997, in Gasc et al. (eds.), Atlas Amph. Rept. Eur.: 98–99, discussed relevant literature and distribution. See accouns by Kuzmin, 1999, Amph. Former Soviet Union: 201–206, and Kuzmin, 2013, Amph. Former Soviet Union, Ed. 2: 115–117, for the former USSR. Braida, 2002, Riv. Idrobiol., 40: 239–246, discussed the distribution in Liguria, Italy. Arnold, 2002, Rept. Amph. Eur., Ed. 2: 52, provided a brief account, figure, and map (in the sense of including Bombina pachypus as a subspecies). Lever, 2003, Naturalized Rept. Amph. World: 140, reported on introduced populations in the vicinity of Devon, England. Barbieri, Bernini, Guarino, and Venchi, 2004, Boll. Zool., Padua, 71: 85–90, reported on distribution and conservation status in Italy. Fromhage, Vences, and Veith, 2004, Mol. Phylogenet. Evol., 31: 308–322, provided molecular data consistent with the recognition of Bombina pachypus but retained it as a member of Bombina variegataVörös, Alcobendas, Martínez-Solano, and García-París, 2005, Mol. Phylogenet. Evol., 38: 705–718, reported on mtDNA introgression of Bombina bombina and Bombina variegata in the Carpathian Basin and Spolsky, Szymura, and Uzzell, 2006, J. Zool. Syst. Evol. Res., 44: 100-104, suggested that the mtDNA data suggested two Pleistocene refugia for Bombina variegataCanestrelli, Cimmaruta, Costantini, and Nascetti, 2006, Mol. Ecol., 15: 3741–3754, discussed molecular phylogeography and Pleistocene biogeography. Bosman, Laan, and van Delft, 2009, in Creemers and van Delft (eds.), Amph. Rept. Nederland: 142–153, provided an account for the species in the Netherlands. Stojanov, Tzankov, and Naumov, 2011, Die Amph. Rept. Bulgariens: 159–167, provided a fairly detailed account and range map for Bulgaria. Cogǎlniceanu, Székely, Samoilă, Iosif, Tudor, Plăiaşu, Stănescu, and Rozylowicz, 2013, ZooKeys, 296: 35–57, provided a dot map for Romania. See detailed account by Lanza, Andreone, Bologna, Corti, and Razzetti, 2007, in Lanza et al. (eds.), Fauna d'Italia, 42 (Amph.): 277–280, for Italian population. Yu, Yang, Zhang, and Rao, 2007, J. Herpetol., 41: 365–377, suggested that Bombina pachypus is the sister taxon of Bombina variegatusHofman, Spolsky, Uzzell, Cogǎlniceanu, Babik, and Szymura, 2007, Mol. Ecol., 16: 2301–2316, discussed phylogeography of Bombina bombina and Bombina variegata and disputed the species distinctiveness of Bombina pachypus, although their results are also consistent with recognition of additional species within nominal Bombina variegata (DRF). Zheng, Fu, and Li, 2009, Mol. Phylogenet. Evol., 52: 70–83, reported molecular evidence on the phylogenetics of populations within the nominal species. Speybroeck, Beukema, and Crochet, 2010, Zootaxa, 2492: 6, discussed their reasons from considering Bombina pachypus a subspecies of Bombina variegataFijarczyk, Nadachowska, Hofman, Litvinchuk, Babik, Stuglik, Gollmann, Choleva, Cogǎlniceanu, Vukov, Džukić, and Szymura, 2011, Mol. Ecol., 20: 3381–339, also regarded Bombina pachypus as a diagnosable subspecies of Bombina bombina, but this is a concept- rather than data-based difference of opinion. Mori and Giovani, 2012, Herpetol. Notes, 5: 407–412, documented the disappearance of this species in areas of Tuscany, Italy. Pabijan, Wandycz, Hofman, Węcek, Piwczyński, and Szymura, 2013, Mol. Phylogenet. Evol., 69: 63–74, considered their Bombina variegata scabra to be the sister taxon of Bombina pachypus + Bombina scabra, but treated them as subspecies. Speybroeck, Beukema, Bok, and Van Der Voort, 2016, Field Guide Amph. Rept. Brit. Eur.: 135–137, provided a brief account (as Bombina variegata pachypus). Guicking, Finke, Wittich, Pfeiffer, Veith, Geske, Braukmann, Weising, and Neubeck, 2017, Salamandra, 53: 201–211, reported on conservation genetics in northern Hesse, Germany. Speybroeck, Beukema, Bok, and Van Der Voort, 2016, Field Guide Amph. Rept. Brit. Eur.: 135–137, provided a brief account (in the sense of including Bombina pachypus) and distribution map. Bülbül, Kurnaz, Eroğlu, Koç, and Kutrup, 2018, Russ. J. Herpetol., 25: 236–238, mapped the range in Turkey and discussed its conservation and threat conditions. Weihmann, Weihmann, and Weihmann, 2019, Conserv. Genetics, 20: 557–569, reported on conservation genetics of populations, native, and introduced in Lower Saxony, Germany. See Dufresnes, 2019, Amph. Eur., N. Afr., & Middle East: 44, for brief summary of identifying morphology and biology as well as a range map and photograph (see p. 194 for photographs and map of subspecies). Pop, Lucaci, and Covaciu-Marcov, 2019, Herpetol. Notes, 12: 1001–1004, reported the species in southern Romanian Moldavia. Dufresnes, Suchan, Smirnov, Denoël, Rosanov, and Litvinchuk, 2021, J. Biogeograph., 48: 548–560, reported on hybridization and introgression with Bombina bombina in Poland and Ukraine. See Dufresnes, 2019, Amph. Eur., N. Afr., & Middle East: 45, for brief summary of identifying morphology and biology as well as a range map and photograph.

External links:

Please note: these links will take you to external websites not affiliated with the American Museum of Natural History. We are not responsible for their content.