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, 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. Synonymy by Pabijan, Wandycz, Hofman, Węcek, Piwczyński, and Szymura, 2013, Mol. Phylogenet. Evol., 69: 63-74. 

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 [no longer recognized]: 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 western Ukraine and Moldova; introduced into England. 

Comment

See Arntzen, 1978, J. Biogeogr., 5: 335-345, for detailed statement of distribution. Schneider, Hussein, and Mamdouh, 1986, Bonn. Zool. Beitr., 37: 49-67, documented strong interpopulational variation of call structures. Szymura and Gollmann, 1996, Naturschutz Rep., 11: 9-15, documented strong electrophoretically distinctions between Bombina variegata variegata, Bombina variegata pachypus, and Bombina variegata scabra. Nöllert and Nöllert, 1992, Die Amph. Eur.: 257-261, provided an account and polygon map. See account by Kuzmin, 1999, Amph. Former Soviet Union: 201-206. Gollmann, Szymura, Arntzen, and Pialek, 1997, in Gasc et al. (eds.), Atlas Amph. Rept. Eur.: 98-99, discussed relevant literature and distribution. See Poche, 1912, Verh. Zool. Bot. Ges. Wien, 61: 403-405, for initial sorting of Bombina bombina and Bombina variegata. Lang, 1988, Brit. Herpetol. Soc. Bull., 26: 6-13, summarized the subspecies and their distribution. 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. Vörös, 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 variegata. 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. Bombina pachypus was emoved 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 Nascetti, Vanni, Bullini, and Lanza, 1982, Boll. Zool., Padua, 49 (Suppl.): 134-135, 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). Considered by some authors (e.g., Mertens and Müller, 1928, Abh. Senckenb. Naturforsch. Ges., 41: 16; Arnold, 2002, Rept. Amph. Eur., Ed. 2: 60) to be a race of Bombina variegataLanza and Vanni, 1991 "1990", in Internatl. Symp. Biogeograph. Aspects of Insularity: 337, doubted the reliability of records of this species from Sicily. Lanza, 1983, Guide Reconoscimento Spec. Animali, Anf. Rett.: 83-86, provided an account as Bombina bombina pachypusBraida, 2002, Riv. Idrobiol., 40: 239-246, discussed the distribution in Liguria, Italy. Fromhage, Vences, and Veith, 2004, Mol. Phylogenet. Evol., 31: 308-322, provided molecular data consistent with the recognition of Bombina pachypus as a distinct species but retained it as a member of Bombina variegataYu, Yang, Zhang, and Rao, 2007, J. Herpetol., 41: 365-377, suggested that Bombina pachypus is the sister taxon of Bombina variegatus. See detailed account by Di Cerbo and Bressi, 2007, in Lanza et al. (eds.), Fauna d'Italia, 42 (Amph.): 280-287, for Italian population. Hofman, 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 that is consistent with both views. 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. 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.  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 bombina, but treated them as subspecies, an arrangement followed here but reluctantly as the notion of subspecies has no particular meaning in any concept of species other than the Biological Species Concept, a concept not consistent with phylogenetic aims. The data as they exist are also consistent with three lineage-species: Bombina pachypus, Bombina variegata, and Bombina scabra (DRF; 23 June 2013). Guicking, Finke, Wittich, Pfeiffer, Veith, Geske, Braukmann, Weising, and Neubeck, 2017, Salamandra, 53: 201–211, reported on conservation genetics in northern Hesse, Germany. 

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