Rana macrocnemis Boulenger, 1885

Class: Amphibia > Order: Anura > Family: Ranidae > Genus: Rana > Species: Rana macrocnemis

Rana macrocnemis Boulenger, 1885, Proc. Zool. Soc. London, 1885: 22. Holotype: BMNH by original designation. Type locality: "Brusa", Turkey.

Rana camerani Boulenger, 1886, Bull. Soc. Zool. France, 11: 597. Syntypes: BMNH (1 specimen), ZMB 9345 (3 specimens) and 9344 (1 specimen), according to XXX. Type locality: "Tabizhuri (2500 m)" and "Achalkalki", Georgia; given as "Tabistskhuri Lake [2500 m elevation] and Akhalkalaki Settlement, Georgia" by Tuniyev, 1999, Adv. Amph. Res. Former Soviet Union, 3: 58. Synonymy by Kuzmin, 1999, Amph. Former Soviet Union: 288-291; Veith, Kosuch, and Vences, 2003, Mol. Phylogenet. Evol., 26: 325.

Rana (Crotaphitis) cameraniSchulze, 1891, Jahresber. Abhandl. Naturwiss. Ver. Magdeburg, 1890: 177.

Rana cameranoiBoettger, 1892, Kat. Batr. Samml. Mus. Senckenb. Naturforsch. Ges.: 8. Incorrect subsequent spelling.

Rana temporaria warenzowi Terentjev, 1923, Proc. First Congr. Russ. Zool. Anat. Hist. Petrograd: 35. Syntypes: Not stated; ZMM A-448 (5 specimens), now lost, according to Dunayev and Orlova, 1994, Russ. J. Herpetol., 1: 65. Type locality: Not stated. Synonymy by Borkin, 1977, Trudy Zool. Inst. Akad. Nauk SSSR, Leningrad, 74: 24-31.

Rana macrocnemis angeloi Bogachev, 1927, Izvestia Azerbaidzhanskogo Gosudorstvennogo Universitete, Otdel Yectestvozaniya y Meditsini, 6: 121. Holotype: GIN Moscow (number not given), articulated skeleton. Type locality: Kisatibi Georgia [Pliocene fossil]

Rana macrocnemis macrocnemisBogachev, 1927, Izvestia Azerbaidzhanskogo Gosudorstvennogo Universitete, Otdel Yectestvozaniya y Meditsini, 6: 121, by implication; Eiselt and Schmidtler, 1971, Ann. Naturhist. Mus. Wien, 75: 384.

Rana kisatibensis Riabinin, 1928 "1927", Ezhiegodnik Russkogo Palaeontologicheskogo Obshchestva, 7: 88. Holotype: GIN (number not given), articulated skeleton. Type locality: Kissatibi Georgia. [Pliocene fossil.] Named as an objective synonym of Rana macrocnemis angeloi Bogachev, 1927.

Rana (Rana) macrocnemisDubois, 1987 "1986", Alytes, 5: 41, by implication; Dubois, 1992, Bull. Mens. Soc. Linn. Lyon, 61: 333; Yuan, Zhou, Chen, Poyarkov, Chen, Jang-Liaw, Chou, Matzke, Iizuka, Min, Kuzmin, Zhang, Cannatella, Hillis, and Che, 2016, Syst. Biol., 65: 835.

Rana (Rana) cameraniDubois, 1987 "1986", Alytes, 5: 41-42, by implication; Dubois, 1992, Bull. Mens. Soc. Linn. Lyon, 61: 333.

Rana macrocnemis cameraniVeith, Kosuch, and Vences, 2003, Mol. Phylogenet. Evol., 26: 325.

Rana macrocnemis macrocnemisVeith, Kosuch, and Vences, 2003, Mol. Phylogenet. Evol., 26: 325.

Rana (Laurasiarana) macrocnemisHillis and Wilcox, 2005, Mol. Phylogenet. Evol., 34: 311, by implication; see Dubois, 2006, Mol. Phylogenet. Evol., 42: 317-330, and Hillis, 2007, Mol. Phylogenet. Evol., 42: 331-338, for discussion.

Rana Holtzi Werner, 1898, Zool. Anz., 21: 222. Syntypes: NHMW 15086 (according to Häupl and Tiedemann, 1978, Kat. Wiss. Samml. Naturhist. Mus. Wien, 2: 27, and Häupl, Tiedemann, and Grillitsch, 1994, Kat. Wiss. Samml. Naturhist. Mus. Wien, 9: 31), MCZ 8131 (according to Barbour and Loveridge, 1929, Bull. Mus. Comp. Zool., 69: 328), MZUT An41 (according to Gavetti and Andreone, 1993, Cat. Mus. Reg. Sci. Nat., Torino, 10: 95 ), and ZFMK 18913-14 (according to Böhme and Bischoff, 1984, Bonn. Zool. Monogr., 19: 183). Type locality: "an einem bisher noch unbekannten See, Maidan Göl, im cilischen Taurus, 2400 mm hoch gefangen", Turkey. 

Rana (Rana) holtzi — Dubois, 1987 "1986", Alytes, 5: 41, by implication.

Rana macrocnemis holtzi — Veith, Kosuch, and Vences, 2003, Mol. Phylogenet. Evol., 26: 325.

Rana holtzi — Baran, Ilgaz, Kumlutas, Olgun, Avci, and Iret, 2007, Turkish J. Zool., 31: 1–7; Afsar, Afsar, and Arikan, 2015, Herpetozoa, Wien, 28: 15–27. 

English Names

Brusa Frog (Rana macrocnemis: Frank and Ramus, 1995, Compl. Guide Scient. Common Names Amph. Rept. World: 108).

Iranian Long-legged Wood Frog (Rana macrocnemis: Ananjeva, Borkin, Darevsky, and Orlov, 1988, Dict. Amph. Rept. Five Languages: 125).

Iranian Long-legged Frog (Rana macrocnemis: Borkin, 1997, in Gasc et al. (eds.), Atlas Amph. Rept. Eur.: 150; Kuzmin, 1999, Amph. Former Soviet Union: 334).

Caucasian Brown Frog (Rana macrocnemis: Borkin, 1997, in Gasc et al. (eds.), Atlas Amph. Rept. Eur.: 150; Tarkhnishvili and Gokhelashvili, 1999, Adv. Amph. Res. Former Soviet Union, 4: v; Arekelyan, Danielyan, Corti, Sindaco, and Leviton, 2012, Herpetofauna of Armenia: 45).

Caucasus Frog (Rana macrocnemis camerani: Frank and Ramus, 1995, Compl. Guide Scient. Common Names Amph. Rept. World: 106).

Banded Frog (Rana macrocnemis camerani: Baloutch and Kami, 1995, Amph. Iran: 157).

Transcaucasian Frog (Rana macrocnemis camerani: Baloutch and Kami, 1995, Amph. Iran: 157).

Long-legged Wood Frog (Safaei-Mahroo, Ghaffari, Fahimi, Broomand, Yazdanian, Najafi-Majd, Hosseinian Yousefkani, Rezazadeh, Hosseinzadeh, Nasrabadi, Rajabizadeh, Mashayekhi, Motesharei, Naderi, and Kazemi, 2015, Asian Herpetol. Res., 6: 266). 

Turkish Frog (Rana holtzi [no longer recognized]: Frank and Ramus, 1995, Compl. Guide Scient. Common Names Amph. Rept. World: 107).

Distribution

West-centraland southeastern Asia Minor of Turkey to Caucasus Mountains of Russia, Georgia, Armenia, and Azerbaijan; thence into northwestern Iran (West Azarbaijan, East Azarbaijan, Ardabil, Zanjan, Kordestan, Kermanshah, Hamedan, Qazvin, and Mazandaran provinces); isolated population in Kopet-Dag mountains of Turkmenistan.

Comment

In the Section Rana, subgenus Rana, Rana temporaria group of Dubois, 1992, Bull. Mens. Soc. Linn. Lyon, 61: 333. The distinction/synonymy of Rana macrocnemis and Rana camerani is controversial; see Delwig, 1928, Zool. Anz., 79: 37-44; Mensi, Lattes, Macario, Salvidio, Giacoma, and Balletto, 1992, Zool. J. Linn. Soc., 105: 293-311, Tarkhnishvili and Gokhelashvili, 1996, Alytes, 14: 27-41; Kuzmin, 1999, Amph. Former Soviet Union: 288-291 (who rejected all subspecies); and Çevik, Arikan, Kaya, and Atatür, 2006, Amphibia-Reptilia, 27: 63-71 for discussion and transfer of the population from Mount Erciyes from Rana camerani to Rana macrocnemis. See also Borkin, 1977, Trudy Zool. Inst. Akad. Nauk SSSR, Leningrad, 74: 24-31; Mensi, Lattes, Macario, Salvidio, Giacoma, and Balletto, 1992, Zool. J. Linn. Soc., 105: 293-311, for discussion of phylogenetic relationships. Kami and Vakilpoure, 1996, Herpetol. Rev., 27: 150, provided a record for Chaharmahal Baktiari Province and commented on the range (as Rana camerani). Tarkhnishvili, Arntzen, and Thorpe, 1999, Herpetologica, 55: 406-417, studied the morphometrics of frogs from Georgia and provided evidence of considerable geographic variation, though not species boundaries between nominal Rana macrocnemis and Rana camerani. Tarkhnishvili, Hille, and Böhme, 2001, Biol. J. Linn. Soc., 74: 141-156, looked at the issue of Rana macrocnemis and Rana camerani using mtDNA and reported on the complex system of hybridization and seeming intergradation between the forms. Ishchenko and Pjastolova, 1973, Zool. Zh., 52: 1733-35; Kuzmin, 1999, Amph. Former Soviet Union: 335-336; Picariello, Scillitani, Feliciello, Bellinello, Maresca, and Chinali, 2000, Riv. Idrobiol., 38: 187, considered Rana camerani to be a junior synonym of Rana macrocnemis. Başoğlu and Özeti, 1973, Türkiye Amphibileri: 103-105, provided accounts for the Turkish populations (as Rana macrocnemis and Rana cameranoi). Borkin, 1997, in Gasc et al. (eds.), Atlas Amph. Rept. Eur.: 150-151, discussed relevant literature and range. Tuniyev, 1999, Adv. Amph. Res. Former Soviet Union, 3: 58-59, summarized the taxonomic history. See Baran, 1969, Sci. Rep. Fac. Sci. Ege Univ., Izmir, 80: 1-78 (who removed Rana holtzi from the synonymy of Rana camerani, where it had been placed by Boulenger, 1898, Tailless Batr. Eur., 2: 298), and Başoğlu and Özeti, 1973, Türkiye Amphibileri: 105-106. Mensi, Lattes, Macario, Salvidio, Giacoma, and Balletto, 1992, Zool. J. Linn. Soc., 105: 293-311, reported on phylogenetic relationships. Veith, Kosuch, and Vences, 2003, Mol. Phylogenet. Evol., 26: 310-327, suggested that Rana macrocnemis macrocnemis, Rana camerani, and Rana holtzi are conspecific, and that other populations of "Rana macrocnemis" (now Rana tavasensis and Rana pseudodalmatina) are outside of this clade. Özeti and Yilmaz, 1994, Türkiye Amfibileri: 153-156, provided an account for Turkish populations. Brief account provided by Günther IN Engelmann, Fritzsche, Günther, and Obst, 1993, Lurche Kriechtiere Eur.: 167-168. Baloutch and Kami, 1995, Amph. Iran: 159-160, provided an account for Iran. Veith, Kosuch, and Vences, 2003, Mol. Phylogenet. Evol., 26: 310-327, and Veith, Schmidtler, Kosuch, Baran, and Seitz, 2003, Mol. Ecol., 12: 185-199, provided evidence that Rana macrocnemis tavasensis and Rana macrocnemis pseudodalmatina are distinct species. See comments under Rana tavasensis and Rana pseudodalmatina. Crochet and Dubois, 2004, In Gasc et al. (eds)., Atlas Amph. Rept. Europe, Ed. 2: 499, discussed the issue of Rana camerani and Rana macrocnemis and thought that they should be regarded as subspecies. Arekelyan, Danielyan, Corti, Sindaco, and Leviton, 2012, Herpetofauna of Armenia: 45-46, provided a brief account for Armenia and environs. Afsar, Afsar, and Arikan, 2014, Herpetozoa, Wien, 26: 115-123, reported a population in the Emir Mountains of western inner Anatolia, Turkey, and reviewed the taxonomic history of the Rana macrocnemis–Rana holtzi–Rana tavasensis controversies. Afsar, Afsar, and Arikan, 2015, Herpetozoa, Wien, 28: 15–27, discussed populations of southeastern Turkey previously of uncertain species assignment. Safaei-Mahroo, Ghaffari, Fahimi, Broomand, Yazdanian, Najafi-Majd, Hosseinian Yousefkani, Rezazadeh, Hosseinzadeh, Nasrabadi, Rajabizadeh, Mashayekhi, Motesharei, Naderi, and Kazemi, 2015, Asian Herpetol. Res., 6: 257–290, reported on distribution and conservation status in Iran. Rana holtzi was emoved from the synonymy of Rana macrocnemis on the basis of morphological differences by Baran, Ilgaz, Kumlutas, Olgun, Avci, and Iret, 2007, Turkish J. Zool., 31: 1–7, where it had been placed by Picariello, Feliciello, Scillitani, Cataudo, Maresca, and Chinali, 2000, Riv. Idrobiol., 38: 167, and Veith, Kosuch, and Vences, 2003, Mol. Phylogenet. Evol., 26: 325. See discussion of Rana holtzi by Afsar, Afsar, and Arikan, 2015, Herpetozoa, Wien, 28: 15–27. See Baran, 1969, Sci. Rep. Fac. Sci. Ege Univ., Izmir, 80: 1-78 (who removed Rana holtzi from the synonymy of Rana camerani, where it had been placed by Boulenger, 1898, Tailless Batr. Eur., 2: XXX), and Başoğlu and Özeti, 1973, Türkiye Amphibileri: 105-106 (who provided an account of Rana holtzi)Mensi, Lattes, Macario, Salvidio, Giacoma, and Balletto, 1992, Zool. J. Linn. Soc., 105: 293-311, reported on phylogenetic relationships. See comment under Rana macrocnemis. Kalaycı, Kalaycı, and Özdemir, 2017, Biochem. Syst. Ecol., 73: 26–34, provided evidence from mt and nuDNA that Rana holtzi represents part of the geographic variation of Rana macrocnemis. Najibzadeh, Veith, Gharzi, Rastegar-Pouyani, Rastegar-Pouyani, Kieren, and Pesarakloo, 2017, Amphibia-Reptilia, 38: 339–250, employing solely two partial mtDNA loci suggested that Rana holtzi, Rana camerani, and Rana macrocnemis are distinct lineages. 

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