Ptychadena "mascareniensis"

Class: Amphibia > Order: Anura > Family: Ptychadenidae > Genus: Ptychadena > Species: Ptychadena "mascareniensis"

Rana savignyi Jan, 1857, Cenni Mus. Civ. Milano: 52. Types: MCSM. Type locality: not stated. Considered a synonym of Rana mascareniensis (along with several other now-recognized species) by Boulenger, 1879, Bull. Soc. Zool. France, 4: 92-94; Boulenger, 1882, Cat. Batr. Sal. Coll. Brit. Mus., Ed. 2: 52. May actually apply to any of the Ptychadena mascareniensis complex (e.g., Ptychadena nilotica) (DRF). See comment. 

Rana marchei Rochebrune, 1885, Bull. Soc. Philomath., Paris, Ser. 7, 9: 90. Types: "Mus. Bouvieri"; presumably to MNHNP but not known to be there. Type locality: "ad ripas rivuli Sangourougou", Senegal. Synonymy with Ptychadena mascareniensis by Noble, 1924, Bull. Am. Mus. Nat. Hist., 49: 220. See comment.

Rana venusta Werner, 1908 "1907", Sitzungsber. Akad. Wiss. Wien, Phys. Math. Naturwiss. Kl., 116: 1892. Syntypes: NHMW (3 specimens), by original designation; NHMW 12235 considered holotype by Häupl, Tiedemann, and Grillitsch, 1994, Kat. Wiss. Samml. Naturhist. Mus. Wien, 9: 33. Type localities: "im Sudan ausschliesslich bei Mongolla", Sudan (3 specimens). Synonymy with Ptychadena mascareniensis by Loveridge, 1936, Field Mus. Nat. Hist. Publ., Zool. Ser., 22: 94; Guibé and Lamotte, 1957, Bull. Inst. Franç. Afr. Noire, Ser. A, 19: 979. See comment.

Rana mascareniensis venustaLoveridge, 1933, Bull. Mus. Comp. Zool., 74: 370; Loveridge, 1936, Field Mus. Nat. Hist. Publ., Zool. Ser., 22: 94; Mertens, 1938, Senckenb. Biol., 20: 425. See comment.

Rana (Ptychadena) mascareniensis venustaPerret and Mertens, 1957, Bull. Inst. Franç. Afr. Noire, Ser. A, 19: 568. See comment.

Distribution

The range is an aggregation of unnamed species and unallocated populations within the Ptychadena mascareniensis complex: From West Africa in Savanna regions to Uganda, Angola, northern Namibia, northern Botswana, and northeastern KwaZulu-Natal (Rep. South Africa); 

Comment

Vences, Kosuch, Rödel, Channing, Glaw, and Böhme, 2004, J. Biogeograph., 31: 593-601, suggested on the basis of mtDNA evidence that nominal Ptychadena mascareniensis is composed of a number of species, with Ptychadena mascareniensis sensu stricto restricted to Madagascar, the Seychelles, and the Mascarenes. Although mtDNA has limited application to identifying lineages of organisms, Dehling and Sinsch, 2013, Zool. Anz., 253: 143-157, employed the same gene to delimit and resurrect the name Ptychadena nilotica for the major Nile River drainage (Clade A), which leaves a hodge-podge of unnamed or nomenclaturally unsorted populations from West, Southwest, and East Africa (e.g., Clade B from Cameroon and Benin; Clade C from Ivory Coast; and Clade Clade D from Kenya) and populations from Southwestern and South Africa that have not yet been sampled. Rather than continue to conflate Ptychadena mascareniensis with mainland populations or with Ptychadena nilotica, I just treat the remaining populations as Ptychadena "mascareniensis". This is highly unsatisfactory, but I do not want to submerge Ptychadena nilotica and Ptychadena mascareniensis into a larger taxon just because the revision of the group is incomplete (DRF, Oct. 2014).  Zimkus, Lawson, Barej, Barratt, Channing, Dash, Dehling, Du Preez, Gehring, Greenbaum, Gvoždík, Harvey, Kielgast, Kusamba, Nagy, Pabijan, Penner, Rödel, Vences, and Lötters, 2016, Mol. Phylogenet. Evol., 106: 254–269, further investigated the mainland group, including refining the concepts of Ptychadena nilotica and including Ptychadena newtoni within the Ptychadena mascareniensis complex. Non-Madagascan, unnamed lineages/species within this complex noted by  Zimkus, Lawson, Barej, Barratt, Channing, Dash, Dehling, Du Preez, Gehring, Greenbaum, Gvoždík, Harvey, Kielgast, Kusamba, Nagy, Pabijan, Penner, Rödel, Vences, and Lötters, 2016, Mol. Phylogenet. Evol., 106: 254–269, are (using their numbers for populations): 4: Ghana along the coast (presumably) to western Cameroon; 5: Mount Nimba highlands of Sierra Leone, Guinea, and Liberia; 6: equatorial region of Gabon, Rep. Congo, western Dem. Rep. Congo, east to northeastern Dem. Rep. Congo, Uganda, western Kenya; 7: southeastern Cameroon, exreme southwestern Cent. Afr. Rep., west-central Rep. Congo; 8: north-central Dem. Rep. Congo; 9: south-central Tanzania. Presumably these lineages will be appropriately named in the near future. 

Accounts provided that apply to this complex were provided by Schmidt and Inger, 1959, Explor. Parc Natl. Upemba, Miss. G.F. de Witte, 56: 76-85, who described Ptychadena mascareniensis hylaea. Hylaea was subsequently considered a distinct species by Lamotte, 1967, Bull. Mus. Natl. Hist. Nat. Paris, Ser. 2, 39: 647-656, but Perret, 1979, Bull. Soc. Neuchatel. Sci. Nat., 102: 7-17, considered Ptychadena bibroni (Hallowell, 1845) to be a West African subspecies (following Boulenger, 1882, Cat. Batr. Sal. Coll. Brit. Mus., Ed. 2: 523) and hylaea to be a synonym of bibroni. See Poynton, 1964, Ann. Natal Mus., 17: 129, Poynton and Broadley, 1985, Ann. Natal Mus., 27: 148-149, and Lambiris, 1988, Lammergeyer, 39: 84. Perret, 1997, Bull. Soc. Neuchatel. Sci. Nat., 120: 77, discussed confusion in the literature between Ptychadena arnei and "Ptychadena bibroni". See Largen, 1997, Tropical Zool., 10: 71, for comments on misidentified Ptychadena anchietae ascribed to this species and for discussion of distribution. Rödel, 2000, Herpetofauna W. Afr., 1: 128-132, discussed the issue of difficulty of identification of species related to Ptychadena mascareniensis and provided a comparison with Ptychadena bibroni, Ptychadena pujoli, Ptychadena mascareniensis hylaea, and Ptychadena arnei. Channing, 2001, Amph. Cent. S. Afr.: 329-331, and Du Preez and Carruthers, 2009, Compl. Guide Frogs S. Afr.: 312-313, provided accounts for the southern African populations. Largen, 2001, Tropical Zool., 14: 337-338, commented on distribution. Andreone, Glaw, Nussbaum, Raxworthy, Vences, and Randrianirina, 2003, J. Nat. Hist., London, 37: 2119-2149, discussed the occurrence of this species on Nosy Be Vences, Kosuch, Rödel, Channing, Glaw, and Böhme, 2004, J. Biogeograph., 31: 593-601, studied molecular phylogeography, provided a map of the range, and suggested that this nominal taxon contains at least five cryptic species: (1) the population in Madagascar (which they regarded as having arrived via over-water dispersal), the Seychelles, and Reunion; (2-3) two in West Africa; and (4-5) two in East Africa. Channing and Howell, 2006, Amph. E. Afr.: 306-308, provided an account for East Africa. Minter and Passmore, 2004, in Minter et al. (eds.), Atlas Frogs S. Afr. Lesotho and Swaziland: 286-287, provided an account for South Africa. Padial and De la Riva, 2004, Rev. Esp. Herpetol., 18: 94, provided records of this species, or one of the complex, from Mauritania. Baha El Din, 2006, Guide Rept. Amph. Egypt: 37-39, provided an account for Egypt and distinguished between it and the Delta population of Ptychadena schillukorum. Pickersgill, 2007, Frog Search: 128-131, provided an account and noted an apparently unnamed species from Arusha, Tanzania, masquerading under this name. Lötters, Wagner, Bwong, Schick, Malonza, Muchai, Wasonga, and Veith, 2007, Fieldguide Amph. Rept. Kakamega Forest: 21-22, provided an account for Ptychadena cf mascareniensis in Kenya. Jacobsen, 2009, Afr. Herp News, 47: 2-20, reported this species from east-north-central Central African Republic. Bwong, Chira, Schick, Veith, and Lötters, 2009, Salamandra, 45: 129-146, discussed the assignment of a population in western Kenya to this taxon and noted that if Ptychadena mascareniensis constitutes a species complex, then the name Rana venusta Werner, 1908, is likely the appropriate name for the northeastern population. See account, photograph, and map for Ethiopia by Largen and Spawls, 2010, Amph. Rept. Ethiopia Eritrea: 172, who noted that records for Eritrea are based on misidentifications. Mercurio, 2011, Amph. Malawi: 256-258, provided an account for Malawi. Measey, Malonza, and Muchai, 2009, Amph. Taita Hills: 38-41, provided a brief account and photo for the Taita Hills, Kenya. Cole, 2009, Field Guide. Rept. Amph. Mauritius: 75-76, provided a brief account for Mauritius. Channing, Rödel, and Channing, 2012, Tadpoles of Africa: 308–309, provided information on comparative larval morphology. Harper, Measey, Patrick, Menegon, and Vonesh, 2010, Field Guide Amph. E. Arc Mts. Tanzania and Kenya: 264–265, provided a brief account and photograph. 

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