Hyperolius Rapp, 1842

Class: Amphibia > Order: Anura > Family: Hyperoliidae > Subfamily: Hyperoliinae > Genus: Hyperolius
143 species

Eucnemis Tschudi, 1838, Classif. Batr.: 35, 76. Type species: Hyla horstocki Schlegel, 1837, by monotypy; see Dubois, 1981, Monit. Zool. Ital., N.S., Suppl., 15: 264, for a discussion of the type species. Homonym of Eucnemis Ahrens, 1812, N. Schrift. Naturforsch. Ges. Halle, 2. (Insecta).

Hyperolius Rapp, 1842, Arch. Naturgesch., 8: 289. Replacement name for Eucnemis Tschudi, 1838.

Epipole Gistel, 1848, Naturgesch. Thierr.: ix. Substitute name for Eucnemis Tschudi, 1838.

EuchnemisBianconi, 1849 "1848", Nuovi Ann. Sci. Nat., Bologna, Ser. 2, 10: 107. Incorrect subsequent spelling of Eucnemis Tschudi, 1838.

Crumenifera Cope, 1862, Proc. Acad. Nat. Sci. Philadelphia, 14: 343. Type species: Crumenifera pusilla Cope, 1862, by monotypy. Synonymy with Rappia by Boulenger, 1882, Cat. Batr. Sal. Coll. Brit. Mus., Ed. 2: 127.

Rappia Günther, 1865, Zool. Rec., 1: 130. Substitute name for Hyperolius Rapp, 1842, thought to be preoccupied by Uperolia Gray. Synonymy by Noble, 1924, Bull. Am. Mus. Nat. Hist., 49: 251; Ahl, 1931, Das Tierreich, 55: 254.

CruminiferaHoffmann, 1878, in Bronn (ed.), Die Klassen und Ordnungen des Thier-Reichs, 6(2): 611. Incorrect subsequent spelling of Crumenifera Cope, 1863.

Nesionixalus Perret, 1976, Arq. Mus. Bocage, Ser. 2, 6: 29. Type species: Hyperolius thomensis Bocage, 1886, by original designation. Synonymy by Drewes, 1984, Occas. Pap. California Acad. Sci., 139: 52.

Chlorolius Perret, 1988, Bull. Soc. Neuchatel. Sci. Nat., 111: 46. Type species: Hyperolius koehleri Mertens, 1940, by original designation. Synonymy by Amiet, 2012, Rainettes Cameroun: 362. Synonymy confirmed by Portik and Blackburn, 2016, Evolution, 70: 2024. 

Alexteroon Perret, 1988, Bull. Soc. Neuchatel. Sci. Nat., 111: 39. Type species: Hyperolius obstetricans Ahl, 1931, by original designation. Synonymy by Dubois, Ohler, and Pyron, 2021, Megataxa, 5: 459, although the evidence provided by Pyron and Wiens, 2011, Mol. Phylogenet. Evol., 61: 557, first provided the evidence of paraphyly of HyperoliusPortik, Streicher, and Wiens, 2023, Mol. Phylogenet. Evol., 188 (107907): 66, also discovered this paraphyly without reference to earlier work. 

Nomina inquirenda - Name(s) unassigned to a living or extinct population

Rappia granulata Tornier, 1896, in Möbius (ed.), Deutsch Ost-Afr., 3: 151. Syntypes: ZMB 4811 (2 specimens), by original designation. Type locality: "Tette", Tanzania. Inadvertent instantiation of name credited by Tornier to Peters.

Rappia fimbriata Tornier, 1896, in Möbius (ed.), Deutsch Ost-Afr., 3: 153. Syntypes: ZMB, by original designation. Type locality: "Gowe Limbareni" and "Bukova", Tanzania. Inadvertent instantiation of name credited by Tornier to Duméril and Bibron.

Hyperolius thoracotuberculatus Ahl, 1931, Mitt. Zool. Mus. Berlin, 17: 98. Holotype: ZMB 36097, according to Tillack, de Ruiter, and Rödel, 2021, Zoosyst. Evol., Berlin, 97: 436. Type locality: "Afrika (ohne genauen Fundort)" (= Africa, without precise locality data). The description in Ahl, 1931, Das Tierreich, 55: 371 Ahl, 1931, Mitt. Zool. Mus. Berlin, 17: 98, appeared subsequently according to :Tillack, de Ruiter, and Rödel, 2021, Zoosyst. Evol., Berlin, 97 436. Warty Reed Frog (Frank and Ramus, 1995, Compl. Guide Scient. Common Names Amph. Rept. World: 68). Not mentioned by Schiøtz, 1999, Treefrogs Afr. Appears to be a member of the Hyperolius viridiflavus group and likely a synonym of one of the members of this group. 

Hyperolius laticeps Ahl, 1931, Mitt. Zool. Mus. Berlin, 17: 69. Holotype: ZMB unnumbered, according to the original publication; given as ZMB 46529 by Tillack, de Ruiter, and Rödel, 2021, Zoosyst. Evol., Berlin, 97: 427. Type locality: "Togo". The description in Ahl, 1931, Das Tierreich, 55: 342, appeared subsequently according to Tillack, de Ruiter, and Rödel, 2021, Zoosyst. Evol., Berlin, 97: 427. Togo Reed Frog (Frank and Ramus, 1995, Compl. Guide Scient. Common Names Amph. Rept. World: 67). Known only from the type locality (Togo). Not mentioned by Schiøtz, 1999, Treefrogs Afr.Segniagbeto, Bowessidjaou, Dubois, and Ohler, 2007, Alytes, 24: 77, presumed it to be a junior synonym of some other species of Hyperolius but did not suggest a candidate. Channing and Rödel, 2019, Field Guide Frogs & Other Amph. Afr.: 396, regarded this nominal taxon as too insufficiently known to treat adequately in their fieldguide. Tillack, de Ruiter, and Rödel, 2021, Zoosyst. Evol., Berlin, 97: 427, noted that the holotype is a juvenile and cannot be confidently assigned to any West African species. 

English Names

Reed Frogs (Broadley, 1971, Puku, 6: 121; Passmore and Carruthers, 1978, J. Herpetol. Assoc. Afr., 19: 7; Passmore and Carruthers, 1979, S. Afr. Frogs: 240; Channing, 2001, Amph. Cent. S. Afr.: 144).

Sedge Frogs (Ananjeva, Borkin, Darevsky, and Orlov, 1988, Dict. Amph. Rept. Five Languages: 71).

Lily Frogs (Passmore and Carruthers, 1978, J. Herpetol. Assoc. Afr., 19: 7; Passmore and Carruthers, 1979, S. Afr. Frogs: 240).

African Reed Frogs (Frank and Ramus, 1995, Compl. Guide Scient. Common Names Amph. Rept. World: 66).

Gulf Frogs (Nesionixalus [no longer recognized]: Frank and Ramus, 1995, Compl. Guide Scient. Common Names Amph. Rept. World: 69).

Koehler's Green Frogs (Chlorolius [no longer recognized]:Frank and Ramus, 1995, Compl. Guide Scient. Common Names Amph. Rept. World: 66).

Egg-guarding Frogs (Alexteroon [no longer recognized]: Channing and Rödel, 2019, Field Guide Frogs & Other Amph. Afr.: 168).

Distribution

Savanna, farmbush, and forests of Africa, south of the Sahara.

Comment

See Poynton and Broadley, 1987, Ann. Natal Mus., 28: 193–227, for discussion of south-central African species, and Lambiris, 1988, Lammergeyer, 39: 134–147, for species of KwaZulu-Natal, Rep. South Africa and a discussion of taxonomic problems in the genus. Schiøtz, 1999, Treefrogs Afr., provided accounts, keys, maps, and photographs of most species and discussed species groups, especially the very difficult Hyperolius viridiflavus complex. Wieczorek, Drewes, and Channing, 2000, J. Biogeograph., 27: 1231–1243, provided a molecular phylogeny of members of Hyperolius and redefined species within the Hyperolius viridiflavus superspecies. Wieczorek, Drewes, and Channing, 2001, Amphibia-Reptilia, 22: 155–166, reported on phylogenetics and taxonomy of the Hyperolius viridiflavus complex. Channing, 2001, Amph. Cent. S. Afr.: 144–185, and Du Preez and Carruthers, 2009, Compl. Guide Frogs S. Afr.: 236–271,provided keys and accounts for all species of southern Africa. Wieczorek, Channing, and Drewes, 1998, Herpetol. J., 8: 29–34, discussed the history and varying classifications of the Hyperolius viridiflavus complex. Amiet, 2005, Rev. Suisse Zool., 112: 271–310, dicussed taxonomic problems in the Hyperolius nasutus complex and made a number of taxonomic changes. Nesionixalus considered a synonym of Hyperolius by Drewes, 1984, Occas. Pap. California Acad. Sci., 139: 52, who considered it, therefore to be in Hyperoliinae. Generic status revalidated by Perret, 1988, Bull. Soc. Neuchatel. Sci. Nat., 111: 35–48, who considered it to be in Hyperoliinae. Previously, Dubois, 1981, Monit. Zool. Ital., N.S., Suppl., 15: 260, placed Nesionixalus in Leptopelinae (now part of Arthroleptidae). Schiøtz, 1999, Treefrogs Afr.: 309 (who provided accounts of most of the species recognized at the time), and Drewes, 2000, Copeia, 2000: 624–626, discussed the controversy surrounding the separation of Nesionixalus from Hyperolius. Mercurio, 2011, Amph. Malawi: 153–211, provided accounts and an identification key for the species of Malawi. Pyron and Wiens, 2011, Mol. Phylogenet. Evol., 61: 543–583, provided an estimate of phylogeny of a number of exemplar species. See accounts for Cameroon species by Amiet, 2012, Rainettes Cameroun: 166–371. Channing, Rödel, and Channing, 2012, Tadpoles of Africa: 201–228, reported on comparative tadpole morphology. Loader, Lawson, Portik, and Menegon, 2015, BMC Res. Notes, 8 (Art. 167): 1–16, revised and provided a key to the species of the Hyperolius spinigularis group of East Africa. Gilbert and Bell, 2017 "2018", Biol. J. Linn. Soc., 123: 1–11, compared call structure of the Hyperolius thomesnsi/Hyperolius moleri/Hyperolius drewesi/Hyperolius olivaceus group in Sao Tome and Principe and Central Africa.  Bell, Parra, Badjedjea, Barej, Blackburn, Burger, Channing, Dehling, Greenbaum, Gvoždík, Kielgast, Kasumba, Lötters, McLaughlin, Nagy, Rödel, Portik, Stuart, VanDerWal, Zassi-Boulou, and Zamudio, 2017, Mol. Ecol., 26: 5224–5244, discussed molecular biogeography, phylogenetics, and systematics of Hyperolius species of west-central Africa. Dehling and Sinsch, 2019, Zool. Anz., 280: 65–77, discussed the evolution of morphspace of species found in the Central African Albertine Rift. Lawson, Liedtke, Menegon, and Loader, 2018, Herpetol. Notes, 11: 873–879, summarized clutch size, egg diameter, male and female adult SVL. Channing and Rödel, 2019, Field Guide Frogs & Other Amph. Afr.: 170–209, provided brief accounts, photographs, and range maps for the species. Portik, Bell, Blackburn, Bauer, Barratt, Branch, Burger, Channing, Colston, Conradie, Dehling, Drewes, Ernst, Greenbaum, Gvoždík, Harvey, Hillers, Hirschfeld, Jongsma, Kielgast, Kouete, Lawson, Leaché, Loader, Lötters, van der Meijden, Menegon, Müller, Nagy, Ofori-Boateng, Ohler, Papenfuss, Rößler, Sinsch, Rödel, Veith, Vindum, Zassi-Boulou, and McGuire, 2019, Syst. Biol., 68: 859–875, provided a relatively dense estimate of phylogeny within the genus. Ernst, Kehlmaier, Baptista, Vaz Pinto, Branquima, Dewynter, Fouquet, Ohler, and Schmitz, 2021, Zool. Anz., 293: 215–224, reported on mitogenomics of the Hyperolius with special reference to former Alexteroon, which they discussed with reference to morphology and molecular placement of the taxa as well as their ranges. As previously found Alexteroon was recovered as nested within Hyperolius. However, they retained Alexteroon as a subgenus and noted that Hyperolius sensu lato will presumably be broken into several genera as the phylogenetics and species boundaries are better formulated. Channing, 2022, Zootaxa, 5134: 301–354, provided useful rearrangement of the Hyperolius viridiflavus complex, which included an mtDNA analysis of the members, summaries of the myriad color patterns and calls, a discussion of taxonomic history, reidentification of GenBank sequences, and a pragmatic taxonomic summary which, while it will not be the last word on species boundaries, is a giant step forward in bringing questions into focus as this giant puzzle is resolved. 

Contained taxa (143 sp.):

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