Hyperolius angolensis Steindachner, 1867

Santos, Bruna S., Marques, Mariana P., Bauer, Aaron M. & Ceríaco, Luis M. P., 2021, Herpetological results of Francisco Newton’s Zoological Expedition to Angola (1903 – 1906): a taxonomic revision and new records of a forgotten collection, Zootaxa 5028 (1), pp. 1-80 : 12-13

Hyperolius angolensis Steindachner, 1867

Material (twenty-nine specimens): Six adult specimens (MHNCUP/ ANF 134–139, formerly UP-MHNFCP- 094966 and UP-MHNFCP-094967), collected from Rio Quanza (impossible to georeference) undetermined province, in 1903; one adult male specimen (MHNCUP/ ANF 173 , formerly UP-MHNFCP-017304), collected from N’Golla Bumba [-9.03333º, 14.60000º, 460 m a.s.l.] Kwanza Norte Province, in 1904 ; one adult specimen (MHNCUP/ ANF 177 , formerly UP-MHNFCP-017311), collected from Golungo Alto [-9.13333º, 14.76667º, 666 m a.s.l.] Kwanza Norte Province, in 1904 ; two adult specimens (MHNCUP/ ANF 180-181 , formerly UP-MHNFCP- 017295), collected from Cabiri [-8.92125º, 13.66654º, 31 m a.s.l.] Bengo Province, in July 1903 ; one adult specimen (MHNCUP/ ANF 182 , formerly UP-MHNFCP-094961), collected from Quilombo [-9.33333º, 14.90000º, 744 m a.s.l.] Kwanza Norte Province, in 1904 ; one adult male specimen (MHNCUP/ ANF 183 , formerly UP-MHNFCP- 017293), collected from Palmyra, Cazengo [-9.30000º, 14.68333º, 403 m a.s.l.] Kwanza Norte Province, in 1903 ; one adult specimen (MHNCUP/ ANF 199 , formerly UP-MHNFCP-094963), collected from Cambondo [-9.16349º, 14.66432º, 394 m a.s.l.] Kwanza Norte Province, date unknown ; four adult specimens (MHNCUP/ ANF 200–203 , formerly UP-MHNFCP-094963), collected from unknown locality, date unknown ; one adult specimen (MHNCUP/ ANF 204 , formerly UP-MHNFCP-094964), collected from unknown locality, in 1904 ; one adult specimen (MHNCUP/ ANF 295 , formerly UP-MHNFCP-017317), collected from unknown locality, date unknown ; two adult specimens (MHNCUP/ ANF 296–297 , formerly UP-MHNFCP-017300; Fig. S8 ), collected from unknown locality, date unknown; Three adult specimens (MHNCUP/ ANF 300–302 , formerly UP-MHNFCP-017297 and UP-MHNFCP-017299), collected from unknown locality, date unknown ; five adult specimens (MHNCUP/ ANF 336–337 , ANF 340–341 , ANF 343 ), collected from unknown locality, date unknown .

Comments: Hyperolius angolensis is one of the most widespread species of reed frogs in Angola, but also one with most complicated taxonomic and nomenclatural history ( Marques et al. 2018). The taxonomic identification of the Angolan specimens is currently subject to three different main interpretations. Authors including Wieczorek et al. (2000), Frétey et al. (2011), Baptista et al. (2019) and Frost (2020) consider the Angolan population as representatives of Hyperolius paralellus Günther, 1858 . Some other authors, as Conradie et al. (2012), Ceríaco et al. (2016b), Marques et al. (2018) or Butler et al. (2019) consider the Angolan populations as members of H. angolensis , a species described from Duque de Brangaça [=Kalandula] region in Malanje Province, by Steindachner in 1867. The decision to use H. angolensis instead of the older nomen parallelus is justified by an apparent homonymy of the paralellus type material, constituted by specimens from South Africa and Angola. The original description of parallelus referred only to material from South Africa (Günther, 1858), and only in a subsequent publication were Angolan specimens noted (Günther “1858” 1859; “ Cape of Good Hope” and “Ambris, Angola ” [= Ambriz] fide Boulenger (1882)). Most recently Channing & Rödel (2019) considered the Angolan population as part of a widely distributed H. viridiflavus , referring to the various populations across the continent, noting that “the advertisement call is very similar and the genetic variation very small between populations”. Without more data, we opt to follow Marques et al. (2018). The authors noted, however, that under H. angolensis there might be several different biological species, given the apparent correspondence of different coloration patterns to specific and distinct geographical distributions ( Marques et al. 2018).

Subsequent to the description of both parallelus and angolensis, Bocage described several species, subspecies and varieties from Angola, namely Hyperolius toulsonii from “Loanda” [= Luanda] ( Bocage 1867), Hyperolius insignis from “Benguella” ( Bocage 1867) (corrected later by Perret (1976a) to “St. Salvador du Congo ” [= M’Banza Congo] and “Novo Redondo” [=Sumbe]), Hyperolius huillensis from “Huilla” ( Bocage 1873), Rappia plicifera from “Caconda” and “Duque de Bragança” ( Bocage 1893b), Rappia marmorata var. marginata from “Duque de Bragança” ( Bocage 1895), Rappia marmorata var. taeniolata from “Duque de Bragança”, “Huilla”, “Caconda” and “Cahata” [= Calandula, Huila, Caconda and Caota] ( Bocage 1895) and Rappia marmorata var. variegata from “Cahata” and “Quindumbo” [= Caota and Quindumbo] ( Bocage 1895). Other authors, such as Peters, Ahl and Monard described Hyperolius vermiculatus from “Malange” [= Malanje] ( Peters 1882a), Hyperolius marungaensis from “Marunga” ( Ahl 1931), Hyperolius decoratus from “Longa” ( Ahl 1931), Hyperolius microtictus from “Longa” ( Ahl 1931) and Hyperolius erythromelanus from “Sangevé” [= Sangueve] ( Monard 1937). The aforementioned have all been synonymized with either H. marmoratus , a species originally described from “Natal” [=KwaZulu- Natal Province], South Africa by Rapp (1842), H. paralellus or H. angolensis (see Marques et al. 2018).

Ferreira (1906) cited an adult male specimen of “ Rappia plicifera ” from “N’Golla Bumba” and one “ Rappia toulsoni ” from “Golungo Alto”, matching the extant specimens MHNCUP/ANF 173 ( Fig. S9 ) and MHNCUP/ ANF 177 ( Fig. S10 ), respectively. Ferreira (1906) also cited two specimens of “ Rappia marmorata var. insignis ” from “Cambondo” and “Quilombo”, of which MHNCUP/ANF 199 matches the former. Specimens MHNCUP/ ANF 200–203 ( Fig. S11 ), do not have locality data, but are identified on their respective historical labels as “ R. marmorata insignis ” and possibly correspond to the cited material from “Quilombo”. We were unable to find matches for some specimens cited by Ferreira (1904), namely the adults of “ Rappia plicifera ” from “Cazengo?”, “Cabiri” and “Cacuaco (Bengo)” and adults and juveniles of “ Rappia marmorata ” from “Gumba” and “Cazengo”. It is possible that among the specimens without information (lacking either locality, date of collection, previous identification or a combination of those – MHNCUP/ANF 295–297, ANF 300–302, ANF 336–337, ANF 340–341, ANF 343) are the matches for the remaining cited specimens. Additionally, extant in MHNC-UP are specimens with the following previous identifications (made by Ferreira) – “ Rappia marmorata ” (MHNCUP/ANF 182, Fig. S12 ), “ Rappia marmorata paralella ” (MHNCUP/ANF 136–139, ANF 180–181, Fig. S13 ), “ Rappia marmorata var insignis ” (MHNCUP/ANF 183), “ Rappia marmorata plicifera ” (MHNCUP/ANF 204, Fig. S14 ) and “ Rappia plicifera ” (MHNCUP/ANF 134–135).

As Bethencourt Ferreira had access to Bocage’s original type specimens while working in Lisbon, we consider that the Newton specimens are representatives of the most approximate interpretation of the lost type material, and therefore, can be of crucial importance for future taxonomic revisions of the group .

Figures

FIGURE 8. Life photo of Meroles reticulatus from Baía dos Tigres. Photo by Luis Ceríaco.

FIGURE S9. Specimen of Hyperolius angolensis (originally identified and/or labelled as Rappia plicifera) (MHNCUP/ANF 173), from N’Golla Bumba. Photo by Bruna Santos.

FIGURE S10. Specimen of Hyperolius angolensis (originally identified and/or labelled as Rappia tolsoni) (MHNCUP/ANF 177), from Golungo Alto. Photo by Bruna Santos.

FIGURE S11. Specimen of Hyperolius angolensis (originally identified and/or labelled as R. marmorata insignis) (MHNCUP/ ANF 200), from unknown locality. Photo by Bruna Santos.

FIGURE S12. Specimen of Hyperolius angolensis (originally identified and/or labelled as Rappia marmorata) (MHNCUP/ANF 182), from unknown locality. Photo by Bruna Santos.

FIGURE S13. Specimen of Hyperolius angolensis (originally identified and/or labelled as Rappia marmorata paralella) (MHNCUP/ANF 138), from unknown locality. Photo by Bruna Santos.

FIGURE S14.Specimen ofHyperolius angolensis (originally identifiedand/or labelled asRappiamarmorata plicifera) (MHNCUP/ ANF 204), from unknown locality. Photo by Bruna Santos.