Hyperolius Rapp, 1842

Marques, Mariana P., Ceríaco, Luis M. P., Blackburn, David C. & Bauer, Aaron M., 2018, Diversity and Distribution of the Amphibians and Terrestrial Reptiles of Angola Atlas of Historical and Bibliographic Records (1840 - 2017), Proceedings of the California Academy of Sciences 65, pp. 1-501 : 89-115

Genus Hyperolius Rapp, 1842

Hyperolius adspersus Peters, 1877 SPRINKLED LONG REED FROG

Hyperolius adspersus Peters 1877a:619 , pl., figs. 6-6a. Holotype: ZMB 9176 fide Laurent (1961:92). Type locality: “Chinchoxo” Cabinda Province, Angola.

Rappia nobrei Ferreira 1904:112 . Syntypes: MHNFCP 017292 [2 specimens] (collector F. Newton). Type locality: “Cabiri” Bengo Province, Angola. Considered a junior synonym of H. adspersus by Ceríaco et al. (2014a:21).

Hyperolius nasutus adspersus : Laurent (1961:92).

Hyperolius granulatus : Laurent (1964a:155), Cei (1977:17).

Hyperolius adspersus : Amiet (2005:274), Frétey et al. (2011:29), Channing et al. (2013b:314), Frost (2016).

Hyperolius cf. adspersus : Ceríaco et al. (2014a:21).

Global conservation status (IUCN): Least Concern.

Global distribution: The species is known from southern Cameroon to the lower Congo Basin in Democratic Republic of Congo to northern Angola , including the Cabinda encalve. Ocurrences in Angola (Map 32): The species is known from the type locality “Chinchoxo” and from scattered localities in northeastern Angola. Cabinda: “Chinchoxo”

[-5.10000, 12.10000] ( Peters 1877a:619; Laurent 1961:92; Amiet 2005:275; Channing et al.

2013b:317; Frost 2016). Bengo: “Cabiri”

[-8.91667, 13.66667] ( Ferreira 1904:112;

Ceríaco et al. 2014a:21). Lunda Sul: “Alto

Chicapa galerie forestière des sources du ruisseau Cuílo” [-10.88333, 19.23333] ( Laurent

1964a:155). Moxico: “Réserve de chasse de

Cameia, 120 km à l’est de Luso” [-11.83333,

21.00000] ( Laurent 1964a:155); “Rives du lac

Calundo” [-11.80000, 20.86667] ( Laurent

1964a:155).

Taxonomic and distributional notes:

This species was described by Peters (1877a) MAP 32. Distribution of Hyperolius adspersus in Angola. as Hyperolius adspersus based on a specimen collected in “Chinchoxo,” in the Cabinda enclave. Ferreira (1904) described Rappia nobrei from the type locality “Cabiri” collected by Francisco Newton. This taxon was previously considered as incertae sedis in part because of the long lack of study of the type specimens ( Frost 2016). A recent study by Ceríaco et al. (2014a) confirmed that this taxon is clearly a member of the genus Hyperolius and is plausibly a junior synonym of H. adspersus . Previously, H. adspersus was considered to be a synonym of H. nasutus Günther 1865 , but Amiet (2005) removed it from synonymy. This species is part of the taxonomically problematic Hyperolius nasutus group, which currently contains 16 recognized species (Channing et al. 2013b; Ceríaco et al. 2014a). Of these 16 species, only three are known to occur near the type locality of Rappia nobrei : Hyperolius adspersus , Hyperolius nasutus Günther, 1865 , and H. benguellensis ( Bocage, 1893) (Channing et al. 2013b; Ceríaco et al. 2014a; see H. benguellensis and H. nasutus accounts). The syntypes of R. nobrei are similar to both H. adspersus and H. benguellensis , which makes it difficult to determine to which species Rappia nobrei should be referred. We here follow Ceríaco et al. (2014a) by considering R. nobrei to be a junior synonym of H. adspersus . Channing et al. (2013b) considered Hyperolius granulatus ( Boulenger, 1901) , previously associated with Hyperolius benguellensis ( Bocage, 1893) , to be a synonym of H. adspersus (Perret 1976a; Schiøtz and Van Daele 2003).

Hyperolius angolensis Steindachner, 1867 ANGOLAN REED FROG

[Note: Because of the complexity of the taxonomy of this species we provide additional commentary on its synonymy/chresonymy]

Hyperolius marmoratus Rapp 1842:289 , pl. 6, figs. 1–2. Type: Not designated (collector unknown) . Type locality: “ Natal ” [= KwaZulu-Natal], South Africa .

Hyperolius parallelus Günther 1858a:326 . Syntypes: BMNH 1947.2.9.41–43 [3 specimens] (collector unknown, probably “Mr. Rich”). Specimens BMNH 1947.2.9.41 and 1947.2.9.42 are depicted in Günther “1858b” 1859: 86, pl. 8, fig. A. Type locality: “Süd-Afrika” later changed to “ South Africa ” and “ Angola ” (Günther “1858b” 1859:86) and to “Cape of Good Hope” and “Ambris, Angola ” [= Ambriz] by Boulenger (1882:121). Synonymized with H. marmoratus by Boulenger (1882:121) and treated as a subspecies by Laurent (1951:38).

Hyperolius marmoratus var. angolensis Steindachner 1867:50 . Syntypes: NHMW 13487.1–2 fide Häupl and Tiedemann (1978:22) (collector F.A.P. Bayão). Type locality: “ Angola ” ( Steindachner 1867:50). Restricted to “Duque de Bragança” [= Calandula] (Ceríaco et al. 2014b:669).

Hyperolius insignis Bocage 1867a:844 , fig. 2. Syntypes: MBL T.21-164, 27-167 [2 specimens] (collector J. A. d’Anchieta [Benguella] and P. Barroso [St. Salvador du Congo]), destroyed by fire 18 March 1978. Type locality: “Benguella,” Angola. Corrected by Perret (1976a: 28) to “St. Salvador du Congo” [= M’Banza Congo] and “Novo Redondo” [= Sumbe], Angola. Synonymized with H. marmoratus by Boulenger (1882:121) and Perret (1976a:27) and treated as a subspecies by Laurent (1951 “ 1952 ”:39). Synonymized with H. parallelus by Loveridge (1953a:329).

Hyperolius Toulsonii Bocage 1867a:845 , fig. 3. Holotype: MBL T.27-275 (collector M. Toulson), destroyed by fire 18 March 1978. Type locality: “Loanda” [= Luanda], Angola. Synonymized with H. marmoratus by Boulenger (1882: 121), recognized as valid by Ahl (1931b:388), and considered a synonym of H. m. insignis by Laurent (1952 “ 1951 ”:391), and later synonymized with H. parallelus (Perret 1976a:27).

Hyperolius huillensis Bocage 1873b:225 . Syntypes: MBL 2-166 [4 specimens] (collector J. A. d’Anchieta), destroyed by fire 18 March 1978. Type locality: “Huilla,” [ Angola] ( Bocage 1873b:225). Considered a subspecies of H. marmoratus by Laurent (1952 “ 1951 ”:390).

Hyperolius vermiculatus Peters 1882a:8 . Lectotype: ZMB 10050, designated by Laurent (1961:88) (collector von F.W.A. von Mechow). Type locality: “Malange” [= Malanje] ( Peters 1882a:8). Considered a subspecies of H. marmoratus by Laurent (1952 “ 1951 ”:88).

Rappia plicifera Bocage 1893:118 . Syntypes: MBL T.22-209 [2 specimens] (collector J. A. d’Anchieta [Caconda] and F. A. P. Bayão [Duque de Bragança]), destroyed by fire 18 March 1978. Häupl and Tiedemann (1978:29) considered NHMW 22895 [2 specimens] as part of the type series. Type locality: “Caconda,” “Duque de Bragança,” Angola ( Bocage 1893:118). Synonymized with H. m. vermiculatus by Laurent (1952 “ 1951 ”:390) and later with H. parallelus by Perret (1976a:27).

Rappia marmorata var. marginata : Bocage 1895a:164: Type: MBL, specimen number(s) not known (collector F. A. P. Bayão), destroyed by fire 18 March 1978. Type locality: “ Duque de Bragança ” [= Calandula], Angola.

Rappia marmorata var. taeniolata Bocage 1895a:164 : Syntypes: MBL, specimen numbers not known (collector F. A. P. Bayão [ Duque de Bragança ], Graça and J. A. d’Anchieta [Huilla] and J. A. d’Anchieta [Caconda and Cahata]), destroyed by fire 18 March 1978. Type locality: “Duque de Bragança,” “Huilla,” “Caconda” and “Cahata” [= Calandula, Huila, Caconda and Caota], Angola.

Rappia marmorata var. variegata Bocage 1895a:164 : Syntypes: MBL (collector J. A. d’Anchieta). Type locality: “Cahata” and “Quindumbo” [= Caota and Quindumbo], Benguela Province, Angola.

Hyperolius marungaensis 1931b:351, fig. 226. Holotype: ZMB uncatalogued (collector Böhm). Type locality: “Marunga,” Angola. Considered a subspecies of H. marmoratus by Laurent (1961:88)

Hyperolius decoratus 1931b:352, fig. 227. Holotype: ZMB uncatalogued, MCZ A-17632 (collectors Brühl and Gleim) – on exchange from ZMB fide Barbour and Loveridge (1946:127). Type locality: “Longa” ( Ahl 1931a:78, 1931b:352), Angola. Considered a subspecies of H. marmoratus by Laurent (1952 “ 1951 ”:392).

Hyperolius microtictus 1931b:353, fig. 228. Syntypes: ZMB [2 specimens] uncatalogued, ZMB 36100 [2 specimens] (collector unknown). Type locality: “Longa” ( Ahl 1931a:80, 1931b:352), Angola. Considered a synonym of H. m. decoratus by Laurent (1961:89).

Hyperolius erythromelanus Monard 1937a:36 . Syntypes: LMCFM [2 specimens] (collector A. Monard). Type locality: “Sangevé” ( Monard 1937a:36), [= Sangueve], Angola. Synonymized with H. m. huillensis by Laurent (1952 “ 1951 ”:390).

Hyperolius marmoratus alborufus : Laurent 1964a:153, fig. 40. Holotype: MD 5679 (collector unknown). Type locality: “Cazombo, Alto Zambeze, Moxico, Angola ” ( Laurent 1964a:153).

Hyperolius huillensis : Bocage (1879c:89).

Hyperolius insignis : Bocage (1867a:844, 1887a:191).

Rappia insignis : Günther (1869:479).

Hyperolius marmoratus : Günther (1864 a:480), Bocage (1866a:55, 1886b:74), Peters (1881:150), Noble (1924:253), Schmidt (1936:131), Mertens (1938a:427), Inger (1959:541), Gavetti and Andreone (1993:103), Ceríaco et al. (2014b:669).

Hyperolius citrinus : Bocage (1879c:89).

Hyperolius Toulsinii : Bocage (1895a:166, 1897a:203), Ferreira (1906:161).

Rappia marmorata : Boulenger (1882:121, 1905:109), Bocage (1895a:164, 1896a:113, 1897b:211), Ferreira (1904:112, 1906:160)

Rappia plicifera : Bocage (1895a:167, 1897a:203), Ferreira (1897b:241, 1904:112, 1906:161), Häupl and Tiedemann (1978:29), Häupl et al. (1994:34).

Rana marmorata var. huillensis : Ferreira (1897b:241).

Hyperolius decoratus : Loveridge (1936a:106), Monard (1937a:35, 1938:89); Barbour and Loveridge (1946:127).

Hyperolius vermiculatus : Monard (1938:88).

Hyperolius microstictus : Monard (1937a:35, 1938:90).

Hyperolius sp. II (ap. angolensis ): Monard (1937a:38, 1938:92).

Hyperolius sp. III (ap. angolensis ): Monard (1937a:38, 1938:93).

Hyperolius sp. I (ap. decoratus ): Monard (1937a:37, 1938:92).

Hyperolius angolensis : Monard (1937a:36, 1938:90), Channing (2001:148), Conradie et al. (2012a:2), Ceríaco et al. (2016b:31), Conradie et al. (2016:14) .

Hyperolius graueri : Mertens (1937a:20).

Hyperolius parallelus : Peters (1877a:618), Laurent (1943a:14), Ceríaco et al. (2014b:669), Frost (2016).

Hyperolius marmoratus angolensis : Laurent (1950a:17, 1954a:80), Frade (1963:254), Laurent (1964a:152), Cei (1977:17), Broadley (1965a:26), Häupl and Tiedemann (1978:22).

Hyperolius marmoratus albofasciatus : Loveridge (1953a:350).

Hyperolius marmoratus parallelus : Laurent (1961:89), Cei (1977:17).

Hyperolius marmoratus huillensis : Laurent (1961:88), Cei (1977:17), Poynton and Haacke (1993:15).

Hyperolius marmoratus vermiculatus : Laurent (1961:88).

Hyperolius marmoratus marungaensis : Laurent (1961:88), Cei (1977:17).

Hyperolius marmoratus insignis : Laurent (1961:89, 1964a:151) Cei (1977:17).

Hyperolius marmoratus alborufus : Laurent (1964a:153), Cei (1977:17).

Hyperolius parallelus alborufus : Schiøtz (1975:185).

Hyperolius parallelus pliciferus : Perret (1976a:27).

Hyperolius parallelus toulsoni : Perret (1976a:27).

Hyperolius parallelus insignis : Perret (1976a:29).

Hyperolius parallelus huillensis : Perret (1976a:29).

Hyperolius erythromelanus : Frost (1985:210).

Hyperolius parallelus-marginatus subgroup: Schiøtz (1999:217).

Hyperolius (Hyperolius) marmoratus : Frétey et al. (2011:32).

Hyperolius (Hyperolius) parallelus : Frétey et al. (2011:32).

Global conservation status (IUCN): Least Concern.

Global distribution: The species is known from Angola and the adjacent countries of Namiba, Democratic Republic of Congo, and Zambia.

Ocurrences in Angola (Map 33): The species is very widespread for almost the entire territory . Cabinda: “ Chinchoxo ” [-5.10000,

12.10000] ( Peters 1877a:618; Bocage

1895a:164; Laurent 1961:89). “Landana”

[-5.21667, 12.15000] ( Laurent 1943a:14).

Zaire: “ St. Salvador du Congo ” [-6.26667,

14.23333] ( Bocage 1887a:191, 1895a:164;

Perret 1976a:29). Luanda: “Loanda”

[-8.83333, 13.26667] ( Bocage 1867a:845,

1895a:166, 1897b:203; Loveridge 1936a:106,

1953a:352; Laurent 1961:88, Perret 1976a:27);

“Longa, Loanda” ( Ahl 1931a:78, 1931b:352;

Barbour and Loveridge 1946:127). Bengo:

“Ambriz” [-7.844312, 13.106493] ( Boulenger

1882:121; Loveridge 1953a:352). Malanje:

“ Tembo Aluma” [-7.70000, 17.28333] ( Boulenger 1905:109); “Bange N’gola” [-8.43333, MAP 33. Distribution of Hyperolius angolensis in Angola.

16.56667] ( Boulenger 1905:109); “Duque de

Bragança ” [-9.10000, 15.95000] ( Bocage 1866a:55, 1866b:74, 1893:118, 1895a:164, 167, 1897a:203; Boulenger 1882:121); “ Malanje ” [-9.55000, 16.35000] ( Peters 1882a:8, Laurent 1961:88); “Capanda” [-9.72841, 15.34585] (Ceríaco et al. 2014b:669); “Cangandala National Park” [-9.84606, 16.72233] (Ceríaco et al. 2016b:31). Lunda Norte: “Dundo” [-7.36667, 20.83333] ( Laurent 1950a:17, 1954a:80, 1964a:152); “Matala rive gauche de la Thshiumbe, 40 km à l’est de Dundo” [-7.43333, 21.16667] ( Laurent 1950a:17); “Carumbo lagoon” [-7.74422, 19.95467] (Conradie et al. 2012a:2); “Muita (Luembe E)” [-7.80000, 21.45000] ( Laurent 1950a:17, 1954a:80); “Andrada (Luembe O)” [-7.68729, 21.37003] ( Laurent 1954a:80); “Caluango, Riv. Camaloa, affl. Caluango” [-8.33333, 19.65000] ( Laurent 1964a:152); “Sombo, riv. Melanda, affl. Chiumbe” [-8.68333, 20.95000] ( Laurent 1954a:80, 1964a:152). Lunda Sul: “Alto Cuílo, mare Tchifuka” [-10.00000, 19.58333] ( Laurent 1964a:152). Moxico: “rives de Lumeje, près du lac Calundo” [-11.51667, 20.76667] ( Laurent 1964a:152); “rives du lac Calundo, 105 km à l’est de Luso” [-11.80000, 20.86667] ( Laurent 1964a:152); “Réserve de chasse de Cameia, 120 km à l’est de Luso” [-11.83333, 21.00000] ( Laurent 1964a:152); “Cazombo (High Zambèze)” [-11.88333, 22.91667] ( Laurent 1964a:153; Schiøtz 1975:185). Kwanza Norte: “N’golla Bumba” [-9.03333, 14.60000] ( Ferreira 1906:161); “Golungo Alto” [-9.13333, 14.76667] ( Boulenger 1905:109; Ferreira 1906:161); “Cambondo” [-9.15963, 14.65827] ( Ferreira 1906:160); “Canhoca” [-9.25000, 14.68333] ( Boulenger 1905:109); “Cazengo” [-9.33333, 14.76667] ( Ferreira 1904:112); “Quilombo” [-9.33333, 14.90000] ( Ferreira 1906:160). Kwanza Sul: “Novo Redondo Sumbe” [-11.20000, 13.85000] ( Bocage 1895a:164; Perret 1976a:29); “Gumba” [-11.26667, 14.28333] ( Ferreira

1904:112). Huambo: “Bimbi” [-11.81667, 15.83333] ( Monard 1937a:35-36, 38; 1938:89, 90, 92). Bié: “Gauca” [-11.18333, 17.45000] ( Schmidt 1936:131); “Chitau” [-11.43333, 17.15000] ( Schmidt 1936:131); “Goedecke Farm, Conjo, 25 km north Genera Machado (Camacopa)” [-12.01667, 17.06667] ( Mertens 1937a:20); “Bihé” [-12.38333, 16.95000] ( Bocage 1879c:89, 1895a:164); “Cubango basin (10)” [-13.71616, 17.09661] ( Conradie et al. 2016:8-9, 14); “Cuando basin (12a)” [-13.59333, 16.87986] ( Conradie et al. 2016:8-9, 14); “Cuando basin (16)” ( Conradie et al. 2016:8-9, 14). Benguela: “Cahata” [-12.35000, 14.81667] ( Bocage 1895a:164); “Quindumbo” [-12.46667, 14.93333] ( Bocage 1895a:164); “Benguella” [-12.58333, 13.41667] ( Bocage 1867a:844, 1887a:191; Günther 1869:479; Boulenger 1882:121; Loveridge 1936a:106, 1953a:352; Laurent 1961:89); “Ebanga” [-12.73333, 14.73333] ( Monard 1937a:35, 37; 1938:89, 92); “Dombe” [-12.95000, 13.10000] ( Bocage 1895a:164; Laurent 1961:89; Gevetti and Andreone 1993:104); “Cubal” [-13.03333, 14.25000] (Mertens 1938:427); “Marco de Canavezes (Cubal da Ganda)” [-13.08333, 14.33333] ( Laurent 1964a:151); “Hanha” [-13.30000, 14.20000] ( Bocage 1867a:844, 1896a:113, 1897b:211; Boulenger 1882:121); “Tongrube neben dem Jamba-Fluβ, Entre Rios” [-13.60000, 16.60000] ( Hellmich 1957a:28). Huíla: “ 12 km W of Bela Vista” [-12.56667, 16.21667] (Poynton and Haacke 1993:15); “Rio Cuce” [-13.51667, 15.20000] ( Ferreira 1897b:241); “Caconda” [-13.73333, 15.06667] ( Bocage 1893:118, 1895a:164, 167, 1897a:203; Perret 1976a:27; Häupl and Tiedemann 1978:29; Häupl et al. 1994:34; Gevetti and Andreone 1993:104); “Kalukembé” [-13.78333, 14.68333] ( Monard 1937a:35, 38; 1938:92); “Sangevé” [-13.88333, 15.83333] ( Monard 1937a:35; 1938:89, 91; Frost 1985:210); “Kuvangu” [-14.46667, 16.30000] ( Monard 1937a:35-38; 1938:90, 93); “Kapelongo” [-14.88333, 15.08333] ( Monard 1937a:36, 1938:90); “Humpata” [-14.93333, 13.51667] (Conradie et al. 2012a:2); “Huilla” [- 15.05000, 13.55000] ( Günther 1865a:480; Bocage 1873b:225, 1895a:164; Boulenger 1882:121, Laurent 1961:88; Perret 1976a:29); “Nuntechite lagoon” [-15.13333, 13.41667] (Poynton and Haacke 1993:15). Cuando Cubango: “Marunga oder Kawende” [-17.45000, 20.03333] ( Ahl 1931a:77, 1931b:351; Laurent 1961:88); “Kuandu” [-16.74487, 19.10136] ( Monard 1937a:36; 1938:90); “Cubango basin (6b)” [-14.67458, 17.73544] ( Conradie et al. 2016:8-9, 14); “(7c)” [-14.43916, 17.81491] ( Conradie et al. 2016:8-9, 14); “Cuando basin (19)” [-14.70213, 17.37772] ( Conradie et al. 2016:8-9, 14); “Cuando basin (21)” [-14.66586, 17.07661] ( Conradie et al. 2016:8- 9, 14); “Cuando basin (22a)” [-14.66622, 16.97842] ( Conradie et al. 2016:8-9, 14); “Cuito basin (23)” [-14.58972, 18.17083] ( Conradie et al. 2016:8-9, 14); “Cuito basin (24)” [-14.60622, 18.46722] ( Conradie et al. 2016:8-9, 14); “Cuito basin (24a)” ( Conradie et al. 2016:8-9, 14); “Cuito basin (26)” [-15.08686, 19.14872] ( Conradie et al. 2016:8-9, 14); “ Cuito basin (27)” [-15.17127, 19.19433] ( Conradie et al. 2016:8-9, 14); “Cuito basin (28)” [-15.13486, 19.19636] ( Conradie et al. 2016:8-9, 14); “Cubango basin (29)” [-17.87291, 19.83333] ( Conradie et al. 2016:9-10, 14); “Cuito basin (30a)” [-17.50875, 20.06594] ( Conradie et al. 2016:9-10, 14); “Cuito basin (30b)” [-17.51194, 20.04305] ( Conradie et al. 2016:9-10, 14); “Cuito basin (30d)” [-17.51327, 20.06111] ( Conradie et al. 2016:9-10, 14); “Cuito basin (32)” [-17.04880, 19.53333] ( Conradie et al. 2016:9- 10, 14); “Cuito basin (35)” [-16.62322, 19.05352] ( Conradie et al. 2016:9-10, 14); “Cuando basin (41b)” [-17.46777, 23.07944] ( Conradie et al. 2016:9-10, 14); “Cuando basin (43)” [-17.53500, 23.18916] ( Conradie et al. 2016:9-10, 14); “Cuando basin (44a)” [-17.57333, 23.26000] ( Conradie et al. 2016:9-10, 14); “Cubango basin (45)” [-16.88350, 18.01180] ( Conradie et al. 2016:9, 12, 14); “Cuito basin (51)” [-14.58970, 18.1711] ( Conradie et al. 2016:9, 12, 14); “Cuito basin (53b)” [-14.56322, 18.44394] ( Conradie et al. 2016:9, 12, 14); “Cuito basin (54)” [-14.46810, 18.35488] ( Conradie et al. 2016:9, 12, 14); “Cuito basin (55)” [-14.68478, 18.67369] ( Conradie et al. 2016:9, 12, 14); “Cuito basin (57)” [-15.45969, 18.76833] ( Conradie et al. 2016:9, 12, 14); “Cuito basin (58)” [-15.38206, 19.06375] ( Conradie et al. 2016:9, 12, 14); “Cuito basin (62)” [-17.50875, 20.06608] ( Conradie et al. 2016:9, 12, 14); “Cuito basin (63)” [-17.93611, 21.10269] ( Conradie et al. 2016:9, 12, 14). Undetermined Locality: “Without precise location” ( Bocage 1887a:191, Laurent 1961:88); “north of Quanza” ( Bocage 1895a:164); “Between Benguela and Bihé” ( Boulenger 1905:109); “Locomi” ( Boulenger 1905:109); “plateaus regions” ( Cei 1977:17); “areas of forest and savanna in the north and northeast of Angola ” ( Cei 1977:17)

Taxonomic and distributional notes: This species was first described as Hyperolius angolensis Steindachner, 1867 based on a single specimen from “ Angola,” now corrected to “Duque de Brangança” [= Calandula] (Ceríaco et al. 2014b); note, however, that Häupl and Tiedemann (1978) cited two syntypes from “ Angola ” deposited in the Naturhistorsches Museum in Wein, Austria. This species has been reported from Angola by many previous authors under a variety of names. This taxon is considered to be a synonym of Hyperolius parallelus Günther, 1858 by some authors such as Wieczorek et al. (2000), Frétey et al. (2011), Ceríaco et al. (2014b), and Frost (2016). Other authors, including Schiøtz (1999), considered it part of the Hyperolius parallelus-marginatus subgroup — corresponding to one of three taxa, parallelus , insignis and angolensis — or instead a subspecies of Hyperolius marmoratus ( Rapp, 1842) as was suggested by Loveridge (1953a) and Poynton and Broadley (1987). Channing (2001) and Conradie et al. (2012a) considered Hyperolius angolensis to be a valid species. Both H. parallelus and H. marmoratus are taxonomically problematic. Poynton and Broadley (1987) regarded all the forms in southern Africa as subspecies of H. marmoratus , while Channing (1999) regarded angolensis and possibly other forms from Angola as specifically distinct from the eastern marmoratus -forms ( Schiøtz 1999). Hyperolius parallelus is represented in southern and southwestern Africa, including Angola ( Schiøtz 1999) whereas, H. marmoratus , is widespread across southern and eastern Africa and appears to be absent from Angola ( Channing 2001; Frost 2016). The nomina parallelus and marmoratus were previously reported from the country under various names associated with many different species and subspecies names ( Ferreira 1906; Loveridge 1953a; Monard 1937 a, 1938; Mertens 1938; Laurent 1964a; Perret 1976a; Frétey et al. 2011). This is explained partly by the considerable intraspecific and interspecific variation in coloration and pattern across populations (Ceríaco et al. 2014b). There are several studies in progress on the genus Hyperolius that will help to clarify species boundaries among these challenging taxa. Given the current state of knowledge, we refer all of these records to Hyperolius angolensis , though we recognize that this may contain multiple biological species.

Hyperolius benguellensis ( Bocage, 1893) BENGUELA LONG REED FROG

Rappia benguellensis Bocage 1893:119 . Syntypes: MBL 27.220-223 (collector J.A. d’Anchieta), destroyed by fire 18 March 1978. Type locality: “Cahata dans l’intérieur de Benguella” ( Bocage 1893:119), [= Caota]

Benguela Province, Angola. Rappia benguellensis : Bocage (1895a:169, 1897a:204, 1897b:211), Ferreira (1906:161). Hyperolius nasutus : Monard (1937a:39, 1938:94). Hyperolius benguellensis : Noble (1924:252), Monard (1937a:34; 1938:87), Frade (1963:254), Perret

(1976a:27), Frost (1985:207, 2016), Poynton and Broadley (1987:208), Amiet (2005:292), Schiøtz and

Van Daele (2003:128), Amiet (2005:275) Schiøtz (2006:62), Conradie et al. (2012a:2), Channing et al.

(2013:317). Hyperolius oxyrhynchus : Laurent (1950a:17). Hyperolius nasutus : Schiøtz (1999:97), Channing et al. (2002:96). Hyperolius (Hyperolius) benguellensis : Frétey et al. (2011:29).

Global conservation status (IUCN): Least Concern.

Global distribution: The species is known from Democratic Republic of Congo (former

Katanga Province), southern Angola and adjacent northern Nambia and Botswana.

Ocurrences in Angola (Map 34): The species is known from western Angola from the type locality “Cahata, Benguella,” although there are some records further north . Kwanza

Norte: “Quilombo” [-9.33333, 14.90000] ( Ferreira 1906:161). Huambo: “Bimbi”

[-11.81667, 15.83333] ( Monard 1938:87). Bié:

“ Cubango basin (11)” [-13.69413, 17.06177]

( Conradie et al. 2016:8-9, 15); “Cubango basin

(12a)” [-16.59333, 16.87986] ( Conradie et al.

2016:8-9, 15); Benguela: “Cahata (Caota)”

[-12.35000, 14.81667] ( Bocage 1893:119

1895a:169, 1897a:204; Perret 1976a:27; Frost

1985:207, 2016; Poynton and Broadley

1987:208; Channing et al. 2002:96; Amiet

2005:275; Channing et al. 2013:318); “Eban- MAP 34. Distribution of Hyperolius benguellensis in Angola.

ga” [-12.73333, 14.73333] ( Monard 1938:87);

“Hanha” [-13.30000, 14.20000] ( Bocage 1897b:211). Huíla: “Indungu” [-14.81667, 16.26667] ( Monard 1938:87); “Zootecnica Plateau, Humpata” [-14.96581, 13.34458] (Channing et al. 2013:317); “Humpata” [-14.23814, 13.43331] (Channing et al. 2013b:317); “Humpata” [-15.03333, 13.40000] (Conradie et al. 2012a:2). Cunene: “Chimporo” [-16.03333, 17.15000] ( Monard 1937a:39, 1938:94). Cuando Cubango: “Cubango basin (5)” [-14.74628, 17.66844] ( Conradie et al. 2016:8-9, 15); “Cubango basin (6b)” [-14.67458, 17.73544] ( Conradie et al. 2016:8-9, 15); “ Cubango basin (7b)” [-14.43377, 17.82957] ( Conradie et al. 2016:8-9, 15); “Cubango basin (7c)” [-14.43916, 17.81491] ( Conradie et al. 2016:8-9, 15); “Cubango basin (8)” [-14.25705, 17.77852] ( Conradie et al. 2016:8-9, 15); “Cubango basin (21)” [-14.66586, 17.07661] ( Conradie et al. 2016:8-9, 15); “Cubango basin (22b)” [-14.66278, 16.96081] ( Conradie et al. 2016:8-9, 15); “Cuito basin (27)” [-15.17127, 19.19433] ( Conradie et al. 2016:8-9, 15); “Cuito basin (30a)” [-17.50875, 20.06594] ( Conradie et al. 2016:9-10, 15); “Cuito basin (30d)” [-17.51327, 20.06111] ( Conradie et al. 2016:9-10, 15); “Cuito basin (32)” [-17.04880, 19.53333] ( Conradie et al. 2016:9-10, 15); “Cuito basin (35)” [-16.62322, 19.05352] ( Conradie et al. 2016: 9-10, 15); “Cuando basin (43)” [-17.53500, 23.18916] ( Conradie et al. 2016:9-10, 15); “Cubango basin (45)” [-16.88350, 18.01180] ( Conradie et al. 2016:9, 12, 15). Undetermined Locality: “Western subregion and Angolan highlands (Angolan coastal or watersheds)” ( Frade 1963:254). Taxonomic and distributional notes: In the description, Bocage notes that the type specimens are similar to Rappia nasuta (Günther, 1865) . In their detailed discussion of the benguelensis-nasutus complex, Poynton and Broadley (1987) had little doubt that the characters used at that time were inadequate to separate all available specimens into nasutus and benguellensis . Schiøtz (1999) concluded that it is difficult to separate H. nasutus and H. benguellensis based on morphology and color patterns. Channing et al. (2002) proposed formal changes in nomenclature and gave detailed lists of synonyms, but Schiøtz (2006) questioned several of these changes. Schiøtz and Van Daele (2003) provided a discussion about the differences in the advertisement calls of nasutus and benguellensis as well as differences in external morphology. Hyperolius benguellensis ( Bocage, 1893) was questionably removed from the synonymy of H. nasutus Günther, 1865 by Amiet (2005). Channing et al. (2013) used molecular data to argue that H. benguellensis is a valid species. The species has only been confirmed from southern Angola, northern Namibia, and northern Botswana where it was found in open grassy habitats, along streams or man-made structures with emergent vegetation (Channing et al. 2013). We follow Poynton and Broadley (1987), Schiøtz and Daele (2003), and Channing et al. (2013) in regarding H. benguellensis as distinct from the sympatric H. nasutus . Hyperolius oxyrhynchus is regarded as a synonym of H. benguellensis (Channing et al. 2013b) , though a record from “Muíta” (Laurent 1950) is probably a misidentification as it is far to the north of the rest of the distribution of H. benguellensis . Laurent’s (1950a) record should probably be referred to another species of the nasutus complex, such as H. adspersus or H. nasutus .

Hyperolius bicolor Ahl, 1931 TWO-COLORED REED FROG (Endemic)

Hyperolius bicolor Ahl 1931a:129 . Syntypes: ZMB, lost (collector K. May). Type locality: “Farenda Bango, Loanda” ( Ahl 1931a:129), [= Fazenda Bango, Luanda], Angola.

Hyperolius bicolor : Frost (1985:208, 2016).

Global conservation status (IUCN):

Data Deficient.

Global distribution: The species is known only from Angola.

Ocurrences in Angola (Map 35): The species is known only from the type locality

“Fazenda Bango, Loanda”. Luanda: “Farenda

Bango” [-9.81767, 14.78916] ( Ahl 1931a:129;

Frost 1985:208, 2016).

Taxonomic and distributional notes:

According to Laurent (in Frost 1985:208,

2016) the species is probably a synonym of

Hyperolius marmoratus insignis , a member of parallelus-marginatus subgroup ( Schiøtz

1999) [see Hyperolius angolensis account].

The validity of this taxon is questionable and difficult to address due to loss of the type specimen. MAP 35. Distribution of Hyperolius bicolor in Angola .

Hyperolius bocagei Steindachner, 1867 BOCAGE’S REED FROG

Hyperolius bocagei Steidachner 1867:51 , pl. 5, fig. 11. Holotype: NHMW 14846 fide Häupl and Tiedemann (1978:22) (Collector F.P. Bayão). Type locality: “ Angola ” (Steindachner 1897:51).

Rappia bocagei var. maculata Ferreira 1906:160 . Type: Not stated (collector F. Newton), see Ceríaco et al. (2014a:25-26) discussion. Type locality: “ Golungo Alto ”, Angola.

Rappia seabrai Ferreira 1906:163 . Holotype: (collector F. Newton), not located, probably lost. Paratype: MNHFCP 018587 (collector F. A. P. Bayão). Type locality: “Quilombo, Rio Luinha ” [= Gonguembo] Kwanza Norte, Angola.

Hyperolius bocagei : Bocage (1873b:225), Noble (1924:252), Parker (1936:143), Laurent (1950a:16, 1954a:79, 1964a:150), Cei (1977:17); Poynton and Broadley (1987:212), Häupl and Tiedemann (1978:22), Frost (1985:208, 2016), Häupl et al. (1994:26), Schiøtz (1999:188), Channing (2001:151), Schiøtz and Van Daele (2003:145), Ceríaco et al. (2014a:24).

Rappia bocagii : Boulenger (1882:126, 1905:109), Bocage (1895a:165, 1897a:203).

Rappia cinctiventris : Bocage (1895a:168), Boulenger (1905:110).

Rappia bocagei : Ferreira (1904:112, 1906:160).

Hyperolius seabrai : Noble (1924:253), Schmidt (1936:132), Monard (1937a:39, 1938:95), Cei (1977:17), Frost (1895:118).

Hyperolius cinctiventris : Noble (1924:252), Monard (1938:93), Inger (1959:541), Cei (1977:17).

Hyperolius (Hyperolius) bocagei : Frétey et al. (2011:29).

Global conservation status (IUCN): Least Concern.

Global distribution: The species is known only from the Democratic Republic of the Congo, to Tanzania, Angola and Zambia.

Ocurrences in Angola (Map 36): The species has been reported from scattered localities mainly in the westerns regions of the country. Zaire: “St. Salvador do Congo”

[-6.26667, 14.23333] ( Bocage 1895a:165,

1897a:203). Luanda: “Cacuaco?” [-8.78333,

13.36667] ( Ferreira 1904:112). Kwanza

Norte: “Canhoca” [-9.25000, 14.68333] ( Boulenger 1905:109); “Golungo Alto” [-9.13333,

14.76667] ( Ferreira 1906:160); “Quilombo,

Rio Luinha ” [-9.33333, 14.90000] ( Ferreira

1906:163; Ceríaco et al. 2014a:24). Kwanza

Sul: “Congulu” [-10.86667, 14.28333] ( Parker

1936:143). Malanje: “Bange N’gola”

[-8.43333, 16.56667] ( Boulenger 1905:110);

“ Duque de Bragança ” [-9.10000, 15.95000]

( Bocage 1873b:225, 1895a:165, 1897a:203;

Ferreira 1906:163; Ceríaco et al. 2014a:24).

MAP 36. Distribution of Hyperolius bocagei in Angola. Lunda Norte: “Dundo” [-7.36667, 20.83333] ( Laurent 1950a:16, 1954a:79, 1964a:150); “Muita (Luembe E)” [-7.80000, 21.45000] ( Laurent 1950a:16). Moxico: “Cazombo” [-11.88333, 22.91667] ( Laurent 1964a:150). Huambo: “Bimbi” [-11.81667, 15.83333] ( Monard 1937a:40 1938:93, 96). Bié: “Chitau” [-11.43333, 17.15000] ( Schmidt 1936:132); “Bihé” [-12.38333, 16.95000] ( Bocage 1895a:168). Benguela: “Ebanga” [-12.73333, 14.73333] ( Monard 1938:93); “Dombe” [-12.95000, 13.10000] ( Bocage 1895a:168). Huíla: “Caconda” [-13.73333, 15.06667] ( Bocage 1895a:165); “Kuvangu” [-14.46667, 16.30000] ( Monard 1937a:40, 1938:93, 95); “Osi” [-15.08333, 15.41667] ( Monard 1938:93). Cunene: “ruisseau Mbalé” [-15.16667, 16.75000] ( Monard 1938:93); “Kamba” [-16.28333, 15.23333] ( Monard 1938:93). Cuando Cubango: “Kuandu” [-16.74487, 19.10136] ( Monard 1937a:40, 1938:93, 95). Undetermined localities: without precise location ( Boulenger 1882:165); “Nana Meya” ( Boulenger 1905:110).

Taxonomic and distributional notes: Hyperolius bocagei Steindacher, 1867 was described based on a specimen from “ Angola ”. However, the type locality of H. bocagei should be considered “Duque de Bragança [= Calandula], Malanje,” as this is the locality from which the specimen collected by Pinheiro Bayão was sent by Bocage to Steidachner (Ceríaco et al. 2014a). Ferreira (1906) described Rappia seabrai based on a single individual deposited in Museu do Porto, collected by Francisco Newton. The holotype is believed to be lost (Ceríaco et al. 2014a) and several authors consider it to be a synonym of H. bocagei (Laurent, pers. comm. in Frost, 1985; Frétey et al., 2011; Ceríaco et al. 2014a; Frost 2016). In the original description, Ferreira (1906) also noted that one specimen referable to H. seabrai from “Duque de Bragança ” [= Calandula] collected by Bayão was found in Museu Bocage, Lisboa, Portugal, and this was considered a paratype by Ceríaco et al. (2014a). The type locality of “Quilombo,” currently Gonguembo in Kwanza Norte

Province, is approximately 140 km west of Caladula Falls in Malanje Province, formerly “Duque de Brangança,” the type locality of H. bocagei and the locality from which the paratype of R. seabrai was collected (Ceríaco et al. 2014a). Schiøtz (1999) and Schiøtz and Van Daele (2003) suggested that H. bocagei might be a junior synonym of a species in the Hyperolius viridiflavus complex. Poynton (1986) considered Hyperolius cinctiventris Cope, 1862 to be a synonym of Hyperolius argus Peters, 1854 , and as this species is restricted to East Africa it is unlikely to correspond to Angolan specimens. The specimens identified by Monard (1938) as cinctiventris deposited in the Musée d’Histoire Naturelle, La-Chaux-de Fond, Switzerland were recently studied and identificated as Hyperolius bocagei (Ceríaco et al. in prep.).

Hyperolius chelaensis Conradie, Branch, Measy and Tolley, 2012 CHELA MOUNTAIN REED FROG (Endemic)

Hyperolius chelaensis Conradie et al. 2012 a:5, figs 5-6. Holotype: PEM A9223 (collector W. Conradie). Type locality: “a small patch (<2.5ha.) of Afromontane forest in a small gorge draining from Serra da Chela above the Estacao Zootecnica, near Humpata, Lubango” (Conradie et al. 2012a:5), Angola.

Hyperolius chelaensis : Conradie et al. (2013:203), Frost (2016).

Global conservation status (IUCN): Data Deficient.

Global distribution: The species is known only from Angola.

Ocurrences in Angola (Map 37): The species is known only from the type locality “Serra de Chela,” Lubango . Huíla: “ Serra de Chela gorge, above the Estação Zootecnica near Humapata, Lubango” [-14.88944, 13.27417] (Conradie et al. 2012a:5; Conradie et al. 2013:203) .

Taxonomic and distributional notes: This species was recently described by Conradie et al. (2012a). According to the original publication, H. chelaensis is the sister species of H. cinereus Monard, 1937 . The type locality is along a small stream that drains from the Serra de Chela escarpment into the ephemeral water courses (e.g., Giraul River) draining west to the Atlantic Ocean ( Conradie et al. 2013). It is expected to be more widespread in the Serra de Chela and Leba mountain ranges. Further MAP 37. Distribution of Hyperolius chelaensis in surveys are needed to establish a full distribu- Angola. tion of the species and its conservation status.

Hyperolius cinereus Monard, 1937 ASHY REED FROG (Endemic)

Hyperolius cinereus Monard 1937a:32 . Syntypes: MHNC 90.0856–7 (collector A. Monard). Type locality: “Kalukembé” and “Bimbi” ( Monard 1937a:32), [= Caluquembe and Bimbe], Angola.

Hyperolius cinereus : Monard (1938:85), Cei (1977:17), Frost (1985:209, 2016), Schiøtz (1999:192), Channing (2001:152), Conradie et al. (2012a:17), Conradie et al. (2013:222) .

Hyperolius (Hyperolius) cinereus : Frétey et al. (2011:30).

Global conservation status (IUCN): Least Concern.

Global distribution: The species is known only from Angola.

Ocurrences in Angola (Map 38): The species is known from southwestern and central Angola. Bié: “western dambo of Cacuchi

River, Bie Province ” [-13.58333, 16.86667]

( Conradie et al. 2013:222); “Cubango basin

(11)” [-13.69413, 17.06177] ( Conradie et al.

2016:8-9, 16). Huambo: “Bimbi” [-11.81667,

15.83333] ( Monard 1937a:32, 1938:85, Channing 2001:153; Conradie et al. 2013:222);

“small stream at Huambo Agriculture Institute,

Huambo Province ” [-12.71667, 15.80000]

( Conradie et al. 2013:222); “1st stream crossing east of Caccuchi River, Bie Province ”

[-13.68333, 17.05000] ( Conradie et al.

2013:222). Benguela: “Zamba-Seidlung bei

Entre Rois, Benguella Province” [-13.01667, MAP 38. Distribution of Hyperolius cinereus in Angola. 14.63333] ( Conradie et al. 2013:222). Huíla: “Kalukembé” [-13.78333, 14.68333] ( Monard 1937a:32, 1938:85; Frost 1985:209, 2016; Channing 2001:153; Conradie et al. 2013:201, 222); “stream under road before Estacao Zootecnica near Humpata, Lubango” [-14.90400, 13.32556] (Conradie et al. 2012a:17; Conradie et al. 2013:222); “waterfall below dams, on plateau above Estacao Zootecnica near Humpata, Lubango” [-14.91425, 13.31386] (Conradie et al. 2012a:17, Conradie et al. 2013:222). Cuando Cubango: “ 5 km south of Soba Matios Military base, Cuando” [-14.25706, 17.77861] ( Conradie et al. 2013:222); “small stream after Muvange river, near Menongue” [-14.73333, 17.66667] ( Conradie et al. 2013:222); “Muvange river crossing, near Menongue” [-14.81667, 17.66667] ( Conradie et al. 2013:222); “Cubango basin (5)” [-14.74628, 17.66844] ( Conradie et al. 2016:8-9, 16); “Cubango basin (6b)” [-14.67458, 17.73544] ( Conradie et al. 2016:8-9, 16); “Cubango basin (7c)” [-14.43916, 17.81491] ( Conradie et al. 2016:8-9, 16); “Cubango basin (8)” [-14.25705, 17.77852] ( Conradie et al. 2016:8-9, 16); “Cubango basin (21)” [-14.66586, 17.07661] ( Conradie et al. 2016:8-9, 16); “Cubango basin (22b)” [-14.66278, 16.96081] ( Conradie et al. 2016:8-9, 16); “Cuito basin (27)” [-15.17127, 19.19433] ( Conradie et al. 2016:8-9, 16); “Cuito basin (30a)” [-17.50875, 20.06594] ( Conradie et al. 2016:9- 10, 16); “Cuito basin (30d)” [-17.51327, 20.06111] ( Conradie et al. 2016:9-10, 16); “Cuito basin (32)” [-17.04880, 19.53333] ( Conradie et al. 2016:9-10, 16); “Cuito basin (35)” [-16.62322, 19.05352] ( Conradie et al. 2016:9-10, 16); “Cuando basin (43)” [-17.53500, 23.18916] ( Conradie et al. 2016:9-10, 16); “Cubango basin (45)” [-16.88350, 18.01180] ( Conradie et al. 2016:9, 12, 16). Undetermined Locality: “areas of forest and savanna in the north and northeast of Angola ” ( Cei 1977:17); “plateaus regions” ( Cei 1977:17).

Taxonomic and distributional notes: Hyperolius cinereus Monard, 1937 was described based on two syntype specimens from “Kalukembé” in Huíla Province and “Bimbi” in Huambo Province ( Monard 1937a). Laurent (1964a) later assigned a small collection of specimens from “Dundo” in Lunda Norte Province to H. cinereus , althought his characterization of the coloration of this species differs from that of Monard. Both Monard (1937a) and Laurent (1964a) examined only preserved material and could only speculate on the coloration in life ( Conradie et al. 2013). Conradie et al. (2013) considered that populations from southern and central Angola, including the localities for Monard’s two syntypes, correspond to H. cinereus . However, the northern populations identified by Laurent (1964a) correspond to a closely related species that they described as Hyperolius raymondi Conradie, Branch and Tolley, 2013 (see H. raymondi ). Both species occur in flood- ed grasslands called “dambos” where water levels reach 30 (northern population) to 50 (southern population) cm deep ( Conradie et al. 2013). Monard (1937a) stated that the syntype from “Bimbi” was a juvenile and differed in several characters. Due to that, Conradie et al. (2013a) argued that this specimen should not be considered part of the type series, instead assigning it only provisionally to H. cinereus , which was followed by Frost (2016). We recently studied the type material in the Musée d’Histoire Naturelle, La-Chaux-de Fond, Switzerland and believe that the “Bimbi” specimen is identifiable as Hyperolius cinereus , and as it was used by Monard in his description it should be recognized as a part of the type series (Ceríaco et al. in prep.).

Hyperolius cinnamomeoventris Bocage, 1866 CINNAMON-BELLIED REED FROG

Hyperolius cinnamomeo-ventris Bocage 1866a:55 , 1866b:75. Holotype: MBL (collector F.A.P. Bayão), not localted by Perret (1976a:24), destroyed by fire 18 March 1978. Type locality: “Duque de Bragança ”

( Bocage 1866a:55), [= Calandula] Malanje Province, Angola. Rappia tristis Bocage 1866a:56 , 1866b:76. Holotype: MBL (collector F.A.P. Bayão), not localted by Perret

(1976a:25), destroyed by fire 18 March 1978. Type locality: “ Duque de Bragança ” ( Bocage 1866a:56) ,

[= Calandula] Malanje Province, Angola. Rappia cinnamomeiventris : Bocage (1895a:172, 1897a:204), Ferreira (1906:163). Rappia cinnamomei-ventris : Bocage (1897b:211), Noble (1924:252, 253), Mertens (1938:429). Hyperolius tristis : Bocage (1895a:171, 1897a:204), Boulenger (1882:121), Perret (1976a:25). Rappia bivittata : Ferreira (1906:161). Hyperolius cinnamomeoventris : Parker (1936:144), Laurent (1950a:16, 1954a:78, 1964a:149), Cei (1977:17),

Perret (1976a:25), Frost (1985:209, 2016), Schiøtz (1999:129), Channing (2001:153), Conradie et al.

(2013:206), Ceríaco et al. (2014a:19), Ceríaco et al. (2016b:37). Hyperolius cinnamomeoventris cinnamomeoventris : Laurent (1961:79), Schiøtz (1975:123). Hyperolius (Hyperolius) cinnamomeoventris : Frétey et al. (2011:30).

Global conservation status (IUCN): Least Concern.

Global distribution: The species is known from central and southern Cameroon south to Angola, extending across the Democratic

Republic of Congo to northwestern Zambia,

and east to Uganda and western Kenya.

Ocurrences in Angola (Map 39): The species is found in western areas along the coast as well as northeastern regions of the country. Lunda Norte: “Dundo” [-7.36667,

20.83333] ( Laurent 1954a:78); “environs de

Dundo, forêt des sources de la Dundundo”

[-7.36667, 20.83333] ( Laurent 1964a:149);

“Matala, rive gauche de la Tshihumbwe, 40 km

à l’est de Dundo” [-7.43333, 21.16667] ( Laurent 1950a:16); “Andrada (Luembe O)”

[-7.68729, 21.37003] ( Laurent 1954a:78);

“ Muita (Luembe E)” [-7.80000, 21.45000]

( Laurent 1950a:16); “riv. Kakuje, affl. gauche

Luembe près du village du “soba”, Santana”

[ -8.56667, 20.56667] ( Laurent 1954a:78). MAP 39. Distribution of Hyperolius cinnamomeoventris Malanje: “ Duque de Bragança ” [-9.10000, in Angola.

15.95000] ( Bocage 1866a:55, 56, 1866b:76, 1895a:171, 172, 1897a:204; Laurent 1961:79; Schiøtz 1975:123, Perret 1976a:25; Frost 1985:209, 2016); “Kalandula” [-9.10000, 15.95000] ( Conradie et al. 2013:206), “Cangandala National Park” [-9.84606, 16.72233] (Ceríaco et al. 2016b:37). Kwanza Norte: “N’golla Bumba” [-9.03333, 14.60000] ( Ferreira 1906:161; Ceríaco et al. 2014a:19); “Rio Luinha” [-9.26667, 14.53333] ( Ferreira 1906:163); “Quilombo” [-9.33333, 14.90000] ( Ferreira 1906:163). Kwanza Sul: “Novo Redondo” [-11.20000, 13.85000] ( Ferreira 1906:163); “Congulu” [-10.86667, 14.28333] ( Parker 1936:144). Benguela: “Hanha” [-13.30000, 14.20000] ( Bocage 1897a:204, 1897b:211; Perret 1976a:25); “Cubal” [-13.03333, 14.25000] (Mertens 1938:429). Undetermined Locality: “Mupèpe” ( Ferreira 1906:163); “areas of forest and savanna in the north and northeast of Angola ” ( Cei 1977:17); “plateaus regions” ( Cei 1977:17).

Taxonomic and distributional notes: Bocage (1866b) described two new species, Hyperolius cinnamomeo-ventris Bocage, 1866 and Rappia tristis (Bocage, 1866) from “Duque de Bragança [= Calandula].” The nomen tristis was synonymized with H. cinnamomeoventris by Laurent (1943b, 1947) and has subsequently been followed by other authors including Perret (1976a). Before the Lisbon fire Perret (1976a) visited the Museu Bocage but did not find the holotype of either of these species. One syntype of Rappia bivittata designated by Ferreira (1906) from “N’Golla Bumba,” Kwanza Sul province seems to represent Hyperolius cinnamomeoventris Bocage, 1866 (Ceríaco et al. 2014a). Currently this species is accepted and recognized as Hyperolius cinnamomeoventris ( Schiøtz 1999; Channing 2001; Frétey et al. 2011; Conradie et al. 2013; Frost 2016). This widespread species represents a complex of species found in Central Africa and several offshore islands and is awaiting taxonomic revision.

Hyperolius concolor (Hallowell, 1844) VARIABLE REED FROG

Ixalus concolor Hallowell 1844:60 . Holotype: ANSP 3216 ( Malnate 1971:350) (collector unknown). Type locality: “ Liberia, W. Africa ” (Hallowell 1844:60), Liberia.

Hyperolius modestus : Bocage (1866a:55, 1866b:74).

Rappia concolor : Bocage (1895a:173), (1897b:211).

Hyperolius concolor : Noble (1924:252), Schiøtz (1999:104), Amiet (2012:264), Frost (2016).

Global conservation status (IUCN): Least Concern.

Global distribution: The species is known from eastern Sierra Leone to western

Cameroon, and along the Atlantic coast south to Angola.

Ocurrences in Angola (Map 40): The species is recorded mostly from western Angola. Malanje: “Duque de Bragança” [-9.10000,

15.95000] ( Bocage 1866a:55, 1866b:74). Bié:

“Bihé” [-12.38333, 16.95000] ( Bocage

1895a:173). Benguela: “Hanha” [-13.30000,

14.20000] ( Bocage 1897b:211). Huíla :

“Caconda” [-13.73333, 15.06667] ( Bocage

1895a:173); “ Huíla ” [-15.05000, 13.55000]

( Bocage 1895a:173). Undetermined Locality:

“ Rio Quando” ( Bocage 1895a:173).

Taxonomic and distributional notes:

The Angolan records of Hyperolius concolor

(Hallowell, 1844) probably represent a MAP 40. Distribution of Hyperolius concolor in Angola.

misidentification, although Frétey et al. (2011) considered the species present in Angola. Unfortunately, the specimens used by Bocage (1866a, 1866 b, 1895a, 1897b) were destroyed in the 1978 fire at the Museu Bocage, Lisboa. Noble (1924) cited one specimen of H. concolor from Angola at the American Museum of Natural History, although he did not provide more precise locality information.

Hyperolius dartevellei Laurent, 1943 DARTEVELLE’S REED FROG

Hyperolius dartevellei Laurent 1943:71 , fig. 3. Holotype: MRAC 38385 (collector E. Dartevelle). Type locality: “Zambi, Bas-Congo”, Democratic Republic of Congo. Hyperolius nasutus adspersus : Laurent (1961:92). Hyperolius dartevellei : Channing et al. (2002:96), Amiet (2015:275), Channing et al. (2013:319), Frost

(2016).

Global conservation status (IUCN): Least Concern.

Global distribution: The species is known from southern Cameroon to the lower Congo Basin, northern Angola and the highlands of northwestern Zambia.

Ocurrences in Angola: The species is known from the most northern parts of Angola as plotted by Channing (2001). Those occurrences are not mapped here.

Taxonomic and distributional notes: Hyperolius dartevellei Laurent, 1943 was recently removed by Channing et al. (2013) from the synonymy of Hyperolius adspersus Peters, 1877 , in which it had been placed by Laurent (1961). Channing et al. (2002) and Amiet (2005) considered it a member of nasutus complex. The species is presently only confirmed from northern Angola, including the Cabinda enclave (Channing et al. 2013).

Hyperolius fuscigula Bocage, 1866 BROWN-THROATED REED FROG (Endemic)

Hyperolius fuscigula Bocage 1866a:56 , 1866b:76. Syntypes: MBL 19-189 [2 specimens] (collector F.A.P.

Bayão), destroyed by fire 18 March 1978. Type locality: “ Duque de Bragança ” ( Bocage 1866a:56) ,

[= Calandula], Angola. Rappia fuscigula : Bocage (1895a:170, 1897a:204). Hyperolius fuscigula : Noble (1924:253), Perret (1976a:26), Frost (2016). Hyperolius cinnamomeoventris : Laurent (1943:67). Hyperolius (Hyperolius) fuscigula : Frétey et al.

(2011:30).

Global conservation status (IUCN):

Data Deficient.

Global distribution: The species is known from Angola.

Ocurrences in Angola (Map 41): The species is known only from the type locality in

Malanje. Malanje: “Duque de Bragança ”

[-9.10000, 15.95000] ( Bocage 1866a:56,

1866b:76, 1895a:170, 1897a:204; Perret

1976a:26).

Taxonomic and distributional notes:

Laurent (1943b) tentatively considered Hyperolius fuscigula Bocage, 1866 to be a synonym of Hyperolius cinnamomeoventris Bocage.

1866. Perret (1976a) noted the poor condition MAP 41. Distribution of Hyperolius fuscigula in Angola.

of the type specimens that were later destroyed in the 1978 fire. Because of the lack of records and loss of the type material, the validity of this taxon is uncertain.

Hyperolius glandicolor Peters, 1878 PETERS’ REED FROG

Hyperolius glandicolor Peters 1878:208 , pl. 2, fig. 9. Syntypes: ZMB 9299 [2 specimens] (collector J.M. Hildebrandt). Type locality: “ Taita ”, Kenya.

Hyperolius glandicolor : Schiøtz (1999:212), Wieczorek et al. (2000:1333), Channing and Howell (2006:162), Conradie et al. (2012a:6).

Global conservation status (IUCN): Least Concern.

Global distribution: The species is known from Kenya, southern Somalia, and Tanzania, Angola and adjacent Democratic Republic of Congo and northern Mozambique.

Ocurrences in Angola (Map 42): The only Angolan record is from the northwest of the country. Malanje: “ Duque de Bragança ” [-9.10000, 15.95000] ( Bocage 1866a:56) .

Taxonomic and distributional notes: This species was removed from the synonymy of Hyperolius marmoratus Rapp, 1842 by Wieczorek et al. (2000), and Schiøtz (1999) has argued it is a member of the viridiflavus subgroup in the viridiflavus complex. The recognized distribution of this species is limited to the eastern coast of equatorial Africa and the associated interior, where it is often associated with higher elevations (Wiezczorek et al. 2000). The Angolan record from “Duque de Bragança ” [= Calandula] likely should be referred to another species of Hyperolius .

MAP 42. Distribution of Hyperolius glandicolor in Angola.

Hyperolius gularis Ahl, 1931 LUANDA REED FROG (Endemic)

Hyperolius gularis Ahl 1931b:408 , fig. 281. Holotype: ZMB uncatalogued (collector K. May), probably lost. Type locality: “Loanda” [= Luanda] Luanda Province, Angola.

Hyperolius gularis : Frost (1985:211, 2014).

Global conservation status (IUCN): Data Deficient.

Global distribution: The species is known only from Angola.

Ocurrences in Angola ( Map 43 ): The species is known only from the type locality “Loanda”. Luanda: “Loanda” [-8.83333, 13.26667] ( Ahl 1931a:125; Frost 1985:211, 2016).

Taxonomic and distributional notes: The taxonomic validity of this species is doubtful and it has not been discussed in recent synopses of Hyperolius by Schiøtz (1999), Channing (2001), or Frétey et al. (2011). However, Laurent (in Frost 1985; Frost 2016) suggested that this species is probably a synonym of Hyperolius marmoratus Rapp, 1842 , which we recognize above as Hyperolius angolensis Steindachner, 1867 (see H. angolensis account).

Hyperolius kivuensis Ahl, 1931 KIVU REED FROG

Hyperolius kivuensis Ahl 1931b:280 ., fig. 151. Holotype: ZMB 36098 (collector Kandt). Type locality: “Kivu- See” ( Ahl 1931a:280), [= Lake Kivu], Democratic Republic of Congo.

Hyperolius multifasciatus : Monard (1937a:33, 1938:87).

Hyperolius kivuensis kivuensis : Laurent (1950:16, 1954a:78, 1964a:149), Poynton and Broadley (1976:196).

Hyperolius quinquevittatus : Laurent (1950a:16, 1954a:79).

Hyperolius kivuensis : Cei (1977:17), Schiøtz (1999:163), Channing (2001:157), Frost (2016).

Hyperolius kivuensis multifasciatus : Pickersgill (2007a:325).

Hyperolius (Hyperolius) kivuensis : Frétey et al. (2011:31).

Global conservation status (IUCN): Least Concern.

Global distribution: The species is widespread in savannas of southwestern Ethiopia, southeastern Sudan, western Kenya, Tanzania,

Uganda, Rwanda, and Burundi, eastern and southern Democratic Rebublic of Congo,

Angola, Zambia, Malawi, and northwestern

Mozambique.

Ocurrences in Angola (Map 44): The species is well documented from northeastern

Angola, though may also occur in southern regions. Lunda Norte: “Dundo” [-7.36667,

20.83333] ( Laurent 1950a:16, 1954a:78-79);

“Matala, rive gauche de la Tshihumbwe, 40 km

à l’est de Dundo” [-7.43333, 21.16667] ( Laurent 1950a:16); “Andrada (Luembe O)”

[-7.70000, 21.38333] ( Laurent 1954a:78);

“ Muita (Luembe E)” [-7.80000, 21.45000]

( Laurent 1950a:16, 1954a:78); “Kossa”

[-7.90000, 21.36667] (Laurent 1950:16).

Lunda Sul: “Alto Chicapa” [-10.88333, MAP 44. Distribution of Hyperolius kivuensis in Angola. 19.23333] ( Laurent 1964a:149). Huambo: “Bimbi” [-11.81667, 15.83333] ( Monard 1938:87). Undetermined Locality: “with no precise identification” ( Laurent 1954a:78), “areas of forest and savanna in the north and northeast of Angola ” ( Cei 1977:17).

Taxonomic and distributional notes: Poynton and Broadley (1987) recognized that Hyperolius kivuensis Ahl, 1931 is easily confused with Hyperolius quinquevittatus Bocage, 1866 , though they stated that the latter is generally smaller and slimmer. Whereas Laurent (1954a) treated H. multifasciatus (Ahl, 1931) as a junior synonym of H. quinquevittatus, Pickersgill (2007a) considered it to be a subspecies of Hyperolius kivuensis . Schiøtz (1999) and Channing (2001) provid- ed some additional records, though without information about the source.

Hyperolius langi Noble, 1924 LANG’S REED FROG

Hyperolius langi Noble 1924:266 , pl. 39, fig. 1. Holotype: AMNH 9983 (collector Mr. Lang). Type locality: “Niapu” Democratic Republic of Congo.

Hyperolius langi : Monard (1937a:40, 1938:95).

Hyperolius platycephalus langi : Cei (1977:17).

Hyperolius langi : Schiøtz (1999:149), Frost (2016).

Global conservation status (IUCN): Least Concern.

Global distribution: The species is known from Angola, the northeastern Democratic Republic of Congo and Uganda.

Ocurrences in Angola (Map 45): The species has been recorded from Benguela

Province. Benguela: “Ebanga” [-12.73333,

14.73333] ( Monard 1937a:40, 1938:95).

Undetermined Locality: “plateaus regions”

( Cei 1977:17).

Taxonomic and distributional notes:

Monard (1937 a, 1938) reported H. langi from

“Ebanga” in southwestern Angola. There are also two other species of Hyperolius that occur at this location, H. benguellensis ( Bocage,

1893) and H. bocagei Steindachner, 1867 .

The species that Monard (1937 a, 1938) identified as H. langi probably corresponds to

H. bocagei , especially as it does not agree with the original description for H. benguellensis .

However, examination of the type material of

H. langi is required to confirm this. MAP 45. Distribution of Hyperolius langi in Angola.

Hyperolius maestus Rochebrune, 1885 CABINDA REED FROG (Endemic)

Hyperolius Maestus Rochebrune 1885:91 . Type: Presumably in MNHN (collector unknown), unrecognized as a type, if still extant, formerly in Museo Bouvieri. Type locality: “Locum Landana dictum, in foliis Musarum” [= Landana, Cabinda enclave], Angola.

Hyperolius maestus : Frost (1985:212, 2014).

Hyperolius (Hyperolius) marmoratus : Frétey et al. (2011:32).

Global conservation status (IUCN): Data Deficient.

Global distribution: The species is known from Angola.

Ocurrences in Angola (Map 46): The species is only known from the type locality “Landana” in the Cabinda enclave. Cabinda: “Landana” [-5.21667, 12.15000] ( Rochebrune 1885:91; Frost 1985:212, 2016).

Taxonomic and distributional notes: The species was described by Rochenbrune (1885) from “Locum Landana dictum, in foliis Musarum” in the Cabinda enclave. Laurent (in Frost 1985), and latter Frétey et al. (2011) considered Hyperolius maestus Rochebrune, 1885 to be a possible synonym of Hyperolius marmoratus Rapp, 1842 . The type specimen was deposited in Museo Bouvier and is now presumably in Muséum national d’Histoire naturelle, Paris and unrecognized as a type if it still exists ( Frost 2016). The species is not mentioned in Schiøtz (1999) or Channing MAP 46. Distribution of Hyperolius maestus in Angola. (2001).

Hyperolius nasutus Günther, 1865 LARGE-NOSED LONG REED FROG

Hyperolius nasutus Günther 1865a:482 , Pl XXXIII, fig. 3. Holotype: BMNH 1947.2 .9.68 (collector J.F. Bayão). Type locality: “ Duque de Bragança ” [= Calandula], Malanje Province, Angola.

Rappia punctulata Bocage 1895a:168 . Holotype: MBL T.23-227 (collector Banyures) destroyed by fire 18 March 1978. Type locality: “sur les bords du Quanza” [= Quanza river ], Angola.

Hyperolius nasutus : Bocage (1866a:55), Noble (1924:253), Schmidt (1936:132), Loveridge (1936a:109, 1953a:362, 1953:362, 1957:333), Monard (1937a:39, 1938:94), Mertens (1938:429), Frade (1963:254), Haacke (1970:279), Schiøtz (1975:97), Poynton and Broadley (1976:206), Cei (1977:17, 18), Frost (1985:214, 2016), Poynton and Haacke (1993:14), Schiøtz (1999:97, 2006:65), Channing (2001:166), Largen (2001:359), Channing et al. (2002:96), Schiøtz and Van Daele (2003:138), Amiet (2005:275), Conradie et al. (2012a:3), Ceríaco et al. (2016b:35), Conradie et al. (2016:11).

Rappia nasuta : Boulenger (1882:127), Bocage (1895a:169, 1897a:204).

Rappia punctulata : Bocage (1897a:204), Ferreira (1904:112), Amiet (2005:275).

Hyperolius punctulatus : Noble (1924:253), Loveridge (1936a:107, 1936b:405), Mertens (1937:20), Monard (1938:86), Frade (1963:254), Perret (1976a:27), Frost (1985:216).

Hyperolius nasutus nasutus : Laurent (1950a:17, 1954a:84, 1964a:154), Hellmich (1957a:29).

Hyperolius (Hyperolius) nasutus : Frétey et al. (2011:32).

Global conservation status (IUCN): Least Concern.

Global distribution: The species is known from Angola south to the Okavango Delta of Botswana and adjacent northern Namibia.

Ocurrences in Angola (Map 47): The species is known from its type locality “Duque de Bragança, Malanje Province ” and is very widespread across almost the whole of the country. Bengo: “Cabiri” [-8.91667, 13.66667] ( Ferreira 1904:112). Kwanza Norte: “ 3 km W of Salazar” [-9.30000, 14.91667] (Poynton and Haacke 1993:14); “Dondo” [-9.68333, 14.43333] ( Loveridge 1936a:107). Kwanza Sul: “Namba” [-11.91667, 14.85000] (Poynton and Haacke 1993:14). Malanje: “Duque de Bragança” [-9.10000, 15.95000] ( Günther 1865a:482; Bocage 1866a:55, 1895a:169, 1897a:204; Boulenger 1882:127; Loveridge 1936a:109, 1936b:405, 1953a:362, 1957:333; Schiøtz 1975:97; Frost 1985:214, 2016; Amiet 2005:275); “Calandula” [-9.07917,

15.79583] (Channing et al. 2013:334); “Kan-

gandala” [-9.82511, 16.91225] (Channing et al.

2013:334); “Cangandala National Park”

[-9.84606, 16.72233] (Ceríaco et al. 2016b:35);

Lunda Norte: “Dundo” [-7.36667, 20.83333]

( Laurent 1954a:84); “environs de Dundo, rives de la Luachimo” [-7.38333, 20.85000] ( Laurent 1950a:17); “Carumbo lagoon” [-7.74422,

19.95467] (Conradie et al. 2012a:3); “Muita

(Luembe E)” [-7.80000, 21.45000] ( Laurent

1950a:17); “Kossa à 100 km au sud-est de

Dundo” [-7.90000, 21.36667] ( Laurent

1950a:17); “rive de la Tchihumbwe 40 km à l’est de Dundo” [-8.01667, 19.31667] ( Laurent

1950a:17); “Village Capaia” [-8.33847,

20.24250] (Conradie et al. 2012a:3). Lunda

Sul: “Dala” [-11.03333, 20.20000] ( Monard

1937a:39, 1938:94); “Alto Chicapa,sources du

MAP 47. Distribution of Hyperolius nasutus in Angola. Cuílo” [-10.88333, 19.23333] ( Laurent 1964a:154); “Alto Cuílo, rives du Cuílo “ [-10.01667, 19.55000] ( Laurent 1964a:154). Moxico: “région du lac Calundo” [-11.80000, 20.86667] ( Laurent 1964a:154); “réserve de chasse de Cameia” [-11.83333, 21.00000] ( Laurent 1964a:154). Bié: “Chitau” [-11.43333, 17.15000] ( Schmidt 1936:132); “General Machado” [-12.01667, 17.06667] Mertens (1937:20). Huambo: “Bimbi” [-11.81667, 15.83333] ( Monard 1937a:39, 1938:94); “Tongrube am Jamba Flüβchen” [-13.60000, 16.60000] ( Hellmich 1957a:29). Benguela: “Cubal” [-13.03333, 14.25000] ( Mertens 1938a:429); “Marco de Canavezes (Cubal da Ganda)” [-13.08333, 14.33333] ( Laurent 1964a:154). Huíla: “Huilla” [-15.05000, 13.55000] ( Bocage 1895a:169, 1897a:204); “Caconda” [-13.73333, 15.06667] ( Bocage 1895a:169, 1897a:204); “Kalukembé” [-15.05000, 13.55000] ( Monard 1938:86); “Nuntechite lagoon” [-15.13333, 13.41667] (Poynton and Haacke 1993:14). Cunene: “Chimporo” [-16.03333, 17.15000] ( Monard 1937a:39, 1938:94).

Cuando Cubango: “Kuandu” [-16.74487, 19.10136] ( Monard 1937a:39, 1938:94). Undetermined Locality: “Margens do Cuanza” ( Bocage 1895a:168, 1897a:204; Loveridge 1936a:107, 1957:333; Perret 1976a:27; Frost 1985:216; Amiet 2005:275); “without precise location” ( Monard 1937a:39, 1938:94); “Western subregion and Angolan highlands (Angolan coastal or watersheds)” ( Frade 1963:254); “Zambezian highlands” ( Frade 1963:254); “areas of forest and savanna in the north and northeast of Angola ” ( Cei 1977:17); “plateaus regions” ( Cei 1977:17); “arid territories along the coast” ( Cei 1977:18).

Taxonomic and distributional notes: While the type specimen of R. punctulata is now lost, Channing et al. (2013) included it as a synonym of H. nasutus , for which there are many junior synonyms. To date, fifteen names have been used for members of the nasutus group, most of which have been synonymized with the nominotypical form ( Schiøtz 1999); a relatively recent list of these synonyms is provided by Amiet (2005). Poynton and Broadley (1987) recognized three species in the southern African savanna, Hyperolius viridis Schiøtz, 1975 , Hyperolius nasutus Günther, 1865 and Hyperolius benguellensis Bocage, 1893 . Schiøtz (1999) including H. benguellensis as a synonym of H. nasutus after arguing that differences in morphology and color pattern are not sufficient for distinguishing these two species. Later, Channing et al. (2002) retained H. benguellensis as a synonym of nasutus , but also divided the group into three cryptic species differing in advertisement call: Hyperolius acuticeps Ahl, 1931 , Hyperolius viridis Schiøtz, 1975 , and H. nasutus (Schiøtz and Van Daele 2003; Schiøtz 2006). Schiøtz (2006) distinguished two species, H. benguellensis and H. nasutus , in the nasutus group for Angola based on the structure of the advertisement call and color pattern. This widespread species is common in savannas and exhibits a preference for humid areas such as reeds growing in deep water, pools, streams, and large rivers ( Schiøtz 1999; Channing 2001).

Hyperolius nitidulus Peters, 1875 PLAIN REED FROG

Hyperolius nitidulus Peters 1875:209 , pl. 3, figs. 4, 4a. Holotype: ZMB 7729 fide Bauer et al. (1995:44) (collector R. Buchholz). Type locality: “Yoruba (Lagos)” ( Peters 1875:209), Nigeria.

Hyperolius nitidulus : Peters (1877a:619), Schiøtz (1999:206), Wieczorek et al. (2000:1238-1241), Rödel et al. (2010:178), Amiet (2012:194), Frost (2016).

Global conservation status (IUCN):

Least Concern.

Global distribution: The species is known from West African extending from

Guinea east into Nigeria, Angola and

Cameroon.

Ocurrences in Angola (Map 48): The species was cited from “Chinchoxo” in the

Cabinda enclave. Cabinda: “Chinchoxo”

[-5.10000, 12.10000] ( Peters 1877a:619).

Taxonomic and distributional notes:

According to Wieczorek et al. (2000), the species Hyperolius nitidulus Peters, 1875 is a

West African species associated with savanna and woodlands ( Schiøtz 1999; Wieczorek et al.

2000; Rödel et al. 2010; Amiet 2012). Based on the distribution of the species delimited by

Wieczorek et al. (2000), the record from north- MAP 48. Distribution of Hyperolius nitidulus in Angola. western Angola is likely a misidentification.

Hyperolius ocellatus (Günther, 1858) GOLDEN-EYED REED FROG

Hyperolius ocellatus Günther 1858a:326 . Lectotype: BMNH 1947.2.9.22, (collector L. Fraser), formerly BMNH 51.11.13.48, by implication of being referred to as “ holotype ” by Perret (1975a:27), Syntypes: BMNH [2 specimens] (collector Mr. Fraser, for the “ Fernando Po ” specimen) ( Günther 1858a:326). Type locality: “ Fernando Po ” [= Bioko, Guinea] fide Perret (1975a:24), who considered “ Angola ” an error. Originally given as “ Fernando Po ” and “ Angola ” by Günther (1858a:326).

Hyperolius Lucani Rochebrune 1885:91 . Type: Presumably in MNHP (collector unknown), uncatalogued as a type, if still extant, formerly in Museo Bouvieri. Type locality: “Locam Landana dictum, in frondibus Borassorum” [= Landana , Cabinda enclave] Cabinda enclave, Angola.

Hyperolius ocellatus : Günther (“1858b” 1859:88), Perret (1975a:24), Cei (1977:17), Frost (1985:214, 2016), Schiøtz (1999:141), Channing (2001:168), Channing et al. (2012:217).

Rappia ocellata : Boulenger (1882:123); Bocage (1895a:165).

Hyperolius lucani : Frost (1985:212, 2016).

Hyperolius (Hyperolius) ocellatus : Frétey et al. (2011:32).

Global conservation status (IUCN): Least Concern.

Global distribution: The species is from tropical Central Africa. It occurs in the Cabinda enclave in Angola, extending north into southeastern Nigeria and Bioko Island, east across Democratic Republic of Congo, with possibly occurences in western Uganda and Rwanda.

Ocurrences in Angola (Map 49): The species is recorded from northwestern Angola in the Cabinda encalve. Cabinda: “Landana” [-5.21667, 12.15000] ( Rochebrune 1885:91; Frost 1985:212, 2016); “ Cabinda ” ( Schiøtz 1999:141; Channing 2001:169).

Taxonomic and distributional notes: Günther (1858a) described Hyperolius ocellatus from “ Fernando Pó ” and “ Angola,” the latter without clear indication of the locality based on two specimens deposited in British Museum. Perret (1975a) considered the type locality “ Angola ” to be in error and designated the lectotype as being from “ Fernando Pó.” MAP 49. Distribution of Hyperolius ocellatus in Angola. Following Günther’s (1858a) description, Rochebrune (1885) described a new species from “Locam Landana dictum, in frondibus Borassorum” in the Cabinda enclave as Hyperolius lucani . This may be a synonym of Hyperolius ocellatus (Laurent in Frost 1985, 2016; Frétey et al. 2011), although it is not mentioned by Schiøtz (1999) or Channing (2001).

Hyperolius platyceps ( Boulenger, 1900) BENITO RIVER REED FROG

Rappia platyceps Boulenger 1900:444 , pl. 27, fig. 4. Syntypes: BMNH 1947.2.9.57–58 and BMNH 1900.2.17.89-90 [2 specimens] (collector G.L. Bates, formerly BMNH 1900.2.17.89–90). Type locality: “Benito River, north of the Gaboon River between 20 and 30 miles inland from the coast, Gaboon” [= Benito River ], Equatorial Guinea.

Rappia bivittata Ferreira 1906:161 , pl. 1. Syntypes: MHNFCP 229 017291, 017296 and 017302, [3 specimens] (collector F. Newton). Type locality: “N’Golla Bumba”, “Quilombo” [= Gonguembo] and “ Rio Luinha ,” Kwanza Norte Province, Angola.

Rappia platyceps var. angolensis Ferreira 1906:161 , pl. 1. Syntype: MHNFCP 017303 (Quilombo) (collector F. Newton), the other syntype from “N’golla Bumba” was not located. Type locality: “N’Golla Bumba” and “Quilombo” [= Gonguembo], Kwanza Norte Province, Angola.

Rappia fasciata Ferreira 1906:164 , pl. 1. Holotype: MHNFCP 017294 (collector F. Newton). Type locality: “Quilombo” [= Gonguembo] Kwanza Norte Province, Angola.

Hyperolius angolanus Ahl 1931b:271 . Replacement name for Rappia platyceps var. angolensis .

Rappia platyceps : Ferreira (1906:161).

Hyperolius platyceps : Noble (1924:253), Ceríaco et al. (2014a:24), Frost (2016).

Hyperolius angolanus Laurent (1950a:15) , Frost (1985:207).

Hyperolius platyceps angolanus : Laurent (1954a:77), Cei (1977:17).

Hyperolius fasciatus : Noble (1924:252), Cei (1977:17), Frost (1985:210).

Hyperolius ferreirai : Frost (1985:210).

Hyperolius (Hyperolius) platyceps : Frétey et al. (2011:33).

Global conservation status (IUCN): Least Concern.

Global distribution: The species is known from southern Cameroon to the southwestern Central African Republic, south to the Democratic Republic of Congo and northwestern Angola.

Ocurrences in Angola (Map 50): The species occurs in the north of the country. Lunda

Norte: “Dundo” [-7.53333, 21.08333] ( Laurent 1950a:15); “Luachimo” [-7.36667,

20.83333] ( Laurent 1954a:77). Kwanza

Norte: “N’golla Bumba” [-9.03333, 14.60000]

( Ferreira 1906:161; Frost 1985:207, 210;

Ceríaco et al. 2014a:23); “ Rio Luinha”

[-9.26667, 14.53333] ( Ferreira 1906:161; Frost

1985:210); “Quilombo” [-9.33333, 14.90000]

( Ferreira 1906:161, 164; Frost 1985:210, 2016;

Ceríaco et al. 2014a:22).

Taxonomic and distributional notes: Ahl

(1931a) elevated Rappia platyceps var.

angolensis Ferreira, 1906 . to full species and proposed the replacement name Hyperolius angolanus because of the the name was preoccupied by Hyperolius marmoratus var.

angolensis Steindachner, 1862 (Ceríaco et al.

MAP 50. Distribution of Hyperolius platyceps in Angola. 2014a). Ferreira (1906) also described as new

Rappia fasciata from “Quilombo” and Rappia bivitatta from “N’golla Bumba”, “Quilombo,” and “ Rio Luinha.” Recently Frétey et al. (2011) and Ceríaco et al. (2014a) recognized Hyperolius platyceps angolensis , Rappia fasciata , and Rappia bivittata as synonyms of H. platyceps .

Hyperolius polli Laurent, 1943 TSHIMBULU REED FROG

Hyperolius polli Laurent 1943b:96 , fig. 20. Holotype: MRAC 656 (collector Achten). Type locality: “Tshimbulu s/Luebi (Kasai)” Democratic Republic of Congo.

Hyperolius polli : Laurent (1954a:78).

Hyperolius polli : Schiøtz (1999:153), Frost (2016)

Hyperolius (Hyperolius) polli : Frétey et al. (2011:33).

Global conservation status (IUCN):

Data Deficient.

Global distribution: The species is known from the Democratic Republic of

Congo and adjacent Angola.

Ocurrences in Angola (Map 51): The single species record is in Lunda Norte

Province near the border with Congo. Lunda

Norte: “ Tshinguvu, Riv. Tshikapa” [-7.61667,

20.50000] ( Laurent 1954a:78).

Taxonomic and distributional notes:

According to Laurent (1954a), the specimen from “Tshinguvu (Tshikapa)” is similar to

Hyperolius cinnamomeoventris Bocage, 1866 ,

but differs in some morphological characters.

Currently it remains poorly known and its validity is uncertain.

MAP 51. Distribution of Hyperolius polli in Angola.

Hyperolius protchei Rochebrune, 1885 ROCHEBRUNE’S REED FROG (Endemic)

Hyperolius Protchei Rochebrune 1885:92 . Type: Presumably in MNHP (collector unknown), uncatalogued as a type, if still extant, formerly in Museo Bouvieri. Not cited by Guibé (1950). Type locality: “Locum Landana dictum, in foliis Musarum” [= Landana, Cabinda enclave], Angola .

Hyperolius protchei : Frost (1985:215, 2016).

Hyperolius (Hyperolius) marmoratus : Frétey et al. (2011:32).

Global conservation status (IUCN):

Data Deficient.

Global distribution: The species is known only from Angola.

Ocurrences in Angola (Map 52): The species is only known from the type locality

“Landana” in the Cabinda enclave. Cabinda:

“Landana” [-5.21667, 12.15000] ( Rochebrune

1885:92; Frost 1985:215, 2016).

Taxonomic and distributional notes:

Laurent (in Frost 1985) considered Hyperolius protchei as a possible synonym of Hyperolius marmoratus Rapp, 1842 , which in Angola should be considered Hyperolius angolensis

Steindachner, 1867 (see Hyperolius angolensis account). The species is not mentioned by

Schiøtz (1999) or Channing (2001).

MAP 52. Distribution of Hyperolius protchei in Angola.

Hyperolius pusillus ( Cope, 1862) WATER LILLY REED FROG

Crumenifera pusilla Cope 1862:343 . Holotype: ANSP 11323 (collector unknown) fide Malnate (1971:352).

Type locality: “Umvoti,” KwaZulu-Natal, South Africa. Hyperolius microps : Bocage (1866a:55, 1866b:75), Frade (1963:254), Noble (1924:253). Rappia microps : Bocage (1895a:173, 1897a:204), Boulenger (1882:127; 1905:110). Hyperolius pusillus : Cei (1977:17), Schiøtz

(1999:185), Channing (2001:176).

Global conservation status ( IUCN):

Least Concern.

Global distribution: The species is rectricted to the eastern Africa coastal lowlands,

from Somalia to KwaZulu-Natal.

Ocurrences in Angola (Map 53): Specimen records are from Malanje and Benguela

Province. Malanje: “ Fort Don Carlos ”

[-7.65000, 17.30000]; ( Boulenger 1905:110);

“Marimba” [-8.36667, 17.03333] ( Boulenger

1905:110); “Bange N’gola” [-8.43333,

16.56667] ( Boulenger 1905:110); “Duque de

Bragança ” [-9.10000, 15.95000] ( Bocage

1866a:55, 1866b:75, 1895a:173, 1897a:204).

Benguela: “Cahata” [-12.35000, 14.81667]

( Bocage 1895a:173, 1897a:204). Undetermi- MAP 53. Distribution of Hyperolius pusillus in Angola.

ned Locality: “bords du Quanza” ( Bocage 1895a:173, 1897:204); “ Rio Quando” ( Bocage 1895a:173, 1897a:204); “Zambezian highlands” ( Frade 1963:254); “plateaus regions” ( Cei 1977:17).

Taxonomic and distributional notes: Boulenger (1882) noted that this is an East African species, although he recognized ( Bocage 1866b) Hyperolius microps as a valid name. Based on this information, the Angolan records possibly belong to another Hyperolius species and it would be important to review this case. Unfortunately, the specimens from Museu Bocage ( Bocage 1866 b, 1895a, 1897a) were destroyed in the 1978 fire, although the specimens studied by Boulenger (1905) still exist in the British Museum. Noble (1924) also cited specimens of H. pusillus and H. microps in the collections of the American Museum of Natural History from Angola, though without precise locality information.

Hyperolius quinquevittatus Bocage, 1866 FIVE-STRIPED REED FROG

Hyperolius quinquevittatus Bocage 1866a:56 . Syntypes: MBL 24-214 , 24-215 [2 specimens] (collector F.A. P.

Bayão). Type locality: “Duque de Bragança” [= Calandula], Malanje Province, Angola. Hyperolius quinquevittatus : Bocage (1866b:77), Frade (1963:254), Noble (1924:253), Laurent (1950A:16,

1954a:79), Perret (1976a:25), Cei (1977:17), Frost (1985:216, 2016), Schiøtz (1999:101), Channing

(2001:178), Channing and Howell (2006:180). Rappia quinquevittata : Bocage (1895a:174). Hyperolius quinquevittatus quinquevittatus : Schiøtz (1975:102), Poynton and Broadley (1987:197), Pickersgill (2007a:360). Hyperolius (Hyperolius) quinquevittatus : Frétey et al. (2011:33).

Global conservation status (IUCN): Least Concern.

Global distribution: The species is known from open savannas at higher elevations in northern Angola, the southern Democratic Republic of Congo, southern Tanzania, and northern Zambia and Malawi, and Mt. Namuli in northern Mozambique.

Ocurrences in Angola (Map 54): The species occurs especially in northern Angola. Malanje: “Duque de Bragança” [-9.10000, 15.95000]

( Bocage 1866a:56, 1866b:77, 1895a:174;

Schiøtz 1975:102; Perret 1976a:25; Frost

1985:216, 2016; Poynton and Broadley

1987:197; Pickersgill 2007a:360). Undetermined Locality: “areas of forest and savanna in the north and northeast of Angola ” ( Cei

1977:17); “plateaus regions” ( Cei 1977:17).

Taxonomic and distributional notes:

According to Perret (1976a), the syntypes described by Bocage (1866a), which were destroyed in the 1978 fire in Lisbon, were studied by Schiøtz (1975) when he concluded that

H. quinquevittatus is similar in morphology to

H. nasutus Günther, 1865 . Poynton and

Broadley (1987) noted, however, that Hyperolius quinquevittatus is more similar to Hyperolius kivuensis even if H. quinquevittatus is in MAP 54. Distribution of Hyperolius quinquevittatus in general much smaller and slimmer. Pickersgill Angola.

(2007a) suspected that the species belongs to the multifasciatus group, while Frétey et al. (2011) included the H. multifasciatus in the synonym of quinquevittatus .

Hyperolius raymondi Conradie, Branch and Tolley, 2013 RAYMOND’S REED FROG (Endemic)

Hyperolius raymondi Conradie et al. 2013:203 , fig. 7. Holotype: PEM A10049 (collectors W. Conradie,

W. Branch, P. Vaz Pinto, S. Batista and N. Batista). Type locality: “Lagoa Carumbo from a dambo near the expedition base camp,” Lunda Norte Province, Angola. Hyperolius cinereus : Laurent (1964a:149), Schiøtz (1999:192), Channing (2001:152). Hyperolius (Hyperolius) cinereus : Frétey et al. (2011:30). Hyperolius raymondi : Frost (2016).

Global conservation status (IUCN): Data Deficient.

Global distribution: The species is known from Angola.

Ocurrences in Angola (Map 55): The species is known from the type locality “ Lagoa

Carumbo” and from surrounding localities in

Lunda Norte Province. Lunda Norte:

“Dundo” [-7.36667, 20.83333] ( Laurent

1964a:149; Channing 2001:153; Conradie et al. 2013:203); “margins Lagoa Samokaza east of Lagoa Carumbo” [-7.73833, 19.98694]

( Conradie et al. 2013:203); “Lagoa Carumbo from a dambo near the expedition base camp”

[-7.74422, 19.95522] ( Conradie et al.

2013:203; Frost 2016); “headwaters of Lovua,

north of village Capaia” [-8.33847, 20.24250]

( Conradie et al. 2013:203).

Taxonomic and distributional notes:

Populations that Laurent (1964a) referred to

H. cinereus were recognized by Conradie et al.

(2013a) as a distinct species and described as MAP 55. Distribution of Hyperolius raymondi in Angola. H. raymondi (see further discussion above in account for H. cinereus ). This newly described species is so far endemic to Angola, but may also occur in southernmost Democratic Republic of Congo and northwest Zambia. It is found in grass-covered “dambo” (water levels less than 30 cm) in the flood plains of the Lulele River, a southern tributary draining north to the larger Congo River ( Conradie et al. 2013).

Hyperolius rhizophilus Rochebrune, 1885 AFRICAN REED FROG (Endemic)

Hyperolius Rhizophilus Rochebrune 1885:92 . Type: Presumably in MNHN (collector unknown), uncatalogued as a type, if still extant, formerly in Museo Bouvieri. Not cited by Guibé (1950). Type locality: “Locum Landana dictum, in foliis rhizophorarum” [= Landana, Cabinda enclave], Cabinda Province, Angola .

Hyperolius rhizophilus Rochebrune, 1885 : Frost (1985:215, 2016).

Hyperolius (Hyperolius) rhizophilus Rochebrune, 1885 : Frétey et al. (2011:33).

Global conservation status (IUCN): Data Deficient.

Global distribution: The species is known only from Angola.

Ocurrences in Angola ( Map 56 ): The species is only known from the type locality “Landana”

in the Cabinda enclave. Cabinda: “Landana”

[-5.21667, 12.15000] ( Rochebrune 1885:92;

Frost 1985:215, 2016).

Taxonomic and distributional notes:

The species was described by Rochebrune

(1885) from “Locum Landana dictum, in foliis rhizophorarum” in the Cabinda enclave. The type specimen was deposited in the Museo

Bouvier, now presumably in the Muséum national d’Histoire naturelle, Paris ( Frost

2016). The species is not mentioned in Schiøtz

(1999) or Channing (2001).

Hyperolius steindachneri Bocage, 1866 STEINDACHNER’S REED FROG Hyperolius steindachneri Bocage 1866a:55 . Holotype: MBL T.25-226 (collector F.A.P. Bayão). Type locality: “Duque de Bragança” [= Calandula], Malanje Province, Angola. Hyperolius machadoi Laurent 1954a:80 , figs. 22, 23. Holotype: MD 495 (collector unknown, presumably

A.B. Machado). Type locality: “Camissombo” Lunda Norte, Angola. Hyperolius steindachneri : Bocage (1866b:75), Noble (1924:253), Loveridge (1936a:103), Perret (1976a:25),

Cei (1977:17), Frost (1985:218, 2016), Poynton and Haacke (1993:15), Schiøtz (1999:144), Channing

(2001:182), Conradie et al. (2012a:3). Rappia steindachneri : Bocage (1895a:171, 1897a:204). Hyperolius steindachneri steindachneri : Laurent (1950a:16, 1954a:79, 1961:75, 1964a:151). Hyperolius machadoi : Frade (1963:254), Cei (1977:17). Hyperolius (Hyperolius) steindachneri : Frétey et al.

(2011:33).

Global conservation status (IUCN):

Least Concern.

Global distribution: The species is known from the southern Democratic Republic of Congo to northern central-east of Angola and Zambia.

Ocurrences in Angola (Map 57): The species occurs in northern Angola. Lunda

Norte: “Dundo” [-7.36667, 20.83333] ( Laurent 1950a:16, 1954a:79, 80, 1964a:151);

“Lagoa Carumbo” [-7.74422, 19.95522] (Conradie et al. 2012a:3); “Camissombo”

[-8.15000, 20.65000] ( Laurent 1954a:80).

Malanje: “ Duque de Bragança ” [-9.10000,

15.95000] ( Bocage 1866a:55, 1866b:75,

MAP 57. Distribution of Hyperolius steindachneri in 1895a:171, 1897a:204; Loveridge 1936a:103; Angola.

Perret 1976a:25; Frost 1985:218, 2016; Poynton and Haacke 1993:15). Undetermined Locality: without precise locality ( Laurent 1954a:79); “areas of forest and savanna in the north and northeast of Angola ” ( Cei 1977:17); “plateaus regions” ( Cei 1977:17).

Taxonomic and distributional notes: The holotype was destroyed in the 1978 fire in the zoological collections in Museu Bocage. Laurent (1954a) described Hyperolius machadoi (Laurent, 1954) from “Camissombo” in Lunda Norte Porvince, but then later Laurent (1964a) synonymized it with H. steindachneri . The only recent report of this species was by Conradie et al. (2012a) who reported several specimens referred to as H. cf. steindachneri from Lagao Carumbo in Lunda Norte.

Hyperolius vilhenai Laurent, 1964 VILHENA’S REED FROG (Endemic)

Hyperolius vilhenai Laurent 1964a:155 . Holotype: MD 6213 (collector A. Barros Machado). Type locality: “galerie forestière de la rivière, Luíta, Poste de Cuílo ” [= Luíta], Lunda Norte Province, Angola.

Hyperolius vilhenai : Cei (1977:17), Frost

(1985:219, 2016), Schiøtz (1999:155); Channing

(2001:184).

Hyperolius (Hyperolius) vilhenai : Frétey et al.

(2011:33).

Global conservation status (IUCN):

Data Deficient.

Global distribution: The species is known only from northernmost Angola.

Ocurrences in Angola (Map 58): The species is known only from the type locality

(northern Angola). Lunda Norte: “galerie forestière de la rivière, Luíta, Poste de Cuílo”

[-8.03333, 19.41667] ( Laurent 1964a:155;

Frost 1985:219, 2016).

Taxonomic and distributional notes: If valid, the species is also likely to be found in the adjacent Democratic Republic of Congo MAP 58. Distribution of Hyperolius vilhenai in Angola. ( Frost 1985, 2016, Schiøtz 1999; Channing

2001).

Figures

MAP 43. Distribution of Hyperolius gularis in Angola.

MAP 56. Distribution of Hyperolius rhizophilus in Angola.