Leptopelis Günther, 1859

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 : 121-126

Genus Leptopelis Günther, 1859

Leptopelis anchietae (Bocage, 1873) ANCHIETA’S TREEFROG (Endemic)

Hylambates anchietae Bocage 1873b:226 . Holotype: MBL T.13-233 (Perret 1976a:22-23) (collector J. A. Anchieta ), destroyed 18 March 1978. Type locality: “l’intérieur de Mossamedes” ( Bocage 1873b:226), [= Namibe] Angola, later corrected to “Huíla” and “Caconda,” Huíla Province by Perret (1976a:22). Hylambates anchietae : Bocage (1895a:177, 1897a:205), Boulenger (1882:133, 1905:110).

Leptopelis anchietae : Schmidt (1936:131), Monard (1938:84), Loveridge (1957:317), Frade (1963:254), Laurent (1964a:147), Perret (1976a:22), Cei

(1977:17), Frost (1985:227, 2016), Schiøtz

(1999:306), Channing (2001:196), Channing et al. (2012:106), Frétey et al. (2011:35).

Leptopelis cf. anchietae : Conradie et al. (2016:10) .

Global conservation status (IUCN): Least Concern.

Global distribution: The species is known only from Angola.

Ocurrences in Angola (Map 67): The species is known from the type locality “Huíla” and from scattered localities throughout much of the western half of Angola. Lunda Sul: “Alto Chicapa” [-10.88333, 19.23333] ( Laurent 1964a:147). Bié: “Chitau” [-11.43333, 17.15000] ( Schmidt 1936:131); “Cubango basin (11)” [-13.69413, 17.06177] ( Conradie et al. 2016:8-9, 10); “Cubango basin (13)” MAP 67. Distribution of Leptopelis anchietae in Angola.

[-13.28061, 16.74722] ( Conradie et al. 2016:8-9, 10). Benguela: “Quindumbo” [-12.46667, 14.93333] ( Bocage 1895a:177, 1897a:205); “Ebanga” [-12.73333, 14.73333] ( Monard 1938:84); “Mount Chininga regions, between 1200 and 1700m ” (Chaning 2001:196). Huíla: “Caconda” [-13.73333, 15.06667] ( Bocage 1897a:205; Perret 1976a:12); “Kalukembé” [-13.78333, 14.68333] ( Monard 1938:84); “near Humpata” [-14.23814, 13.43331] Channing et al. (2012:106); “Kuvangu” [-14.46667, 16.30000] ( Monard 1938:84); “Huíla” [-15.05000, 13.55000] ( Bocage 1895a:177, 1897a:205; Perret 1976a:22). Namibe: “l’interieur de Mossamedes (Mossamedes)” [-15.20000, 12.15000] ( Bocage 1873b:226; Boulenger 1882:133; Loveridge 1957:317); “Chiyaka district” [vic. -15.08333, 12.73333] (Perret 1976:22). Undetermined Locality: “Western subregion and Angolan highlands ( Angola coastal or watersheds)” ( Frade 1963:254); “areas of forest and savanna in the north and notheast of Angola ” ( Cei 1977:17).

Taxonomic and distributional notes: Perret (1976a) suggested that Leptopelis anchietae (Bocage, 1873) is closely related to Leptopelis nordequatorialis Perret, 1966 and L. oryi Inger, 1968 .

Leptopelis aubryi ( Duméril, 1856) GABON FOREST TREEFROG

Hyla Aubryi Duméril 1856:561 . Syntypes: MNHN 833 , 1571 and 4603 (collector J. Aubry-Lecomte). Type locality: “ Gabon.”

Hylambates Aubryi : Peters (1877a:618), Bocage (1895a:181).

Leptopelis aubryi : Laurent (1954a:75), Loveridge (1957:318), Cei (1977:17), Schiøtz (1999:266), Frétey et al. (2011:35), Channing et al. (2012:107), Forst (2016).

Global conservation status (IUCN): Least Concern.

Global distribution: A forest-associated species distributed broadly in Central Africa, extending from southeastern Nigeria through

Cameroon and Gabon, east into the Central

African Republic, and south through the Democratic Republic of Congo to northern Angola.

Ocurrences in Angola (Map 68): The species occurs in the Cabinda enclave and in the extreme northeast of Lunda Norte

Province. Cabinda: “Chinchoxo” [-5.10000,

12.10000] ( Peters 1877a:618; Bocage

1895a:176); “cotê de Loango” ( Bocage

1895a:181); “ Cabinda enclave” (Channing et al. 2012:107). Lunda Norte: “Dundo”

[-7.36667, 20.83333] ( Laurent 1954a:75).

Taxonomic and distributional notes:

Many species of Leptopelis in Central Africa have been confused with Leptopelis aubryi ,

which has a broad distribution and occurs both within forests and along the margins of forests. MAP 68. Distribution of Leptopelis aubryi in Angola.

Leptopelis bocagii (Günther, 1865) BOCAGE’S TREEFROG

Cystignathus Bocagii Günther 1865a:481 , pl. 33, fig. 2. Holotype: not stated, destroyed by fire 18 March 1978 (collector F.P. Bayão). Two specimens received by BMNH from MBL may be types fide Frost (2016). Type locality: “Duque de Bragance” ( Günther 1865a:482), [= Calandula], Malanje Province, Angola;

Hylambates angolensis Bocage 1893:119 . Syntypes: MBL T.14-242-244 [3 specimens] (collector J.A. d’Anchieta), destroyed by fire 18 March 1978. Type locality: “Cahata” [= Caota], “Quindumbo”, “Quissange”, Benguela Province and “Caconda” Huíla Province, Angola .

Hylambates bocagei var. leucopunctata Ferreira 1904:113 . Syntype: MHNFCP 017324 (collected by F. Newton). Type locality: “Gumba” ( Ferreira 1904:113), [= Gumba, Serra de Selles] Kwanza Norte Province, Angola.

Cystignathus Bocagii : Bocage (1866a:54).

Hylambates bocagii : Boulenger (1882:133), Bocage (1895a:176, 1897a:205).

Hylambates angolensis : Bocage (1895a:179, 1896a:113, 1897a:205, 1897b:211), Perret (1976a:23).

Hylambates bocagei var. leucopunctata : Ferreira (1904:113).

Hylambates bocagei : Ferreira (1906:164).

Leptopelis angolensis : Schmidt (1936:131), Monard (1938:84), Frade (1963:254).

Leptopelis bocagei : Loveridge (1933:393, 1953a:342, 1957:317), Laurent (1950a:15, 1954a:76), Hellmich (1957a:30), Inger (1959:539, 541), Cei (1977:17), Schiøtz (1975:14), Frost (1985:228, 2016), Poynton and Broadley (1987:174), Schiøtz (1999:289), Channing (2001:198), Largen (1977:96, 2001:349), Frétey et al. (2011:36), Ceríaco et al. (2014a:25), Ceríaco et al. (2016b:25).

Leptopelis parbocagii : Poynton and Broadley (1987:171), Schiøtz and Van Daele (2003:146).

Leptopelis cf. bocagii : Amiet (2012:466).

Global conservation status (IUCN): Least Concern.

Global distribution: This savanna species has a broad distribution extending from Ethiopia in the north, south to Angola, northeastern Namibia, Zambia, Zimbabwe, and Mozambique. Ocurrences in Angola (Map 69): The species occurs from the western regions to northeastern Angola. Lunda Norte: “Dundo”

[-7.36667, 20.83333] ( Laurent 1954a:76;

Ceríaco et al. 2014a:26); “Muita (Luembe E)”

[-7.80000, 21.45000] ( Laurent 1950a:15,

1954a:76; Ceríaco et al. 2014a:26). Kwanza

Norte: “N’Golla Bumba” [-9.03333,

14.60000] ( Ferreira 1906:164; Ceríaco et al.

2014a:26); “ Rio Luinha ” [-9.26667, 14.53333]

( Ferreira 1906:164; Ceríaco et al. 2014a:26);

“Quilombo” [-9.33333, 14.90000] ( Ferreira

1906:164; Ceríaco et al. 2014a:26). Kwanza

Sul: “Gumba” [-11.26667, 14.28333] ( Ferreira

1904:113; Loveridge 1957:317; Ceríaco et al.

2014a:25). Malanje: “Duque de Bragance

(Duque de Brangança)” [-9.10000, 15.95000]

( Günther 1865a:482; Bocage 1866a:54, MAP 69. Distribution of Leptopelis bocagei in Angola. 1895a:176, 1897a:205; Loveridge 1933:393, 1953a:342, 1957:317; Schiøtz 1975:14; Perret 1976a:23; Frost 1985:228, 2016; Poynton and Broadley 1987:174; Largen 2001:349; Ceríaco et al. 2014a:27), “Cangandala National Park” [-9.84606, 16.72233] (Ceríaco et al. 2016b:25). Bié: “Chitau” [-11.43333, 17.15000] ( Schmidt 1936:131). Huambo: “Bimbi” [-11.81667, 15.83333] ( Monard 1938:84). Benguela: “Quibula” [-12.28333, 14.68333] ( Bocage 1893:119, 1895a:179, 1897a:205); “Cahata” [-12.35000, 14.81667] ( Bocage 1893:119, 1895a:179, 1897a:205; Perret 1976a:23); “Quissange” [-12.43333, 14.05000] ( Bocage 1893:119, 1895a:179, 1897a:205; Loveridge 1933:393; Perret 1976a:23); “Quindumbo” [-12.46667, 14.93333] ( Bocage 1893:119, 1895a:179, 1897a:205; Perret 1976a:23); “Entre Rios” [-13.01667, 14.63333] ( Hellmich 1957a:30); “Hanha” [-13.30000, 14.20000] ( Bocage 1896a:113; 1897b:211). Huíla: “Caconda”

[-13.73333, 15.06667] ( Bocage 1893:133, 1895a:179, 1897a:205; Loveridge 1957:317; Perret 1976a:23).

Taxonomic and distributional notes: Perret (1976a) noted that the synonymy of L. angolensis with L. bocagei was difficult to evaluate due to the differences in maturity between the type specimens of these two species, which were lost in the 1978 fire in Museu Bocage. Poynton and Broadley (1987) described a similar species, Leptopelis parbocagii based on five specimens collected at Mabwe on the eastern shore of Lake Upemba, in what is now the Democratic Republic of Congo. Because of their morphological similarity and overlapping distributions, the identification of these two species remains difficult ( Schiøtz 1999; Schiøtz and Van Daele, 2003). Leptopelis bocagii is a widespread species in southern and central Africa ( Channing 2001), though it may comprise several cryptic species ( Largen 1977; Amiet 2012). Ferreira (1904) described the variety leucopunctata , currently considered a synonym of Leptopelis bocagii (Günther, 1865) . Ceríaco et al. (2014a) provided more details about the taxonomic identity and nomenclature of L. bocagii and located a syntype from “Gumba, Sa [Serra] de Selles = Angola,” collected by Francisco Newton during the 1903–1905 expedition in Angola, in the Museu de História Natural da Universidade do Porto, Portugal.

Leptopelis cynnamomeus ( Bocage, 1893) ANGOLA FOREST TREEFROG

Hylambates cynnamomeus Bocage 1893:120 . Holotype: MBL T.16-250 (collector J.A. d’Anchieta), destroyed by fire 18 March 1978. Type locality: “Quilengues,” Huíla Province, Angola. Hylambates cynnamomeus : Bcoage (1895a:180, 1897a:205). Leptopelis viridis cinnamomeus : Laurent (1964a:148), Cei (1977:17). Leptopelis cynnamomeus : Schiøtz (1975:24), Frost (1985:229, 2016), Poynton and Broadley (1987:177),

Schiøtz (1999:299), Channing (2001:200), Schiøtz and Van Daele (2003:146), Frétey et al. (2011:36). Leptopelis bocagii : Loveridge (1957:317).

Global conservation status (IUCN): Least Concern.

Global distribution: The species is known from central Angola, northwestern Zambia, and southern Democratic Republic of Congo.

Ocurrences in Angola (Map 70): The species is known from the type locality “Quillengues” in Huíla, as well as from central and northeastern Angola , next to the boundary with the

Democratic Republic of Congo and Zambia.

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

( Laurent 1964a:148; Channing 2001:201).

Moxico: “Cazombo” [-11.88333, 22.91667]

( Laurent 1964a:148; Channing 2001:201).

Huíla: “Quillengues” [-14.06667, 15.08333]

( Bocage 1893:120, 1895a:180, 1897a:205;

Loveridge 1957:317; Schiøtz 1975:24; Perret

1976a:22; Frost 1985:229, 2016; Channing

2001:201).

Taxonomic and distributional notes:

Perret (1976a) examined the holotype from

Angola in the collection of Museu Bocage and condisered two paratypes from “Boloma” in

Guinée-Bissau to represent Leptopelis viridis

(Günther, “1868” 1869). MAP 70. Distribution of Leptopelis cynnamomeus in Angola.

Leptopelis jordani Parker, 1936 CONGULU FOREST TREEFROG (Endemic)

Leptopelis jordani Parker 1936:144 . Holotype: BMNH 1947.2.19.95 [formerly BMNH 1936.8.1.226] (collector K. Jordan). Type locality: “Congulu (700-800m)” [= Congulo] Kwanza Sul Province, Angola. Leptopelis jordani Parker, 1936 : Frost (1985:230, 2016), Frétey et al. (2011:36).

Global conservation status (IUCN):

Data Deficient.

Global distribution: This species is known only from Angola.

Ocurrences in Angola (Map 71): The species is known only from the type locality

“Congulu” in western Angola. Kwanza Sul:

“ Congulu (700-800m)” [-10.86667, 14.28333]

( Parker 1936:144; Frost 1985:230, 2016).

Taxonomic and distributional notes: In his description, Parker (1936) noted similarities to the widespread Leptopelis aubryi

( Duméril, 1856). There have been no recent records since the original description and the validity of this taxon remains in question. This species is not mentioned in Schiøtz (1999) or

Channing (2001).

MAP 71. Distribution of Leptopelis jordani in Angola.

Leptopelis marginatus (Bocage, 1895) QUISSANGUE FOREST TREEFROG (Endemic)

Hylambates marginatus Bocage 1895a:178 . Holotype: MBL T.218-241 (collector J.A. d’Anchieta), destroyed by fire 18 March 1978. Type locality: “Quissange, dans l’intérieur de Benguella” [= Quissange] Benguela Province, Angola.

Hylambates marginatus : Bocage (1897a:205), Cei (1977:17).

Leptopelis bocagii : Loveridge (1933:393).

Leptopelis anchietae : Loveridge (1957:317).

Leptopelis marginatus : Perret (1976a:23), Forst

(1985:230, 2016), Frétey et al. (2011:36).

Global conservation status (IUCN):

Data Deficient.

Global distribution: This species is known only from Angola.

Ocurrences in Angola (Map 72): The species is known only from the type locality in estern Angola. Benguela: “Quissange”

[-12.43333, 14.05000] ( Bocage 1895a:179,

1897a:205; Loveridge 1933:393, 1957:317;

Perret 1976a:23, Frost 1985:230, 2016).

Taxonomic and distributional notes:

Perret (1976a) concluded that L. marginatus was a valid species, in contrast to Loveridge

(1933) who recognized it as a synonym of

Leptopelis bocagii (Günther, 1865) . There have MAP 72. Distribution of Leptopelis marginatus in been no other recent records and the holotype Angola .

was destroyed in the 1978 fire in Museu Bocage. This species is not mentioned Schiøtz (1999) or Channing (2001). This should be recognized as a nomen dubium.

Leptopelis notatus ( Peters, 1875) COMMON FOREST TREEFROG

Hylambates notatus Peters 1875:205 , pl. 2, figs. 1, 1a. Holotype: ZMB 8471 (collector R. Buchholz) fide

Bauer et al. (1995:43). Type locality: “Cameruns” ( Peters 1875:206), [= Cameroon]. Leptopelis tessmanni : Laurent (1950a:15), Frade (1963:254). Leptopelis notatus : Laurent (1964a:147), Cei (1977:17), Schiøtz (1999:264), Frétey et al. (2011:36), Channing et al. (2012:116), IUCN SSC Amphibian Specialist Group (2013a), Frost (2016).

Global conservation status (IUCN): Least Concern.

Global distribution: This is a widespread species occurring in and near forests extending from Cameroon, Gabon, Republic of Congo,

the western Democratic Republic of Congo,

and into northernmost Angola.

Ocurrences in Angola (Map 73): The species occurs in the northern Angola. Lunda

Norte: “Dundo” [-7.36667, 20.83333] ( Laurent 1964a:147); “Mussalomuca” [-7.43333,

20.45000] ( Laurent 1964a:147); “rive de la

Tchihumbwe, 40 km à l’est de Dundo”

[-8.01667, 19.31667] ( Laurent 1950a:15).

Lunda Sul: “Alto Cuílo” [-10.01667,

19.55000] ( Laurent 1964a:147). Undetermined Locality: “areas of forest and savanna in the north and northeast of Angola ” ( Cei

1977:17).

Taxonomic and distributional notes: MAP 73. Distribution of Leptopelis notatus in Angola. Frétey et al. (2011) questioned the presence of this species in Angola.

Leptopelis viridis (Günther, “1868” 1869) RUSTY FOREST TREEFROG

Hylambates viridis Günther “1868” 1869:487. Holotype: BMNH 1947.2 .10.23 (purchased from Mr. Stevens),

formerly BMNH 65.5 .3.63. Type locality: “ West Africa ” .

Hylambates viridis : Bocage (1873b:226, 1895a:176, 1897a:205), Boulenger (1882:134).

Leptopelis viridis : Frétey et al. (2011:37), IUCN SSC Amphibian Specialist Group (2013b), Frost (2016).

Global conservation status (IUCN): Least Concern.

Global distribution: This species is found in savannas across western and Central Africa, extending south into Angola.

Ocurrences in Angola (Map 74): The species has been recorded from Malanje Province. Malanje: “Duque de Bragança” [-9.10000, 15.95000] ( Bocage 1873b:206, 1895a:179, 1897a:205).

Taxonomic and distributional notes: Bocage (1873 b, 1895a, 1897a) cited a single juvenile from “Duque de Bragança” collected by Bayão whereas Boulenger (1882) cited another individual from Angola collected by J.J. Monteiro, although without locality. Many recent summaries of the distribution of this species, including Schiøtz (1999), IUCN SSC Amphibian Specialist Group (2013b), and Frost (2016), have excluded Angola from its distribution though Frétey et al. (2011)