Afrotyphlops Broadley and Wallach, 2009

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 : 292-299

Genus Afrotyphlops Broadley and Wallach, 2009

Afrotyphlops angolensis (Bocage, 1866) ANGOLA BLIND SNAKE

Onychocephalus angolensis Bocage 1866a:46 , 1866b:65. Holotype: MBL T79.1134 (collector F.A.P. Bayão),

destroyed by fire on 18 March 1978. Type locality: “le district du Duque de Bragança ” [= Calandula] ,

Malanje Province, Angola. Onychocephalus Kraussii : Bocage (1873a:252). Onychocephalus angolensis : Bocage (1879b:95). Typhlops congicus : Bocage (1895a:63). Typhlops punctatus punctatus : Loveridge (1957:242). Typhlops angolensis adolfi : Laurent (1964a:88). Typhlops angolensis : Laurent (1964c:424), Roux-Estève (1974a:490; 1974b:46), McDiarmid et al. (1999:90),

Spawls et al. (2004:288). Typhlops angolensis angolensis : Manaças (1973:189) Rhinotyphlops angolensis : Broadley and Cotterill (2004:44), Chirio and LeBreton (2007:332). Afrotyphlops angolensis : Broadley and Wallach (2009:29), Ceríaco et al. (2014b:672), Wallach et al.

(2014:13), Hedges et al. (2014:20).

Global conservation status (IUCN): Not Evaluated.

Global distribution: The species occurs in both forest and savanna from Cameroon south through the Congo Basin to northern Angola and northeast Zambia and east to Uganda and Kenya.

Ocurrences in Angola (Map 253): The species occurs in the northeastern Angola. Kwanza Norte: “Canzele” [-8.30000, 15.18333] ( Laurent 1964c:424). Malanje: “Duque de Bragança” [-9.10000, 15.95000] ( Bocage 1866a:46, 1866b:65, 1873a:252; Loveridge 1957:242; Laurent 1964c:424; Roux-Estève 1974a:490, 1974b:57; Broadley and Wallach 2009:29; Wallach et al. 2014:13). Lunda Norte: “Dundo” [-7.36667, 20.83333] ( Laurent 1964b:88; Roux-Estève

1974b:57); “Cassange” [-9.58333, 17.86667]

( Bocage 1879b:95). Moxico: “Calombe, Luso”

[-11.83333, 19.93333] ( Manaças 1973:189).

Huambo: “ Nova Lisboa ” [-12.76667,

15.73333] ( Roux-Estève 1974b:57).

Taxonomic and distributional notes: The species was first named by Bocage (1866a),

who provided a diagnosis and description, but the status of the species as new was indicated conditionally, as “ Onychocephalus angolensis

Nov. sp.?” The name was used without such a caveat in a subsequent publication Bocage

(1866b). Both publications appeared in the

November 1866 issue of the same journal. See

Roux-Estève (1974b), McDiarmid et al. (1999)

and Broadley and Wallach (2009) for comprehensive chresonymies and details of distribution elsewhere in Africa. MAP 253 Distribution of Afrotyphlops angolensis in Angola.

Afrotyphlops anomalus (Bocage, 1873) ANGOLAN GIANT BLIND SNAKE (Endemic)

Onychocephalus anomalus Bocage 1873a:248 , pl. 1, fig. 3. Syntypes: MBL 1177 A–D (four specimens), (collector J.A. d’Anchieta and J.J. da Graça), destroyed by fire 18 March 1978. Type locality: “Huilla, l’intérieur de Mossamedes” [= Huíla], Huíla Province, Angola.

Typhlops (Onychocephalus) Anchietae Bocage 1886a:172 . Holotype: MBL 1871 , (collector J. Anchieta), destroyed by fire 18 March 1978. Type locality: “Huilla” [= Huíla ], Huíla Province, Angola.

Typhlops anchietae : Boulenger (1893:40, 1915:197), Bocage (1895a:63, 1897a:198), Monard (1937b:103).

Typhlops anomalus : Boulenger (1893:47), Bocage (1895a:70), Ferreira (1897b:243)

Typhlops anomalus : Monard (1937b:103, 105).

Rhinotyphlops anomalus : Roux-Estève (1974a:495, 1974b:193), McDiarmid et al. (1999:78).

Megatyphlops anomalus : Broadley and Wallach (2009:48), Wallach et al. (2014:423).

Afrotyphlops anomalus : Hedges et al. (2014:20).

Global conservation status (IUCN): Not Evaluated.

Global distribution: The species is endemic to Angola.

Ocurrences in Angola (Map 254): The species is restricted to the southwestern areas of the country. Cabinda: “Loango (?)” [-5.15000, 12.16667] ( Boulenger 1893:47, 1915:197; Monard 1937b:103). Huambo: “Bimbi” [-11.81667, 15.83333] ( Monard 1937b:103, 105; Broadley and Wallach 2009:48). Benguela: “Quibula” [-12.28333, 14.68333] ( Bocage 1895a:70; Monard 1937b:103; Broadley and Wallach 2009:48); “Cahata” [-12.35000, 14.81667] ( Bocage 1895a:70; Monard 1937b:103; Broadley and Wallach 2009:48); “Quindumbo [-12.46667, 14.93333] ( Bocage 1895a:70; Broadley and Wallach 2009:48); “Benguella” [-12.58333, 13.41667] ( Bocage 1895a:70; Boulenger 1915:197; Monard 1937b:103); “Ebanga” [-12.73333, 14.73333] ( Monard 1937b:103, 105; Broadley and Wallach 2009:48). Huíla: “Huilla” [-15.05000, 13.55000] ( Bocage 1873b:248, 252, 1886a:172, 1895a:63, 70, 1897a:198; Monard 1937b:103; Roux-Estève 1974b:194; Broadley and Wallach 2009:48; Wallach et al. 2014:423); “Caconda” [-13.73333, 15.06667] ( Bocage 1895a:70; Ferreira 1897:243; Monard 1937b:103; Roux-Estève 1974b:194; Broadley and Wallach 2009:48); “Kuvangu (Vila-da-Ponte)” [-14.46667, 16.30000] ( Monard 1937b:103, 105; Broadley and Wallach 2009:48). Namibe: “Biballa” [-14.76667, 13.36667] ( Bocage 1895a:70; Monard 1937b:103; Broadley and Wallach 2009:48);

“ Chiyaka district ” [vic. -15.08333, 12.73333]

( Roux-Estève 1974b:194; Broadley and Wallach 2009:48); “Mossamedes” [-15.20000,

12.15000] ( Bocage 1873b:248; Boulenger

1893:47).

Taxonomic and distributional notes:

Hedges et al. (2014) moved all species previously allocated to Megatyphlops Broadley and

Wallach, 2009 to Afrotyphlops based on their phylogenetic analysis, which demonstrated that members of the former group are embedded within a clade containing the type specimen of the latter. Boulenger (1893, 1915)

identified a specimen from “Loango,” a former kingdom within Cabinda Province as

M. anomalus . However, this is certainly a misidentification or error in locality, as Roux- MAP 254. Distribution of Afrotyphlops anomalus in Angola.

Estève (1974a) demonstrated that this species was only reliably known from the mountainous regions of Benguela and Huíla, south of the Cuanza River. Wallach et al. (2014) also listed Bié and Cuanza Sul in the range of the species, but this does not appear to be based on previously published records. McDiarmid et al. (1999) included northern Namibia in the stated distribution of this species, but we are not aware of any confirmed specimens outside of Angola. See Broadley and Wallach (2009) for chresonymy and maps of global distribution.

Afrotyphlops lineolatus ( Jan, 1864) COMMON LINED BLIND SNAKE

Typhlops (Ophthalmidion) lineolatus Jan 1864:24 . Holotype, UUMZ 725 (collector A. Afzel), lost fide RouxEstève (1974b) and Hahn (1980). Type locality: “Sierra-Leona” [= Sierra Leone]. Typhlops bocagei Ferreira 1904:114 . Syntypes: MBL specimen numbers not known (collector F. Newton)

(destroyed July 1943 during WWII fide Broadley and Wallach 2009:40). Type locality: “Cabicula, Bom

Jesus (margens do Quanza)” [= Bom Jesus], Bengo Province, Angola. Typhlops Boulengeri Bocage (1893:117) . Lectotype: BMNH 1946.1 .11.18, formerly BMNH 1893.12 .27.12

(collector J. Anchieta) designated by Broadley and Wallach (2009:40). Type locality: “Quindumbo, dans l’intérieur de Benguella” [= Quindumbo], Benguela Province, Angola. Onychocephalus liberiensis (part): Bocage (1866a:46, 1873a:252). Onychocephalus lineolatus : Bocage (1873a:252, 1866b:65). Typhlops eschrichtii (part): Günther (1876b:678). Typhlops (Ophthalmidion) Eschrichtii var. lineolata : Peters (1877a:614). Typhlops (Ophthalmidion) Eschrichtii var. intermedia : Peters (1877a:614). Typhlops (Ophthalmidion) Eschrichtii (part): Peters (1881:147). Typhlops (Ophthalmidion) Kraussii (part): Bocage (1887a:180). Typhlops lineolatus : Boulenger (1893:43), Roux-Estève (1974b:76), McDiarmid et al. (1999:107), Spawls et al. (2004:290). Typhlops punctatus var. intermedia (part): Bocage (1895a:66). Typhlops punctatus var. lineolata : Bocage (1895a:66). Typhlops punctatus var. lineolatus : Bocage (1896a:112), Ferreira (1903:9). Typhlops punctatus (part): Boulenger (1900a:50, 1905:112), Monard (1937b:103, 104), Themido (1941:9).

Typhlops boulengeri : Bocage (1895a:64, 1897a:198), Ferreira (1900a:50, 1906:167), Boulenger (1915:196),

Monard (1937b:103, 104), Laurent (1964a:89). Typhlops punctatus punctatus (part): Parker (1936:120), Mertens (1937a:11, 1938a:438), Laurent (1950a:7),

Hellmich (1957a:70), Loveridge (1957:242). Typhlops bocagei : Monard (1937b:103). Typhlops boulengeri boulengeri : Laurent (1964c:414). Typhlops lineolatus lineolatus : Roux-Estève (1974a:492). Rhinotyphlops lineolatus : Broadley and Cotterill (2004:45). Rhinotyphlops lineolatus lineolatus : Chirio and LeBreton (2007:338). Afrotyphlops lineolatus : Wallach and Broadley (2009:41), Wallach et al. (2014:14), Hedges et al. (2014:20).

Global conservation status (IUCN): Not Evaluated.

Global distribution: The species is widely distributed in the savannas of sub-Saharan Africa from Senegal east to Sudan and Ethiopia and south to northern Tanzania, Katanga ( Democratic Republic of Congo) and Angola as far south as Benguela.

Ocurrences in Angola (Map 255): The species occurs mainly in western Angola, however there are records in Lunda Norte Province.

Cabinda: “Chinchoxo” [-5.10000, 12.10000]

( Peters 1877a:614); “Landana” [-5.21667,

12.15000] ( Roux-Estève 1974b:104). Zaire :

“ S. Salvador do Congo ” [-6.26667, 14.23333]

( Bocage 1895a:66, 1887a:180; Monard

1937b:103; Roux-Estève 1974b:104). Lunda

Norte: “rivière Muari (afluente de la Luachimo), dans le environs de Dundo” [-7.28333,

20.93333] ( Laurent 1950a:7); “Dundo”

[-7.36667, 20.83333] (Laurent 1950:7,

1964a:89); “Muita” [-7.80000, 21.45000]

( Laurent 1950a:7). Lunda Sul: “Mutianvo”

[-11.45000, 19.33333] ( Themido 1941:9).

Malanje: “ Duque de Bragança ” [-9.10000,

15.95000] ( Bocage 1873a:252; Bocage MAP 255. Distribution of Afrotyphlops lineolatus in Angola.

1895a:66; Monard 1937b:103); “Cassange”

[-9.58333, 17.86667] ( Bocage 1895a:66; Monard 1937b:103); “ Malanje ” [-9.55000, 16.35000] ( Peters 1881:147). Bengo: “Cabicula, Bom Jesus (margem do Quanza)” [-9.16667, 13.56667] ( Ferreira 1904:114; Monard 1937b:103; Loveridge 1957:242). Kwanza Norte: “Golungo Alto” [-9.13333, 14.76667] ( Boulenger 1905:112; Monard 1937b:103; Roux-Estève 1974b:104); “Cambondo” [-9.15963, 14.65827] ( Ferreira 1906:167; Monard 1937b:103, 104); “ Rio Luinha” [-9.26667, 14.53333] ( Ferreira 1906:167; Monard 1937b:103, 104; Wallach and Broadley 2009:41); “N’dalla Tando, Cazengo” [-9.30000, 14.91667] ( Ferreira 1903:9; Monard 1937b:103, Roux-Estève 1974b:104; Wallach and Broadley 2009:41). Kwanza Sul: “Quirimbo” [-10.68333, 14.26667] ( Parker 1936:120); “Congulu” [-10.86667, 14.28333] ( Parker 1936:120). Benguela: “Quindumbo” [-12.46667, 14.93333] ( Bocage 1893:117, 1895a:64, 1897a:198; Loveridge 1933:216, 1957:241; Monard 1937b:103, 104; Laurent 1964a:89, 1964c:414; Roux-Estève 1974b:104; Wallach and Broadley 2009:41); “Benguella” [-12.58333, 13.41667] ( Boulenger 1915:196; Loveridge 1957:242); “Ebanga” [-12.73333, 14.73333] ( Monard 1937b:103, 105); “Entre Rios” [-13.01667, 14.63333] ( Hellmich 1957a:70); “Cubal” [-13.03333, 14.25000] ( Mertens 1937a:11, 1938a:438); “Hanha” [-13.30000, 14.20000] ( Bocage 1896a:112; Roux-Estève 1974b:104). Undetermined Locality: ( Günther 1876b:678; Ferreira 1900:50);. “Cuango” ( Peters 1881:147) (Malanje Province impossible to georeference: See History Section for more detailed information).

Taxonomic and distributional notes: There was long confusion between this taxon and Afrotyphlops punctatus ( Leach, 1819) as reflected in the complex synonymies/chresonymies of these two taxa ( McDiarmid et al. 1999). Authorship of this species is sometimes attributed to Jan (1863) (e.g., McDiarmid et al. 1999), however, this use is as a nomen nudum. See Broadley and Wallach (2009) for comprehensive chresonymy and distribution in eastern and central Africa.

Afrotyphlops mucruso ( Peters, 1854) ZAMBEZI BLIND SNAKE

Onychocephalus mucruso Peters 1854:621 . Lectotype: ZMB 3963 (collector W.C.H. Peters), designated by Loveridge (1933:216). Type locality: “Macanga” ( Peters 1854:621), [= Makanga], Mozambique.

Typhlops mucruso : Boulenger (1893:46), Bocage (1895a:67).

Typhlops schlegelii mucruso : Loveridge (1933:216, 1957:241).

Typhlops schlegeli mucruso : Laurent (1950a:7, 1964a:90),

Rhinotyphlops schlegelii dinga (part): Roux-Estève (1974b:164).

Typhlops schlegelii mucruso : Broadley (1990:47).

Rhinotyphlops schlegelii (part): McDiarmid et al. (1999:84).

Rhinotyphlops mucruso : Spawls et al. (2004:292), Broadley and Cotterill (2004:45).

Megatyphlops mucruso : Broadley and Wallach (2009:52), Bates et al. (2014:313), Wallach et al. (2014:423).

Afrotyphlops mucruso : Hedges et al. (2014:20).

Global conservation status (IUCN): Not

Evaluated.

Global distribution: The species is known from the savannas and forest-savanna mosaic from East Africa, from coastal Kenya south to central Mozambique, Zimbabwe and

Limpopo Province, South Africa, and west through the southern provinces of the Democratic Republic of Congo to northeast Angola.

Ocurrences in Angola (Map 256): The species occurs in the extreme northeast of the country, in Lunda Norte and Sul provinces.

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

( Laurent 1950a:7; Roux-Estève 1974b:178);

“Muita” [-7.80000, 21.45000] ( Laurent

1950a:7; “Camisombo” [-8.15000, 20.65000]

( Laurent 1964a:89); “Calonda” [-8.41667,

20.53333] ( Laurent 1964a:89). Lunda Sul: MAP 256. Distribution of Afrotyphlops mucruso in “Alto Chicapa” [-10.88333, 19.23333] (Lau- Angola.

rent 1964a:89).

Taxonomic and distributional notes: See Roux-Estève (1974b), Bauer et al. (1995, 2002) and McDiarmid et al. (1999) for discussions of the original type series of Onychocephalus mucruso . Hedges et al. (2014) moved all species previously allocated to Megatyphlops Broadley and Wallach, 2009 to Afrotyphlops based on their phylogenetic analysis, which demonstrated that members of the former group are embedded within a clade containing the type specimen of the latter. Previously treated as a subspecies, Rhinotyphlops schlegelii mucruso ( Peters, 1854) , but elevated to full species status and placed in the new genus Megatyphlops by Broadley and Wallach (2009). The status of large typhlopids in Angola previously assigned to Megatyphlops by Broadley and Wallach (2009) as mucruso and schlegelii remains problematic. See Broadley and Wallach (2009) for comprehensive chresonymy and maps of global distribution.

Afrotyphlops schlegelii ( Bianconi, 1849) SCHLEGEL’S GIANT BLIND SNAKE Typhlops schlegelii Bianconi 1849:183 , pl. 6, figs. 2, 2a-2d. Holotype: MZUB specimen numbers not known

(collector C. Fornasini ). Type locality: “Mozambico” [= Mozambique] (Inhambane, Mozambique fide

Roux-Estève 1974b). Typhlops (Onychocephalus) humbo Bocage 1886a:171 . Syntypes: MBL 1887 [2 specimens], destroyed by fire

18 March 1978 (collector J.A. d’Anchieta). Type locality: “Quissange” Benguela Province, Angola. Onychocephalus Petersii Bocage 1873a:248 , pl. 1, fig. 3. Syntypes: MBL 1868 (2 specimens), destroyed by fire 18 March 1978 (collector J. Anchieta). Type locality: “Biballa” [= Bibala] Namibe Province, Angola. Onychocephalus Schlegelii : Bocage (1873a:250). Typhlops Petersii : Bocage (1886a:172, 1895a:68, 1897a:199). Typhlops (Onychocephalus) humbo Bocage 1886a:171 . Syntypes: MBL 1887 (2 specimens) (collector

J. Anchieta), destroyed by fire 18 March 1978. Type locality: “Quissange” Benguela Province, Angola. Typhlops humbo : Bocage (1887c:210, 1895a:66, 1897a:198), Boulenger (1893:46). Typhlops hottentotus : Bocage 1893:117. Holotype: MBL 1867 (collector J. Anchieta), destroyed by fire 18

March 1978. Type locality: “Humbe” Cunene Province, Angola. Typhlops hottentotus : Bocage (1895a:69, 1897a:198), Boulenger (1896:588). Typhlops schlegeli mucruso : Loveridge (1933:216, 1957:241), Mertens (1938a:438). Typhlops mucruso ( = humbo , hottentotus , petersii ): Monard (1937b:103). Rhinotyphlops schlegelii petersii : Roux-Estève (1974a:495; 1974b:166), Branch (1998:54). Typhlops schlegelii petersii : Broadley (1990:48). Rhinotyphlops schlegelii (part): McDiarmid et al. (1999:84). Megatyphlops schlegelii : Broadley and Wallach (2009:48), Bates et al. (2014:314), Wallach et al. (2014:423). Afrotyphlops schlegelii : Hedges et al. (2014:20)

Global conservation status (IUCN): Not Evaluated.

Global distribution: The species is known from southern Mozambique, northern KwaZulu-Natal to Swaziland, through the northeastern provinces of South Africa, central Botswana,

northern Namibia and southwestern Angola.

Ocurrences in Angola (Map 257): The species occurs in southwestern Angola.

Benguela: “Quissange” [-12.43333, 14.05000]

( Bocage 1886a:171, 1887c:210, 1895a:66,

1897a:198; Boulenger 1896:588; Loveridge

1933:216; Monard 1937b:103; Roux-Estève

1974:495, 1974b:180; Broadley and Wallach

2009:51); “Benguella” [-12.58333, 13.41667]

( Boulenger 1893:46), “Caimbambo”

[-13.01667, 14.01667] (Broadley and Wallach

2009:51); “Cubal” [-13.03333, 14.25000]

(Mertens 1938:438). Huíla: “Caconda”

[-13.73333, 15.06667] ( Bocage 1895a:68;

Monard 1937b:103); “ 10 km SE of Joao de

Almeida” [-15.18333, 13.68333] (Broadley MAP 257. Distribution of Afrotyphlops schlegelii in and Wallach 2009:51). Namibe: “ Biballa ” Angola .

[-14.76667, 13.36667] ( Bocage 1873a:249, 252, 1886a:172, 1895a:68, 1897a:199; Loveridge 1933:216; Monard 1937b:103). Cunene: “Humbe, sur les bords du Cunene (Humbe)” [-16.68333, 14.90000] ( Bocage 1893:117, 1895a:69; Boulenger 1896:588; Monard 1937b:103; Broadley and Wallach 2009:51); “Erickson’s Drift, Cunene river” [-17.26944, 14.525 E] (Broadley and Wallach 2009:51).

Taxonomic and distributional notes: There are three juvenile specimens referable to this species that are extant in the Bianconi collection of Mozambiquan reptiles in Bologna, but none of these seem to be consistent with the type specimen. The date of publication is often confused because the description appeared in the journal issue for August and September 1848, but was not published until April 1849, and because Bianconi (1850:183, pl. 6, fig. 2) published a second description, marked “Nobis” of the same animal. Further, Bianconi (1849) notes that he had described this species, and another, in the session of 1847, but this refers to the oral sessions of the Società Agraria, e dell’ Accademia delle Scienze dell’ Istituto di Bologna, for which the Nuovi Annali delle Scienze Naturali was the publication outlet. “ Hedges et al. (2014) moved all species previously allocated to Megatyphlops Broadley and Wallach, 2009 to Afrotyphlops based on their phylogenetic analysis, which demonstrated that members of the former group are embedded within a clade containing the type specimen of the latter. Until recently, four subspecies were recognized under the name Rhinotyphlops schlegelii (brevis, schlegelii , petersii , mucruso ) following Roux-Estève (1974b) and Hahn (1980). Currently Onychocephalus petersii (Bocage, 1873) is placed in the synonymy of Afrotyphlops schlegelii ( Bianconi, 1849) , whereas the others are recognized as valid species ( Roux-Estève 1974a; 1974b; Broadley 1990; Branch 1998; Broadley and Wallach 2009). The complex nomenclatural history of this and related taxa, including the interpretation of type specimens has been reviewed by McDiarmid et al. (1999). See Broadley and Wallach (2009) for comprehensive chresonymy and maps of global distribution.

Afrotyphlops schmidti ( Laurent, 1956) SCHMIDT’S BLIND-SNAKE

Typhlops schmidti Laurent 1956:71 , figs. 9–11, pl. 8, fig. 4. Holotype: MRAC 17996 (collector not mentioned). Type locality: “Nyunzu, Terr. D’Albertville, Tanganika” [= Nyunza, Tanganyika Province (formerly Katanga Province), Democratic Republic of Congo fide Wallach and Broadley 2009:43].

Typhlops schmidti : Laurent (1964a:89), McDiarmid et al. (1999:119).

Rhinotyphlops schmidti : Broadley et al. (2003:42),

Broadley and Cotterill (2004:45)

Afrotyphlops schmidti : Broadley and Wallach

(2009:43), Wallach et al. (2014:15), Hedges et al.

(2014:20).

Global conservation status (IUCN): Not

Evaluated.

Global distribution: A savanna species distributed in eastern Angola, through northern

Zambia and southern Democratic Republic of

Congo (Katanga provinces).

Ocurrences in Angola (Map 258): The species occurs in eastern Angola. Moxico:

“environs du lac Calundo, village du chef Sá-

MAP 258. Distribution of Afrotyphlops schmidti in Mussamba (environs du lac Calundo)” Angola.

[-11.80000, 20.86667] ( Laurent 1964a:89; Broadley and Wallach 2009:43); “Cazombo” [-11.88333, 22.91667] ( Laurent 1964a:89; Broadley and Wallach 2009:43).

Taxonomic and distributional notes: The type locality of Typhlops schmidti in the Tanganika district (now province) of the Democratic Republic of Congo, has been misinterpreted by some subsequent authors to refer to Tanzania (e.g., McDiarmid et al. 1999). Laurent (1964a) was the first to cite this species from Angola. See Broadley and Wallach (2009) for chresonymy and maps of global distribution.