Naja Laurenti, 1768

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 : 368-375

Genus Naja Laurenti, 1768

Naja anchietae Bocage, 1879 ANCHIETA’S COBRA

Naja Anchietae Bocage 1879a:98 , 1879c:89. Lectotype: MBL 1987 (collectors H.C. Capello and R. Ivens), designated by Broadley (1995:31), destroyed by fire 18 March 1978. Type locality: “Caconda,” Huíla Province, Angola.

Naja Anchietae : Bocage (1879c:98, 1895a:133, 1897a:202).

Naja anchietae : Ferreira (1900ab:134), Monard (1937b:138), Bogert (1940:90), Frade (1963:252), Broadley (1968c:7), Broadley and Cotterill (2004:47), Broadley and Wüster (2004:101), Dobiey and Vogel (2007:63), Ceríaco et al. (2014b:671), Ceríaco et al. (2016b:87).

Naja haje anchietae : Mertens (1937:23), Bogert (1940:90), Laurent (1964a:118), Manaças (1981:26), Broadley (1966c:23), Spawls and Branch (1995:70), Trape et al. (2009:20).

Naja anulifera ancheitae : Broadley (1995:31).

Naja (Uraeus) anchietae : Wallach et al. (2009:31).

Uraeus anchietae : Wallach et al. (2014:762).

Global conservation status (IUCN): Not Evaluated.

Global distribution: The species is known from western parts of southern Africa, from Angola and northern Namibia through western Zambia and northern Botswana to northwestern Zimbabwe and Lake Bangweulu, northeastern Zambia.

Ocurrences in Angola (Map 331): The species occurs in mostly in the southern Angola however, there are some records from more northern provinces. Lunda Sul: “Alto Cuílo”

[-10.01667, 19.55000] ( Laurent 1964a:118,

Manaças 1981:26); “Alto Chicapa”

[-10.93333, 19.15000] ( Laurent 1964a:118,

Manaças 1981:26). Malanje: “Capanda”

[-9.72841, 15.34585] (Ceríaco et al.

2014b:671); “Cangandala National Park”

[-9.84606, 16.72233] (Ceríaco et al. 2016b:87).

Benguela: “Hanha” [-13.30000, 14.20000]

( Ferreira 1900b:134; Manaças 1981:26).

Huíla: “Caconda” [-13.73333, 15.06667]

( Bocage 1879a:98, 1879c:89, 1895a:133,

1897a:202; Ferreira 1900b:134; Manaças

1981:26; Broadley 1995:31); “Kalukembé”

[-13.78333, 14.68333] ( Monard 1937b:138;

MAP 331. Distribution of Naja anchietae in Angola. Manaças 1981:26); “Vila da Ponte” [-14.46667, 16.30000] (Manaças 1981:26); “Kuvangu” [-14.46667, 16.30000] ( Monard 1937b:138; Manaças 1981:26); “Capelongo” [-14.91667, 15.08333] ( Bogert 1940:90; Manaças 1981:26); “Huilla” [-15.05000, 13.55000] ( Bocage 1895a:133, 1897a:202; Ferreira 1900b:134; Manaças 1981:26); “vu l’espèce dans la région du Kuluï supérieur” [-15.41667, 15.73333] ( Monard 1937b:138). Cunene: “Mupa” [-16.18333, 15.75000] ( Monard 1937b:138; Manaças 1981:26); “Humbe” [-16.68333, 14.90000] ( Bocage 1895a:133, 1897a:202; Ferreira 1900b:134; Manaças 1981:26). Undetermined Locality: “Rio Cuando” ( Bocage 1895a:133, 1897a:202; Manaças 1981:26).

Taxonomic and distributional notes: Wallach et al. (2014) attributed the lectotypoe designation for Naja anchietae to Broadley and Wallach (2004), however, Broadley (1995) had already refered to MBL 1987 as the lectotype a decade earlier. Mertens (1937a), Bogert (1940) and subsequent authors treated Naja anchietae as a subspecies of Naja haje ( Linnaeus, 1758) , until Broadley (1995) included it as a subspecies of Naja annulifera Peters, 1854 . Recent studies suggest that these two taxa are morphologically and genetically distinct evolutionary lineages that should be considered as two separate species (Broadley and Wüster 2004). Wallach et al. (2009) assigned the non-spitting African cobras to the subgenus Uraeus Wagler, 1830 and subsequently treated Uraeus as a full genus ( Wallach et al. 2014). Most literature has considered N. anchietae to be restricted in Angola to the south (e.g., Broadley 1966c, 1968c, 1995; Spawls and Branch 1995; Broadley and Wüster 2004; Broadley and Cotterill 2004; Dobiey and Vogel 2007; Trape et al. 2009:20; Wallach et al. 2014) but there are now numerous records from more northerly areas ( Laurent 1964a; Manaças 1981; Ceríaco et al. 2014b, 2016b).

Naja annulata Buchholz and Peters, 1876 RINGED WATER COBRA

Naja annulata Buchholz and Peters 1876 in Peters 1876:119. Holotype: ZMB 8772 (collector R. Buchholz). Type locality: “Dorfe Mbusu (Eliva Sonangeam am Ogowe)” [= Eliva Sonange, near Ogooué-Maritime], Gabon.

Boulengerina annulata : Spawls and Branch (1995:55), Broadley et al. (2003:106), Nagy et al. (2005:224), Wüster et al. (2007:445), Wallach et al. (2014:119).

Boulengerina annulata annulata : Dobiey and Vogel (2007:37).

Naja (Boulengerina) annulata : Wallach et al. (2009:31), Branch and Conradie (2015:200).

Global conservation status (IUCN): Not Evaluated.

Global distribution: The the nominate subspecies is known from southern Cameroon to the southern Central African Republic and throughout the northern and central portions of the Democratic Republic of Congo. It extends southwards through People’s Republic of Congo, Gabon and Equatorial Guinea, and reaches Angola along the lower Zaire River. The subspecies N. a. stormsi (Dollo, 1886) occurs extralimitally in the areas around Lakje Tanganyika and Lake Kivu.

Ocurrences in Angola (Map 332): The species presumably occurs in the Cabinda enclave and in the northernmost portions of the country. Lunda Norte: “Along the edge of the Luele River, near the base camp, 5 km from the Carumbo Lagoon” [-7.75294, 19.95672] (Branch and Conradie 2015:200) .

Taxonomic and distributional notes: Water cobras ( Boulengerina ) have long been known to make Naja paraphyletic (Slowinski and Keogh 2000, Nagy et al. 2005, Wüster et al. 2007, Wallach et al. 2009). Wüster et al. (2007) recovered the species Paranaja multifasciata ( Werner, 1902) as the sister taxon of Boulengerina annulata (Buchholz and Peters, 1876) and Naja melanoleuca Hallowell, 1857 and noted that the three constitute a clade of primarily forest inhabiting cobras. Wüster et al. MAP 332. Distribution of Naja annulata in Angola. (2007), therefore, considered both Boulengerina and Paranaja as synonyms of Naja . Wallach et al. (2009) recognized this clade as the subgenus Boulengerina Dollo, 1886 , whereas Wallach et al. (2014) treated this group as generically distinct. Spawls and Branch (1995) and Dobiey and Vogel (2007) both suggested that the species probably occurs in Cabinda and in the Zaire region of northwest Angola. A subadult Naja annulata from Lunda Norte Province (Branch and Conradie 2015), is the first voucheredd record for the country.

Naja melanoleuca Hallowell, 1857 FOREST COBRA

Naja haje ( var. melanoleuca ) Hallowell 1857:61. Syntypes: ANSP 6875–78 (collector H. A. Ford). Type locality: “Gaboon country, West Africa ” [= Gabon] .

Naja haje : Bocage (1866a:51, 1895a:132), Peters (1877a:618).

Naja haje haje : Loveridge (1957:291).

Naja melanoleuca : Boettger (1898:120), Boulenger (1905:114, 1915:219), Ferreira (1900b:133, 1903:12), Parker (1936:126), Bogert (1940:87), Laurent (1950a:10, 1954a:60), Hellmich (1957b:72), FitzSimons (1962:300), Broadley (1968c:5), Manaças (1981:27), Broadley (1990:291), Spawls and Branch (1995:71), Branch (1998:107), Broadley and Cotterill (2004:47), Chippaux (2006:225), Trapé and Mané (2006:198), Dobiey and Vogel (2007:69), Bates et al. (2014:404), Wallach et al. (2014:120), Ceríaco et al. (2017:141).

Naja melanoleuca melanoleuca : Laurent (1964a:120), Thys van den Audenaerde (1966:35), Chirio and LeBreton (2007:582).

Naja (Boulengerina) melanoleuca : Wallach et al. (2009:31).

Boulengerina melanoleuca : Wallach et al. (2014:120).

Global conservation status (IUCN): Not Evaluated.

Global distribution: The N. melanoleuca species complex is known from forest and savanna through west, central and east Africa, from Senegal east to western Ethiopia, southern Somalia, south to Angola on the west coast and KwaZulu-Natal, South Africa on the east coast. Populations in the east of the range are partly disjunct. With the recent elevation of N. subfulva to specific status (Ceríaco et al. 2017), the global distribution of N. melanoleuca will need to be reevaluated. Ocurrences in Angola (Map 333): The species occurs from the northeast of the country, including areas adjacent with Zambia in

Moxico Province (pers. obs.) to the northwest,

including Cabinda, and throughout western

Angola, exclusive of arid regions. Cabinda:

“Chinchoxo” [-5.10000, 12.10000] ( Bocage

1866a:51; Ferreira 1900b:133; Manaças

1981:27); “ Cabinda ” [-5.55000, 12.18333]

( Peters 1877a:618; Bocage 1895a:132, Boettger 1898:120). Lunda Norte: “Dundo”

[-7.36667, 20.83333] ( Laurent 1950a:10,

1954a:60, 1964a:120; Thys van den Audenaer-

de 1966:35; Manaças 1981:27); “Dundo, R.

Capemba” ( Thys van den Audenaerde

1966:35); “Dundo, Barrage de la Luachimo”

[-7.38333, 20.85000] (Thys van den Audenaer-

de 1966:35); “Dundo, Cacanda” [-7.40000, MAP 333. Distribution of Naja melanoleuca in Angola. 21.80000] ( Thys van den Audenaerde 1966:35); “Cossa” [-7.93333, 21.38333] ( Thys van den Audenaerde 1966:35); “R. Camaiala” [-8.05000, 18.61667] ( Thys van den Audenaerde 1966:35). Malanje: “Duque de Bragança” [-9.10000, 15.95000] ( Bocage 1866a:51, 1895a:132; Ferreira 1900b:133; Manaças 1981:27); “Pungo-Andongo” [-9.66667, 15.58333] ( Boulenger 1905:114; Manaças 1981:27). Kwanza Norte: “Piri-Dembos” [-8.56667, 14.50000] ( Hellmich 1957b:72; Manaças 1981:27; Ceríaco et al. 2017:141); “Ambaca” [-9.26667, 15.18333] (Manaças 1981:27);

“N’dalla Tando” [-9.30000, 14.91667] ( Ferreira 1903:12; Manaças 1981:27); “Cazengo” [-9.33333, 14.76667] (Manaças 1981:27). Kwanza Sul: “Libolo-Luati” [-9.98333, 14.90000] ( Hellmich 1957b:72; Manaças 1981:27; Ceríaco et al. 2017); “Quirimbo” [-10.68333, 14.26667] ( Parker 1936:126; Manaças 1981:27); “Congulu” [-10.86667, 14.28333] ( Parker 1936:126; Manaças 1981:27). Huambo: “Galanga” [-12.06667, 15.15000] ( Bocage 1895a:132). Benguela: “Cahata” [-12.35000, 14.81667] ( Bocage 1895a:132; Ferreira 1900b:133; Manaças 1981:27); “Hanha” [-12.25000, 13.75000] ( Bogert 1940:87; Manaças 1981:27). Huíla: “Sanguengue” [-12.36667, 16.20000] ( Hellmich 1957b:72); “Huilla” [-15.05000, 13.55000] ( Ferreira 1900b:133; Manaças 1981:27). Namibe: “Capangombe” [-15.10000, 13.15000] ( Ferreira 1900b:133; Manaças 1981:27).

Taxonomic and distributional notes: Broadley and Cotterill (2004) referred to Naja melanoleuca as a “species complex” that might contain several cryptic species. It was being reviewed by Broadley et al. at the time of the senior author’s death. Wallach et al. (2009) assigned Naja melanoleuca to the subgenus Boulengerina and later ( Wallach et al. 2014) treated Boulengerina as generically distinct. Although Spawls and Branch (1995) and Dobiey and Vogel (2007) considered N. melanoleuca to be restricted to northern Angola, including the Cabinda enclave, it is much more widely spread throughout the country. Ceríaco et al. (2017) presented molecular evidence to support Naja subfulva Laurent, 1955 , as a valid species and not a mere “savannah form” or subspecies of N. melanoleuca as considered by many previous authors. The two species are easily told apart by a combination of morphological characters and coloration, but to separate the Angolan records, many of them from before the original description of subfulva , will require a detailed review of all available specimens. Despite N. melanoleuca being considered more of a forest dweller and subfulva a savannah species, they are sympatric in parts of Angola (LMPC pers. obs.), frustrating attempts to infer the identity of records based solely on distribution.

Naja mossambica Peters, 1854 MOZAMBIQUE SPITTING COBRA

N [aja]. mossambica Peters 1854:625 . Lectotype: ZMB 2811 (collector W.C.H. Peters), designated by Golay et al. (1993:188). Type locality: “Tette and Sena” [= Tete, Vila de Sena], Mozambique, restricted to “Tette” as a result of lectotype designation.

Naja mossambica mossambica : Manaças (1981:28).

Naja mossambica : Broadley (1966c:25, 1974:156), Broadley (1990:295), Spawls and Branch (1995:74), Broadley and Cotterill (2004:47), Dobiey and Vogel (2007:71), Würster et al. (2007:438)

Naja mossambica mossambica : Manaças (1981:28).

Naja (Afronaja) mossambica : Wallach et al. (2009:32), Conradie et al. (2016:22) .

Afronaja mossambica : Wallach et al. (2014:10).

Global conservation status (IUCN): Not Evaluated.

Global distribution: The species is widespread in eastern and southern Africa, occuring from southern Tanzania southwestwards to extreme southeastern Angola, northern Namibia and Botswana, and southwards to eastern South Africa and Swaziland.

Ocurrences in Angola ( Map 334 ): The species occurs in the extreme southeastern regions of Cuando Cubango Province and there is an isolated record in Namibe Province. Namibe: “Maconjo” [-15.01667, 13.20000] ( Broadley 1974:156; Manaças 1981:28). Cuando Cubango: “Cuito basin (30a)” [-17.50875, 20.06594] ( Conradie et al. 2016:9, 10, 22); “Cuando basin (44b)” [-17.56916, 23.27305] ( Conradie et al. 2016:9, 10, 22).

Taxonomic and distributional notes: Dobiey and Vogel’s (2007) distribution map shows that the Mozambique Spitting Cobra in Angola is restricted to the extreme southeastern regions of Cuando Cubango Province, near the Zambian and Namibian borders. However, Broadley (1974,

1990) reported a record from “Maconjo” that represents an isolated western Angolan locality,

where it is sympatric with Naja (Afronaja)

nigricincta Bogert, 1940 . Wallach et al. (2009)

proposed nomenclatural changes regarding the genus Naja : the subgeneric name Naja is applicable to the Asiatic cobras, whereas African spitting cobras, including N. mossambica , were placed in a new subgenus, Afronaja Wallach,

Wüster and Broadley, 2009. Wallach et al.

(2014) used Afronaja at the generic rank.

Naja multifasciata Werner, 1902 MANY-BANDED COBRA

Naia multifasciata Werner 1902:347 . Holotype: IRSNB 2015 (collector L. Rom). Type locality: “der oberen Maringa” [= Upper Maringa River , Tshuapa], Democratic Republic of Congo.

Paranaja multifasciata : Spawls and Branch (1995:82), Nagy et al. (2005:224), Dobiey and Vogel (2007:80), Wüster et al. (2007:445).

Naja (Boulengerina) multifasciata : Wallach et al. (2009:31).

Boulengerina multifasciata : Wallach et al. (2014:121).

Global conservation status (IUCN): Not Evaluated.

Global distribution: The species is known from Cameroon to Congo.

Ocurrences in Angola: The species is expected to occur in the Cabinda enclave and potentially Zaire Province in northwestern Angola (Spawls and Branch 1995; Dobiey and Vogel 2007) but there are no vouchered records .

Taxonomic and distributional notes: Wüster et al. (2007) recovered the species Paranaja multifasciata ( Werner, 1902) as the sister taxon to Boulengerina annulata (Buchholz and Peters, 1876) plus Naja melanoleuca Hallowell, 1857 and they considered both Paranaja and Boulengerina as synonyms of Naja . Althoughh long placed in the monotypic genus Paranaja, Wallach et al. (2009) , based on phylogenetic information, assigned Naja multifasciata to the subgenus Boulengerina and later ( Wallach et al. 2014) treated Boulengerina as generically distinct.

Naja nigricincta Bogert, 1940 WESTERN BARRED SPITTING COBRA

Naja nigricollis var. fasciata Bocage 1895a:136 . Lectotype: MBL 1968 (collector A.P. De Carvalho), destroyed by fire 18 March 1978, designated by Broadley (1974:158). Type locality: “Benguella … Dondo … Capangombe,” Angola, restricted to “Benguella” through lectotype designation. Preoccupied by Naja fasciata Laurenti, 1768 , syn. Naja naja (Linneaus, 1758) .

Naja nigricollis nigricinctus Bogert 1940:89 , pl. 1, fig. 1. Holotype: AMNH 51823 (collector A.S. Vernay, H. Lang and R. Boulton). Type locality: “Muninho (101 km east of Mossamedes via railroad),” Namibe Province, Angola.

Naja nigricollis : Bocage (1866a:51, 1866b:71, 1867b:228), Ferreria (1900b:134), Monard (1937b:136-137).

Naja nigricollis nigricinctus : FitzSimons (1962:303).

Naja mossambica nigricincta : FitzSimons (1962:306), Manaças (1981:28).

Naja nigricincta : Broadley (1966c:25), Würster et al. (2007:445).

Naja nigricollis nigricincta : Broadley (1968c:7), Spawls and Branch (1995:78).

Naja nigricincta nigricincta : Broadley (1974:158, 1990:294), Dobiey and Vogel (2007:72).

Naja (Afronaja) nigricincta : Wallach et al. (2009:32).

Afronaja nigricincta : Wallach et al. (2014:10).

Global conservation status (IUCN): Not Evaluated.

Global distribution: The nominate form of the species is known from southwestern Angola and central and northern Namibia. Naja nigricincta woodi Pringle, 1955 is extralimital, occurring in southern Namibia and western South Africa.

Ocurrences in Angola (Map 335): The species is known from southwestern Angola.

Benguela: “ Benguela ” [-12.58333, 13.41667]

( Bocage 1866a:51, 1866b:71, 1867b:228,

1895a:136; Monard 1937b:136; Broadley

1974:158; Manaças 1981:28); “ Rio Equimina”

[-13.20000, 14.78333] ( Broadley 1974:158;

Manaças 1981:28). Namibe: “Munhino”

[-14.91667, 13.00000] ( Bogert 1940:89; Fitz-

Simons 1962:306; Broadley 1968c:7,

1974:158, 1990:294; Manaças 1981:28; Wallach et al. 2014:10); “Maconjo” [-15.01667,

13.20000] ( Broadley 1974:158; Manaças

1981:28); “Capangombe” [-15.10000,

13.15000] ( Bocage 1895a:136); “Foz do Cunene ” [-17.28333, 11.80000] ( Broadley

1974:158; Manaças 1981:28).

MAP 335. Distribution of Naja nigricincta in Angola.

Taxonomic and distributional notes:

Bocage (1895a) recognized several varieties of Naja nigricollis Reinhardt, 1843 based on coloration ( var. occidentalis ; var. melanoleuca ; var. fasciata ). Naja nigricollis nigricinctus Bogert, 1940 is a synonym of Bocage’s (1895a) var. fasciata , but since the older name was preoccupied by Naja fasciata Laurenti (1768) , a synonym of Naja naja ( Linnaeus, 1758) , Broadley (1966c) accept- ed Bogert’s (1940) as the valid name of this species, treating it as a subspecies of N. nigricollis . Years later he treated this form as a subspecies of N. mossambica Peters, 1854 ( Broadley 1968c, 1974). Bocage had recorded N. n. var. fasciata from “Dondo,” however Broadley (1974) examined the Museu Bocage specimens and only found one specimen from that locality, which he nominat- ed as the lectotype of Naja nigricollis var. occidentalis ( Bocage 1895a) (= Naja nigricollis Reinhardt, 1843 ) whereas the N. nigricollis var. fasciata speciemens were from “ Benguela ” and “Maconjo,” Spawls and Branch (1995) still recognized nigricincta as a subspecies of nigricollis , whereas Wüster et al. (2007) treated it as a separate evolutionary species, with N. n. woodi as a subspecies.

Naja nigricollis Reinhardt, 1843 BLACK-NECKED SPITTING COBRA

Naja nigricolli s Reinhardt 1843:269, pl. 3, figs. 5–7. Lectotype: ZMUC 65204 (collector J.R. Chenon) designated by F. W. Braestrup in Hughes and Barry (1969:1027). Type locality: “Guineakysten” ( Reinhardt 1843:271) [= coast of Ghana fide Hughes and Barry (1969:1027)], probably near “Christiansborg Castle, now a suburb of Accra,” [ Ghana], where the collector was stationed (Rasmussen and Hughes 1997:15).

Naja nigricollis var. occidentalis Bocage 1895a:135 . Lectotype: MBL 1963 (collector F.A.P. Bayão), destroyed by fire 18 March 1978, designated by Broadley (1974:156). Type locality: “Dondo … Quissange, Quillengues, Huilla et Humbe [ Angola] … Bissau [ Guinea-Bissau],” restricted to “Dondo” [= Sumbe] Kwanza Norte Province, Angola through the designation of the lectotype.

Naja nigricollis var. melanoleuca Bocage 1895a:136 (non Hallowell, 1857). Lectotype: MBL 1972 (collector J.A. d’Anchieta), destroyed by fire 18 March 1978, designated by Broadley (1974:156). Type locality: “Catumbella et … Caconda” restricted to “Caconda,” Angola through the designation of the lectotype.

Naja nigricollis : Bocage (1867b:228, 1896a:113), Peters (1881:149), Boettger (1898:120), Ferreria (1900b:134), Boulenger (1905:114), Schmidt (1933:14), Monard (1937b:136-137), Broadley (1966c:25), Manaças (1981:28), Spawls and Branch (1995:78), Dobiey and Vogel (2007:74), Würst et al. (2007:438).

Naia nigricollis : Boulenger (1896:378).

Naja nigricollis nigricollis : Mertens (1938a:442), Laurent (1950a:10, 1954a:60), Hellmich (1957a:74, 1957b:73), Loveridge (1957:292), FitzSimons (1962:302), Broadley (1974:156, 1990:292).

Naja nigricollis occidentalis : Laurent (1964a:119), Thys van den Audenaerde (1966:35), Broadley (1968c:7).

Naja (Afronaja) nigricollis : Wallach et al. (2009:32).

Afronaja nigricollis : Wallach et al. (2014:10).

Global conservation status (IUCN): Not Evaluated.

Global distribution: The species is widely distributed from southern Mauritania and Senegal east to Ethiopia and Somalia, and south to Zambia, Malawi, the southern Democratic Republic of Congo, and Angola. Absent from the forests of Central and West Africa, with disjunct populations along the lower Congo River, in southwestern Angola and elsewhere.

Ocurrences in Angola (Map 336): The species very widespread, from coastal areas to the interior highlands of Angola. Zaire:

“Ambrizette” [-7.23333, 12.86667] ( Bocage

1895a:135; Boettger 1898:120; Manaças

1981:28). Malanje: “ Malanje ” [-9.55000,

16.35000] ( Peters 1881:149; Bocage

1895a:135; Manaças 1981:28). Kwanza

Norte: “Golungo Alto” [-9.13333, 14.76667]

( Boulenger 1905:114; Monard 1937b:136);

“Dondo” [-9.68333, 14.43333] ( Bocage

1895a:135; Ferreira 1900b:134; Monard

1937b:136; Loveridge 1957:292; Hellmich

1957b:73; Broadley 1968c:7; Manaças

1981:28). Kwanza Sul: “Libolo-Luati”

[-9.98333, 14.90000] ( Hellmich 1957b:73;

Manaças 1981:28); “Dondo a Humbe” ( Ferreira 1900b:134). Bié: “Chitau” [-11.43333, MAP 336. Distribution of Naja nigricollis in Angola. 17.15000] ( Schmidt 1933:14). Benguela: “Catumbella” [-12.43333, 13.55000] ( Bocage 1867b:228, 1895a:135; Ferreira 1900b:134; Manaças 1981:28); “Quissange” [-12.43333, 14.05000] ( Bocage 1895a:135; Ferreira 1900b:134; Monard 1937b:136; Loveridge 1957:292; Broadley 1968c:7; Manaças 1981:28); “Chivitidi/Entre Rios” [-13.01667, 14.63333] ( Hellmich 1957a:74; Manaças 1981:28); “Cubal” [-13.03333, 14.25000] Mertens 1938:442; Manaças 1981:28); “Hanha” [-13.30000, 14.20000] ( Bocage 1896a:113; Manaças 1981:28). Huíla: “Caconda” [-13.73333, 15.06667] ( Bocage 1895a:135; Ferreira 1900b:134; Monard 1937b:136; Manaças 1981:28); “Quillengues” [-14.06667, 15.08333] ( Bocage 1895a:135; Monard 1937b:136; Loveridge 1957:292; Broadley 1968c:7; Manaças 1981:28); “Kuvangu/Kubangu/Vila da Ponte”

[-14.46667, 16.30000] ( Monard 1937b:136-137; Manaças 1981:28); “Capelongo” [-14.88333, 15.08333] (Manaças 1981:28); “Huilla” [-15.05000, 13.55000] ( Bocage 1895a:135; Monard 1937b:136; Loveridge 1957:292; Broadley 1968c:7; Manaças 1981:28); “Osi” [-15.08333, 15.40000] ( Monard 1937b:136-137; Manaças 1981:28); “Gambos” [-15.76667, 14.10000] (Manaças 1981:28). Namibe: “Capangombe” [-15.10000, 13.15000] ( Ferreira 1900b:134; Monard 1937b:136; Manaças 1981:28). Cunene: “Humbe” [-16.68333, 14.90000] ( Bocage 1895a:135; Ferreira 1900b:134; Monard 1937b:136; Loveridge 1957:292; Broadley 1968c:7; Manaças 1981:28); “Mupanda” [-17.13333, 15.76667] ( Monard 1937b:136-137; Manaças 1981:28). Cuando Cubango: “Kayundu” [-15.70000, 17.45000] ( Monard 1937b:136-137; Manaças 1981:28).

Taxonomic and distributional notes: Bocage’s (1895a) locality of “Ambrizette” in Zaire Province has been considered questionable and has been excluded from some distribution maps for the species (Spawls and Branch 1995; Dobiey and Vogel 2007). However, Dobiey and Vogel (2007) do include Cabinda within the range of the species, although we are not aware of any published records. These authors also indicate occurrence in northcentral Namibia, although only a single specimen, from Katima Mulilo in the eastern Caprivi Strip has been documented from that country ( Broadley 1974). The taxonomy of N. nigricollis still requires further study ( Wüster et al. 2007). Wallach et al. (2009) placed the African spitting cobras in a new subgenus, Afronaja Wallach, Wüster and Broadley, 2009 , with Naja nigricollis as the type species and Wallach et al. (2014) have treated this as a full genus.

Naja subfulva Laurent, 1955 SAVANNA COBRA

Naja melanoleuca subfulva Laurent 1955:61 . Holotype: MRAC 17514 (collector Van der Borght). Type locality: “Lwiro, 1850 m, Terr. de Kabare , Kivu,” Democratic Republic of the Congo.

Naja subfulva : Ceríaco et al. (2017:141).

Global conservation status (IUCN): Not Evaluated.

Global distribution: The N. melanoleuca species complex is known from forest and savanna through west, central, and east Africa, from Senegal east to western Ethiopia, southern Somalia, south to Angola on the west coast and KwaZulu-Natal, South Africa on the east coast. Populations in the east of the range are partly disjunct. With the recent elevation of N. subfulva to specific status (Ceríaco et al. 2017), the global distribution of both this taxon and N. melanoleuca sensu stricto will need to be reevaluated.

Ocurrences in Angola (Map 337): The species should occur across the country, except in the more heavily forested areas in the north, but currently the only confirmed records are for Kwanza Norte, Kwanza Sul and Malanje provinces. Malanje: “Cacuso” [-9.421023, 15.746679] (Ceríaco et al. 2017:141) . Kwanza: Norte: “Piri-Dembos” [-8.56667, 14.50000] (Ceríaco et al. 2017:141). Kwanza Sul: “Libolo-Luati” [-9.98333, 14.90000] (Ceríaco et al. 2017:141) .

Taxonomic and distributional notes: See Naja melanoleuca account. MAP 337. Distribution of Naja subfulva in Angola.

Figures

MAP 334. Distribution of Naja mossambica in Angola.