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

publication ID

https://doi.org/ 10.5281/zenodo.13159758

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https://treatment.plazi.org/id/3A1F87CA-4E08-4B04-A58E-C4C9FB89F94C

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Felipe

scientific name

Naja Laurenti, 1768
status

 

Genus Naja Laurenti, 1768 View in CoL

Naja anchietae Bocage, 1879 ANCHIETA’S COBRA

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

Naja anchietae View in CoL : 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 View in CoL : 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 View in CoL : 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] GoogleMaps

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

Huíla: “Caconda” [-13.73333, 15.06667] GoogleMaps

( 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 View in CoL COBRA

Naja annulata Buchholz and Peters 1876 View in CoL 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 View in CoL : 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) GoogleMaps .

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 View in CoL COBRA

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

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

Naja haje haje View in CoL : Loveridge (1957:291).

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

Naja (Boulengerina) melanoleuca View in CoL : 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] GoogleMaps

( 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 View in CoL MOZAMBIQUE SPITTING COBRA

N [aja]. mossambica Peters 1854:625 View in CoL . 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 View in CoL : 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 View in CoL : Manaças (1981:28).

Naja (Afronaja) mossambica View in CoL : Wallach et al. (2009:32), Conradie et al. (2016:22) View Cited Treatment .

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 View MAP ): 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 View in CoL . Wallach et al. (2009)

proposed nomenclatural changes regarding the genus Naja View in CoL : the subgeneric name Naja View in CoL 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 View in CoL COBRA

Naia multifasciata Werner 1902:347 View in CoL . Holotype: IRSNB 2015 View Materials (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 View in CoL : 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 View in CoL WESTERN BARRED SPITTING COBRA

Naja nigricollis var. fasciata Bocage 1895a:136 View in CoL . 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 View in CoL , syn. Naja naja (Linneaus, 1758) View in CoL .

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

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

Naja nigricollis nigricinctus : FitzSimons (1962:303).

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

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

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

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

Naja (Afronaja) nigricincta View in CoL : 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] GoogleMaps

( 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 View in CoL 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 View in CoL . 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 View in CoL (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 View in CoL : 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 View in CoL : 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 View in CoL : Laurent (1964a:119), Thys van den Audenaerde (1966:35), Broadley (1968c:7).

Naja (Afronaja) nigricollis View in CoL : 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 View in CoL COBRA

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

Naja subfulva View in CoL : 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) GoogleMaps . 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) GoogleMaps .

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

Kingdom

Animalia

Phylum

Chordata

Class

Squamata

Family

Elapidae

Kingdom

Animalia

Phylum

Chordata

Class

Squamata

Family

Elapidae

Genus

Naja

Loc

Naja Laurenti, 1768

Marques, Mariana P., Ceríaco, Luis M. P., Blackburn, David C. & Bauer, Aaron M. 2018
2018
Loc

Uraeus anchietae

WALLACH V., K. L. & WILLIAMS, AND & J. BOUNDY 2014: 762
2014
Loc

Boulengerina melanoleuca

WALLACH V., K. L. & WILLIAMS, AND & J. BOUNDY 2014: 120
2014
Loc

Afronaja mossambica

WALLACH V., K. L. & WILLIAMS, AND & J. BOUNDY 2014: 10
2014
Loc

Boulengerina multifasciata

WALLACH V., K. L. & WILLIAMS, AND & J. BOUNDY 2014: 121
2014
Loc

Afronaja nigricincta

WALLACH V., K. L. & WILLIAMS, AND & J. BOUNDY 2014: 10
2014
Loc

Afronaja nigricollis

WALLACH V., K. L. & WILLIAMS, AND & J. BOUNDY 2014: 10
2014
Loc

Naja (Uraeus) anchietae

WALLACH, V., W. & WUSTER, AND & D. G. BROADLEY 2009: 31
2009
Loc

Naja (Boulengerina) annulata

CONRADIE, W. & W. R. BRANCH & G. WATSON 2015: 200
WALLACH, V., W. & WUSTER, AND & D. G. BROADLEY 2009: 31
2009
Loc

Naja (Boulengerina) melanoleuca

WALLACH, V., W. & WUSTER, AND & D. G. BROADLEY 2009: 31
2009
Loc

Naja (Afronaja) mossambica

CONRADIE, W. & R. BILLS & W. R. BRANCH 2016: 22
WALLACH, V., W. & WUSTER, AND & D. G. BROADLEY 2009: 32
2009
Loc

Naja (Boulengerina) multifasciata

WALLACH, V., W. & WUSTER, AND & D. G. BROADLEY 2009: 31
2009
Loc

Naja (Afronaja) nigricincta

WALLACH, V., W. & WUSTER, AND & D. G. BROADLEY 2009: 32
2009
Loc

Naja (Afronaja) nigricollis

WALLACH, V., W. & WUSTER, AND & D. G. BROADLEY 2009: 32
2009
Loc

Paranaja multifasciata

WUSTER, W. & S. CROOKES & I. INEICH & Y. MANE & C. E. POOK & J. - F. TRAPE & D. G. BROADLEY 2007: 445
NAGY, Z. T. & N. VIDAL & M. VENCES & W. R. BRANCH & O. S. G. PAUWELS & M. WINK & U. JOGER 2005: 224
2005
Loc

Boulengerina annulata

WALLACH V., K. L. & WILLIAMS, AND & J. BOUNDY 2014: 119
WUSTER, W. & S. CROOKES & I. INEICH & Y. MANE & C. E. POOK & J. - F. TRAPE & D. G. BROADLEY 2007: 445
NAGY, Z. T. & N. VIDAL & M. VENCES & W. R. BRANCH & O. S. G. PAUWELS & M. WINK & U. JOGER 2005: 224
BROADLEY, D. G. & C. T. DORIA & J. WIGGE 2003: 106
2003
Loc

Naja anulifera ancheitae

BROADLEY, D. G. 1995: 31
1995
Loc

Naja nigricincta nigricincta

BROADLEY, D. G. 1990: 294
BROADLEY, D. G. 1974: 158
1974
Loc

Naja nigricollis nigricincta

BROADLEY, D. G. 1968: 7
1968
Loc

Naja mossambica

BROADLEY, D. G. 1990: 295
BROADLEY, D. G. 1974: 156
BROADLEY, D. G. 1966: 25
1966
Loc

Naja nigricincta

BROADLEY, D. G. 1966: 25
1966
Loc

Naja melanoleuca melanoleuca

THYS VAN DEN AUDENAERDE, D. F. E. 1966: 35
LAURENT, R. F. 1964: 120
1964
Loc

Naja nigricollis occidentalis

BROADLEY, D. G. 1968: 7
THYS VAN DEN AUDENAERDE, D. F. E. 1966: 35
LAURENT, R. F. 1964: 119
1964
Loc

Naja nigricollis nigricinctus

FITZSIMONS, V. F. 1962: 303
1962
Loc

Naja mossambica nigricincta

FITZSIMONS, V. F. 1962: 306
1962
Loc

Naja melanoleuca subfulva

LAURENT, R. F. 1955: 61
1955
Loc

Naja haje anchietae

TRAPE, J-F. & L. CHIRIO & D. G. BROADLEY & W. WUSTER 2009: 20
BROADLEY, D. G. 1966: 23
LAURENT, R. F. 1964: 118
BOGERT, C. M. 1940: 90
1940
Loc

Naja nigricollis nigricinctus

BOGERT, C. M. 1940: 89
1940
Loc

Naja nigricollis nigricollis

BROADLEY, D. G. 1990: 292
BROADLEY, D. G. 1974: 156
FITZSIMONS, V. F. 1962: 302
HELLMICH, W. 1957: 74
HELLMICH, W. 1957: 73
LAURENT, R. F. 1954: 60
LAURENT, R. F. 1950: 10
MERTENS, R. 1938: 442
1938
Loc

Naja anchietae

BROADLEY, D. G. 1968: 7
FRADE, F. 1963: 252
BOGERT, C. M. 1940: 90
MONARD, A. 1937: 138
1937
Loc

Naia multifasciata

WERNER, F. 1902: 347
1902
Loc

Naja melanoleuca

BATES, M. F. & W. R. BRANCH & A. M. BAUER & M. BURGER & J. MARAIS & G. J. ALEXANDER & M. S. DE VILLIERS 2014: 404
WALLACH V., K. L. & WILLIAMS, AND & J. BOUNDY 2014: 120
CHIPPAUX, J. - P. 2006: 225
BRANCH, W. R. 1998: 107
BROADLEY, D. G. 1990: 291
BROADLEY, D. G. 1968: 5
FITZSIMONS, V. F. 1962: 300
HELLMICH, W. 1957: 72
LAURENT, R. F. 1954: 60
LAURENT, R. F. 1950: 10
BOGERT, C. M. 1940: 87
PARKER, H. W. 1936: 126
BOULENGER, G. A. 1915: 219
BOULENGER, G. A. 1905: 114
FERREIRA, J. B. 1903: 12
FERREIRA, J. B. 1900: 133
BOETTGER, O. 1898: 120
1898
Loc

Naia nigricollis

BOULENGER, G. A. 1896: 378
1896
Loc

Naja nigricollis var. fasciata

BROADLEY, D. G. 1974: 158
BOCAGE, J. V. B. 1895: 136
1895
Loc

Naja nigricollis var. occidentalis

BROADLEY, D. G. 1974: 156
BOCAGE, J. V. B. 1895: 135
1895
Loc

Naja nigricollis var. melanoleuca

BROADLEY, D. G. 1974: 156
BOCAGE, J. V. B. 1895: 136
1895
Loc

Naja Anchietae

BROADLEY, D. G. 1995: 31
BOCAGE, J. V. B. 1879: 98
BOCAGE, J. V. B. 1879: 89
1879
Loc

Naja Anchietae

BOCAGE, J. V. B. 1897: 202
BOCAGE, J. V. B. 1895: 133
BOCAGE, J. V. B. 1879: 98
1879
Loc

Naja annulata Buchholz and Peters 1876

PETERS, W. C. H. 1876: 119
1876
Loc

Naja nigricollis

BROADLEY, D. G. 1966: 25
MONARD, A. 1937: 136
SCHMIDT, K. P. 1933: 14
BOULENGER, G. A. 1905: 114
BOETTGER, O. 1898: 120
BOCAGE, J. V. B. 1896: 113
PETERS, W. C. H. 1881: 149
BOCAGE, J. V. B. 1867: 228
1867
Loc

Naja haje

BOCAGE, J. V. B. 1895: 132
PETERS, W. C. H. 1877: 618
BOCAGE, J. V. B. 1866: 51
1866
Loc

Naja nigricollis

MONARD, A. 1937: 136
BOCAGE, J. V. B. 1867: 228
BOCAGE, J. V. B. 1866: 51
BOCAGE, J. V. B. 1866: 71
1866
Loc

Naja haje

HALLOWELL, E. 1857: 61
1857
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