Bitis Gray, 1842

Marques, Mariana P., Ceríaco, Luis M. P., Blackburn, David C. & Bauer, Aaron M., 2018, Diversity and Distribution of the Amphibians and Terrestrial Reptiles of Angola Atlas of Historical and Bibliographic Records (1840 - 2017), Proceedings of the California Academy of Sciences 65, pp. 1-501 : 309-315

Genus Bitis Gray, 1842

Bitis arietans ( Merrem, 1820) PUFF ADDER

Vipera (Echidna) arietans Merrem 1820:152 . Lectotype: specimen described and illustrated by Seba

(1735:55, pl. 54, fig. 4), now lost (collector unknown), designated by Wallach et al. (2014:611). Type locality: “Vorgebürge der guten Hoffnung/Promontorio bonae spei” ( Merrem 1820:152), [= Cape of Good

Hope] Western Cape Province, Southern South Africa. Echidna arietans : Bocage (1866a:53). Clotho arietans : Günther (1865a:480). Echida arietans : Bocage (1879c:89). Bitis arietans : Bocage (1887a:190, 1887c:211), Ferreira (1897b:245), Boulenger (1915:221), Schmidt

(1933:15), Loveridge (1936a:45), Monard (1937b:141-142), Themido (1941:11), Hellmich (1957a:75),

Spawls and Branch (1995:114), Branch (1998:114), Lenk et al. (1999:31), Dobiey and Vogel (2007:104),

Barlow et al. (2013:1134), Wallach et al. (2014:89), Branch and Conradie (2015:200), Ceríaco et al.

(2016a:87), Conradie et al. (2016:23). Vipera arietans : Bocage (1895a:149, 1896a: 113). Cobra lachesis : Mertens (1937a:16). Bitis lachesis : Mertens (1938a:442), Bogert (1940:99), Laurent (1950a:11, 1954a:62). Bitis arietans arietans : Hellmich (1957b:74), Laurent (1964a:127), Thys van den Audenaerde (1966:36),

Manaças (1981-82:35), Branch and McCartney (1992:2), Bates et al. (2014:331).

Global conservation status (IUCN): Not Evaluated.

Global distribution: The species is known from most of sub-Saharan Africa, extending to the Arabian Peninsula.

Ocurrences in Angola (Map 269): The species occurs in the entire country with exception of the desert regions of the far southwest, as well as some mountainous areas.

Zaire: “ S. Salvador do Congo ” [-6.26667,

14.23333] ( Bocage 1887a:190, 1895a:149;

Manaças 1981:35). Bengo: “Ambriz”

[-7.844312, 13.106493] (Manaças 1981:35).

Kwanza Norte: “Mucoso, Dondo” [-9.53333,

14.65000] ( Hellmich 1957a:75; Manaças

1981:35). Malanje: “Duque de Bragança ”

[-9.10000, 15.95000] ( Bocage 1866a:53,

1895a:149; Monad 1937b:141; Manaças

1981:35); “Cangandala National Park”

[-9.84606, 16.72233] (Ceríaco et al.

2016a:87). Lunda Norte: “Dundo” [-7.36667,

20.83333] ( Laurent 1950a:11, 1954a:62,

MAP 269. Distribution of Bitis arietans in Angola. 1964a:127; Manaças 1981:35); “Capaia”

[-7.33556, 20.21681] (Branch and Conradie 2015:200); “Muíta, Luembe E” [-7.80000, 21.45000] ( Laurent 1950a:11, 1954a:62; Manaças 1981:35); “Sombo” [-8.68333, 20.95000] (Laurent 1954:62); “Maludi” [-8.03333, 21.33333] ( Thys van den Audenaerde 1966:36). Lunda Sul: “Alto Chicapa” [10.93333, 19.15000] ( Laurent 1964a:127; Manaças 1981:35). Moxico: “environs du lac Calundo (Lago Calundo)” [-11.80000, 20.86667] ( Laurent 1964a:127; Manaças 1981:35). Bié: “Chitau” [-11.25000, 17.16667] ( Schmidt 1933:15; Bogert 1940:99; Manaças 1981:35); “Cubango basin (9)” [-14.00269, 17.40500] ( Conradie et al. 2016:8-9, 23). Huambo: “Bela Vista” [-12.56667, 16.21667] (Manaças 1981:35; Hellmich 1957b:74); “ Huambo ” [-12.83333, 15.66667]

( Bogert 1940:99; Themido 1941:11; Manaças 1981:35); “Cuma” [-12.86667, 15.06667] ( Loveridge 1936a:45; Manaças 1981:35). Benguela: “Quissange” [-12.43333, 14.05000] ( Bocage 1887c:211); “Benguella” [-12.58333, 13.41667] ( Bocage 1866a:53); “Entre Rios” [-13.01667, 14.63333] ( Hellmich 1957a:75; Manaças 1981:35); “Cubal” [-13.03333, 14.25000] (Mertens 1937:16, 1938:442; Manaças 1981:35); “Equimina” [-13.20000, 14.78333] ( Bocage 1895a:149; Manaças 1981:35); “Hanha” [-13.30000, 14.20000] ( Bocage 1896a:113; Manaças 1981:35). Huíla: “Capelongo” [-14.91667, 15.08333] ( Bogert 1940:99; Manaças 1981:35); “Sangevé” [-13.88333, 15.83333] ( Monard 1937b:141-142; Manaças 1981:35); “ Rio Calae” [-13.96667, 16.03333] ( Bocage 1879c:89, 1895a:149; Monad 1937b:141; Manaças 1981:35); “Kuvangu (Vila-da-Ponte)” [-14.46667, 16.30000] ( Monard 1937b:141, 142; Manaças 1981:35); “Kampulu” [-15.21667, 16.11667] ( Monard 1937b:141-142; Manaças 1981:35); “ Rio Cabindongo” ( Bocage 1895a:149; Monad 1937b:141; Manaças 1981:35). Namibe: “Mossamedes” [-15.20000, 12.15000] ( Günther 1865a:480; Manaças 1981:35). Cunene: “ruisseau Mbalé (Bale)” [-15.16667, 16.75000] ( Monard 1937b:141-142; Manaças 1981:35). Cuando Cubango: “approximately 33 km ESE of Cuito Cuanavale” [-15.28333, 19.50000] (Branch and McCartney 1992:2); “Cuito basin (61)” [-16.98919, 19.40614] “Cuando basin (38)” (not collected) [-17.58830, 22.65694] ( Conradie et al. 2016:8, 10, 23). Undetermined Locality: “from almost all localities that he [J. Anchieta] visited” ( Bocage 1895a:149).

Taxonomic and distributional notes: Merrem (1820) had indications to numerous earlier works, including Seba (1735) and Gronovius (1756). In addition, his scale counts (ventrals + caudals), though similar to those given by Gronovius are not exactly the same, suggesting that he made scale counts on at least one specimen himself. Merrem had access to some of Seba’s original specimens through the collections of Willem Xaver Janssen (1760–1793) and Friedrich Heinrich Graf von Borcke (1776–1825), successive owners of Seba specimens originally sold at auction ( Anonymous 1752) (Bauer and Günther 2013). Surviving B. arietans from this collection ( ZMB 2879) does not correspond to the Seba image indicated by Merrem, although it may have been examined by him and could have been part of the basis for his description. Wallach et al. (2014) appear to be the first to have explicitly designated a lectotype for the puffadder.

The species has significant phylogeographic structure ( Lenk et al. 1999; Barlow et al. 2013; Bates et al. 2014) and the implications for its systematics status are being investigated. Currently, two subspecies are recognized: Bitis arietans arietans ( Merrem, 1820) widespread in sub-Saharan Africa and Bitis arietans somalica Parker, 1949 restricted to Somalia ( Branch 1998; Bates et al. 2014). Some authors (e.g., Spawls and Branch 1995; Vogel and Dobiey 2007) considered B. arietans widespread in Angola although with no occurences in northwest regions of the country, whereas Wallach et al. (2014) only cited localities in the eastern regions of Lunda Norte and Moxico Provinces. See McDiarmid et al. (1999) for a chresonomy of the species.

Bitis caudalis (Smith, 1839) HORNED ADDER

Vipera ocellata Smith 1838:92 . Syntypes: BMNH 1865.5.4.153a-e (collector A. Smith). Type locality: “sandy districts north of the Cape Colony,” South Africa. Junior primary homonym of Vipera ocellata Latreille ,

1801 [= Vipera aspis Linnaeus, 1758 ]. Vipera (Cerastes) caudalis Smith 1839 : pl. 7, first of two accompanying unnumbered text pages. Syntypes:

BMNH 1865.5.4.153a-e (collector A. Smith) . Type locality: “sandy districts north of the Cape Colony ,”

South Africa (verbatim from Smith 1838). Nomen substitutum for Vipera ocellata Smith, 1838 . Cerastes caudalis : Bocage (1867c:227, 1870:68). Vipera caudalis : Günther (1865a:480), Bocage (1895a:150). Bitis caudalis : Boulenger (1915:221), Frade (1963:253), Haacke (1984:174), Manaças (1981:37), Visser (1981:7), Spawls and Branch (1995:105), Branch (1998:116), Dobiey and Vogel (2007:110), Bates et al. (2014:334), Wallach et al. (2014:90).

Bitis caudalis caudalis : Laurent (1964a:128).

Global conservation status (IUCN): Not Evaluated.

Global distribution: The species is known from the arid western region of southern Africa, throughout Karoo and Kalahari, from southern Zimbabwe and the western half of South Africa to southern Angola – the northern limit for the species.

Ocurrences in Angola (Map 270): The species occurs in the southwestern Angola,

with preference of hot and dry open areas in sandy regions. Luanda: “Loanda” [-8.83333,

13.26667] ( Bocage 1895a:150; Manaças

1981:37). Benguela: “ Benguela ” [-12.58333,

13.41667] ( Manaças 1981:37). Huíla : “ Huíla ”

[-15.08333, 13.55000] (Manaças 1981:37).

Namibe: “Capangombe” [-15.10000,

13.15000] ( Bocage 1895a:150; Monard

1937b:141; Manaças 1981:37); “Mossamedes”

[-15.20000, 12.15000] ( Günther 1865a:480;

Bocage (1867c:227; Manaças 1981:37);

“ Désert de Moçâmedes , à 35 km au sud de la ville” [-15.20000, 12.15000] ( Laurent

1964a:128); “ Rio Coroca ” [-15.78333,

12.06667] ( Bocage 1895a:150; Monard

1937b:141; Manaças 1981:37). Undetermined

MAP 270. Distribution of Bitis caudalis in Angola.

Locality: “with no precise location” ( Bocage 1870:68).

Taxonomic and distributional notes: Although Smith’s (1838) Vipera ocellata was preoccupied, Smith (1839) noted only “As it does not appear probable that the genis Vipera will ultimately be divisible into subgenera, I have thought it advisable to substitute for ocellata . — a term which under such circumstances would cause some confusion, — that of caudalis .” Preliminary phylogeographic analysis indicates appreciable regional divergence (A. Barlow et al. unpubished data in Bates et al. 2014). Visser (1981) provided a map with the Transvaal Museum accessions since 1962 for Bitis caudalis . Haacke (1984) subsequently provided a map with some records for the species in Namibe Province, without explicit locality information, however the two maps are very similar and probably represent the same data, showing the southwestern distribution of the species in the country. The Angolan record from “Loanda/ Luanda ” ( Bocage 1895a; Manaças 1981) is doubtful and certainly represents a misidentification.

Bitis gabonica Duméril, Duméril and Bibron, 1854 GABON ADDER

Echidna Gabonica Duméril, Duméril and Bibron 1854:1428 , pl. 80 bis, figs. 1–3. Lectotype: MNHH 4012

(collector C.-E. Aubry-Lecomte), designated by Hughes and Barry (1969:1030). Type locality: “côte du

Gabon ” ( Duméril, Duméril and Bibron 1854:1428), Gabon. Echidna rhinoceros : Bocage (1866a:53). Vipera (Bitis) rhinoceros : Peters (1877a:618). Bitis rhinoceros : Bocage (1887a:191). Vipera rhinoceros : Bocage (1895a:149, 1896a:113, 1897b:211). Bitis gabonica : Boulenger (1915:222), Laurent (1950a:11), Spawls and Branch (1995:116-117), Lenk et al.

(1999:31), Lenk et al. (2001:94), Chippaux (2006:260), Chirio and LeBreton (2007:602), Dobiey and Vogel (2007:111), Wallach et al. (2014:90), Oliveira et al. (2016:42).

Bitis gabonica gabonica : Laurent (1954a:62, 1964a:128), Loveridge (1957:302), Thys van den Audenaerde (1966:36), Manaças (1981:36).

Global conservation status (IUCN): Not Evaluated.

Global distribution: The species is known from forest-savanna mosaic of West and Central Africa from Benin to Angola and is very widespread in the Democratic Republic of Congo, extending marginally into adjacent areas such as southern Nigeria and northern Zambia, with isolated populations in South Sudan, Kenya, and eastern southern Africa, the southern most occurring in northeastern KwaZulu-Natal.

Ocurrences in Angola (Map 271): The species occurs in northern regions of Angola including Cabinda enclave and the northeastern areas of Moxico Province. Cabinda:

“Chinchoxo” [-5.48333, 12.13333] ( Peters

1877a:618; Bocage 1895a:149; Manaças

1981:36); “ Cabinda ” [-5.55000, 12.18333]

( Bocage 1866a:53, 1887a:191, 1895a:149;

Manaças 1981:36). Zaire: “S. Salvador do

Congo ” [-6.26667, 14.23333] ( Bocage

1887a:191, 1895a:149; Manaças 1981:36).

Malanje: “Cacuso” [-9.4994, 15.1613] ( Oliveira et al. 2016:42); “Kalandula” [-91956,

15.9026] [-9.2097, 15.8906] [-91907, 15.9159]

[-9.3872, 15.9441] ( Oliveira et al. 2016:42).

Kwanza Norte: “N’dala Tando” [-9.30000,

14.91667] (Manaças 1981:36). Benguela :

MAP 271. Distribution of Bitis gabonica in Angola. “Hanha” [-13.30000, 14.20000] ( Bocage

1896a:113, 1897b:211; Manaças 1981:36); “Caimbambo” [-13.1677, 13.7663] ( Oliveira et al. 2016:42). Moxico: “environs du lac Calundo (Lago Calundo)” [-11.71667, 20.80000] ( Laurent 1964a:128; Manaças 1981:36). Lunda Norte: “Dundo” [-7.3667, 20.8267] ( Oliveira et al. 2016:42).

Taxonomic and distributional notes: McDiarmid et al. (1999) incorrectly cited MNHN 4020 as a holotype specimen. The description was based on a skin sent by Aubry-Lecomte and a specimen from the Ménagerie du Muséum. Angolan specimens of Bitis gabonica have occasionally been cited as Bitis rhinoceros (Schlegel, 1855) — a closely related species endemic to West Africa ( Lenk et al. 1999; Lenk et al. 2001; Chipaux 2006; Bates et al. 2014; Wallach et al. 2014).

Bitis heraldica ( Bocage, 1889) ANGOLAN ADDER (Endemic)

Vipera heraldica Bocage 1889:127 . Holotype: MBL 2127 (collectors H.C. Capello and R. Ivens), destroyed by fire 18 March 1978. Type locality: “sur les bords de la rivière Calae, l’un des affluents du Cunene ” [= Calai River banks] Huíla Province, Angola.

Vipera heraldica : Bocage (1895a:151).

Bitis peringueyi : Boulenger (1896:495, 1905:114, 1915:221), Monard (1937b:143), Bogert (1940:101).

Bitis heraldica : Ferreira (1897b:245), Hellmich (1957b:75), Mertens (1958:146), Manaças (1981:38), Spawls and Branch (1995:104), Dobiey and Vogel (2007:113), Wallach et al. (2014:91).

Global conservation status (IUCN): Not Evaluated.

Global distribution: The species is known from western Angola, chiefly in association with the high plateau.

Ocurrences in Angola (Map 272): The species is known from the central highlands in

Angola. Zaire: “S. Salvador do Congo”

[-6.26667, 14.23333] (Manaças 1981:38).

Bengo: “Bengu” [-8.71667, 13.40000]

( Monard 1937b:141; Mertens 1958:147;

Manaças 1981:38; Spawls and Branch

1995:105; Wallach et al. 2014:91). Kwanza

Sul: “Mombolo” [-12.16667, 14.83333]

( Bogert 1940:101; Mertens 1958:146; Manaças 1981:37-38). Huambo: “Bela Vista (Sanguengue)” [-12.56667, 16.21667] ( Hellmich

1957b:75; Mertens 1958:146; Manaças

1981:38; Spawls and Branch 1995:105). Bié:

“ Bié ” [-12.38333, 16.95000] ( Mertens

1958:147; Manaças 1981:38; Spawls and

Branch 1995:105) Huíla: “Caconda”

MAP 272. Distribution of Bitis heraldica in Angola. [-13.73333, 15.06667] ( Bocage 1895a:151;

Ferreira 1897b:245; Monard 1937b:141; Mertens 1958:147; Manaças 1981:38; Spawls and Branch 1995:105); “Kalukembé (Caluquembe)” [-13.78333, 14.68333] ( Monard 1937b:141, 143; Mertens 1958:147; Manaças 1981:38; Spawls and Branch 1995:105); “sur les bords de la rivière Calae, l’un des affluents du Cunene (Rio Calae)” [-13.96667, 16.03333] ( Bocage 1889:127, 1895a:151; Monard 1937b:141; Mertens 1958:147; Manaças 1981:38; Spawls and Branch 1995:105). Undetermined Locality: “Between Benguella and Bihé” ( Boulenger 1905:114; Monard 1937b:141).

Taxonomic and distributional notes: This species was described by Bocage (1889) based on one specimen from “sur les bords de la rivière Calae, l’un des affluents du Cunene ”. This species was later considered by Boulenger (1896) a synonym of Bitis peringueyi ( Boulenger, 1888) although Ferreira (1897) doubted this conclusion. Later, Mertens (1958) restablished B. heraldica based on distinctive morphological characters between the two species and refered Angolan material from “Kalukembé” ( Monard, 1937b) and from “Namba, Mombolo” ( Bogert, 1940) to B. heraldica .

Bitis nasicornis ( Shaw, 1792) RHINOCEROS VIPER

Coluber Nasicornis : Shaw in Shaw and Nodder 1792: pl. 94, first of three accompanying unnumbered pages of Latin description). Holotype: BMNH, lost fide Golay et al. (1993) (donated by E. Jenkins). Type locality: “interiori Africa” [“likely to be from Ghana ” fide Hughes and Barry 1969:1030].

Bitis nasicornis : Boulenger (1915:222), Parker (1936:126), Hellmich (1957b:76), Loveridge (1957:302), Manaças (1981:36), Spawls and Branch (1995:118), Lenk et al. (1999:31), Lenk et al. (2001:96), Chirio and LeBreton (2007:604), Dobiey and Vogel (2007:117), Wallach et al. (2014:91).

Global conservation status (IUCN): Not Evaluated.

Global distribution: The species is restricted to the rainforest of West Africa, known from Guinea to Gabon, extending east to Uganda, western Kenya, and northern Tanzania, having its southern limit in the northern regions of Angola and Zambia. Wallach et al. (2014) considered records from Spawls et al. (2004) from northeastern Tanzania to be unconfirmed.

Ocurrences in Angola (Map 273): The species is limited to the northwestern of the country,

including the Cabinda enclave, although there are no published records from the latter area.

Kwanza Norte: “Piri-Dembos” [-8.56667,

14.50000] ( Hellmich 1957b:76). Kwanza Sul :

“Quirimbo” [-10.68333, 14.26667] ( Parker

1936:126, Manaças 1981:36).

Taxonomic and distributional notes:

This species is often attributed to Shaw (1802),

where the species is description is largely repeated, or to Shaw and Nodder (1792) (e.g.,

Wallach et al. 2014). As noted by McDiarmid et al. (1999), evidence suggests that Shaw alone is responsible for the description. Shaw

(1792) specifically stated that the specimen had been presented to the British Museum from

Edward Jenkins, who had obtained it from “the master of a Guinea vessel. Hoser (2013) erect-

ed a new taxon, B. hoserae , for B. nasicornis MAP 273. Distribution of Bitis nasicornis in Angola. east of the Dahomey Gap, but did not mention Angola or Angolan populations explicitly.

Bitis peringueyi ( Boulenger, 1888) PERINGUEY’S ADDER

Vipera peringueyi Boulenger 1888:141 . Holotype: SAM 1852 (collector L.A. Péringuey), lost fide Stander in Golay et al. (1993). Type locality: “Damaraland, 10 miles east of Walfisch Bay” [= 10 miles east of Walvis Bay], Erongo Region, Namibia.

Bitis peringueyi : Manaças (1981:37), Spawls and Branch (1995:108), Branch (1998:119), Dobiey and Vogel (2007:118), Wallach et al. (2014:92).

Global conservation status (IUCN): Least Concern.

Global distribution: The species occurs limited to sandy regions of the Namib Desert, in two disjunct populations corresponding to the northern and southern sand seas of the Namib. In Namibia from Lüderitz north to Roikop, and then from northern Namibia northward to Namibe

Province, southern Angola.

Ocurrences in Angola (Map 274): The species is limited to the Namib Desert in

Namibe Province. Namibe: “ Porto Alexandre”

[-15.80000, 11.83333] (Spawls and Branch

1995).

Taxonomic and distributional notes:

Boulenger (1896) considered Bitis heraldica

( Bocage, 1889) to be a synonym of

B. peringueyi which resulted in an incorrect distribution range for the species in Angola

(see Bitis heraldica account). The species was cited for southern Angola by several authors

(Manças 1981, Branch 1998, Dobiey and Vogel

2007, Wallach et al. 2014), although the locality “Porto Alexandre”, currently known as MAP 274. Distribution of Bitis peringueyi in Angola.

“Tômboa,” is the first explicit published record for the species in the country (Spawls and Branch 1995). Wallach et al. (2014) erroneously considered B. peringueyi present in Kwanza Sul Province, presumably based on Bogert’s (1940) record of B. heraldica (as “ B. peringueyi ”).