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PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 
Series 4, Volume 65, Supplement II 
(Bogert 1940:99; Themido 1941:11; Mananas 1981:35); “Cuma” [-12.86667, 15.06667] (Loverid- 
ge 1936a:45; Managas 1981:35). Benguela: “Quissange” [-12.43333, 14.05000] (Bocage 
1887c:211); “Benguella” [-12.58333, 13.41667] (Boeage 1866a:53); “Entre Rios” 
[-13.01667, 14.63333] (Hellmich 1957a:75; Managas 1981:35); “Cubal” [-13.03333, 14.25000] 
(Mertens 1937:16, 1938:442; Managas 1981:35); “Equimina” [-13.20000, 14.78333] (Bocage 
1895a:149; Managas 1981:35); “Hanha” [-13.30000, 14.20000] (Bocage 1896a:113; Mananas 
1981:35). Hmla: “Capelongo” [-14.91667, 15.08333] (Bogert 1940:99; Mana 9 as 1981:35); “San- 
geve” [-13.88333, 15.83333] (Monard 19376:141-142; Managas 1981:35); “Rio Calae” 
[-13.96667, 16.03333] (Bocage 1879c:89, 1895a:149; Monad 1937b:141; Managas 1981:35); 
“Kuvangu (Vila-da-Ponte)” [-14.46667, 16.30000] (Monard 1937b:141, 142; Managas 1981:35); 
“Kampulu” [-15.21667, 16.11667] (Monard 19376:141-142; Mananas 1981:35); “Rio Cabindon- 
go” (Bocage 1895a:149; Monad 19376:141; Managas 1981:35). Namibe: “Mossamedes” 
[-15.20000, 12.15000] (Gunther 1865a:480; Managas 1981:35). Cunene: “ruisseau Mbale (Bale)” 
[-15.16667, 16.75000] (Monard 1937b:141-142; Managas 1981:35). Cuando Cubango: “approx¬ 
imately 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 + cau- 
dals), 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 spec¬ 
imens 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 (Anony¬ 
mous 1752) (Bauer and Gunther 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. ari¬ 
etans 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 Mex¬ 
ico 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 hixmdieus, 1758]. 
Vipera {Cerastes) caudalis Smith 1839: pi. 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: Gunther (1865a:480), Bocage (1895a: 150). 
Bitis caudalis: Boulenger (1915:221), Frade (1963:253), Haacke (1984:174), Mananas (1981:37), Visser 
