308 
PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 
Series 4, Volume 65, Supplement II 
Family Viperidae Oppel, 1811 
Genus Athens Cope, 1862 
Athens squamigera (Hallowell, “1854” 1855) Variable Bush Viper 
Echis squamigera Hallowell “1854” 1855:193. Holotype: ANSP 6949 (collector M.P. Bellonni-Duchaillu). 
Type locality: “Near the river Gaboon, Guinea” (Hallowell “1854” 1855:193), Gabon River, Gabon. 
Athens squamigera: Peters (1881:150), Bocage (1887a:189, 1895a:152), Laurent (1954a:62), Loveridge 
(1957:304), Frade (1963:252), Spawls and Branch (1995:99), Broadley (1998a: 128), Ernst and Rodel 
(2002:55), Chirio and LeBreton (2007:596), Dobiey and Vogel (2007:100), Wallach et al. (2014:62) 
Athens Lueani: Rochebmne (1885:89). 
Athens squamiger Boulenger (1905:114, 1915:222), Ferreira (1906:169). 
Athens squamigera squamigera (Hallowell): Laurent (1954a:62), Hellmich (1957b:76), Managas (1981:39). 
? Athens squamigera squamigera (Hallowell): Laurent (1964a: 128). 
Global conservation status (lUCN): Not Evaluated. 
Global distribution: The species is endemic to West and Central Africa. 
Ocurrences in Angola (Map 268): The 
species occurs in northern Angola, including 
the Cabinda enclave. Cabinda: “Foz do Luan- 
go” [-5.15000, 12.16667] (Mana9as 1981:39); 
“Landana” [-6.21667, 12.15000] (Rochebmne 
1885:90, Loveridge 1957:304, Mananas 
1981:39). Lunda Norte: “Riv. Kundueji, envi¬ 
rons de Dundo, Lunda” [ -7.36667, 20.83333] 
(Laurent 1964a: 128); “Dundo, rives de la Lua- 
chimo” [-7.38333, 20.85000] (Laurent 
1954a:62; Managas 1981:39); “foret de la Lua- 
chimo” [-7.38333, 20.85000] (Laurent 
1964a: 128). Kwanza Norte: “Piri-Dembos” 
[-8.53333, 14.43333] (Hellmich 1957b:76; 
Managas 1981:39); “Golungo Alto” [-9.13333, 
14.76667] (Boulenger 1905:114; Ferreira 
1906:169; Mananas 1981:39). Undetermined 
Locality: “Cuango” (Peters 1881:150; Bocage 
1887a:189; 1895a:152; Managas 1981:39) 
(Malanje Province impossible to georeference: See History Section for more detailed information). 
Taxonomic and distributional notes: The date of the description of this taxon has variably 
been given as 1854, 1855 (Wallach et al. 2014), or 1856 (McDiarmid et al. 1999; Broadley 1998a). 
Although the contents of volume 7 of the Proceedings of the Academy of Natural Sciences of 
Philadelphia covers the years 1854 and 1855 and the printed cover page indicates 1856, part V of 
the volume, in which Hallowll’s paper appeared, was printed in 1855. Several taxonomic revisions 
of the genus Atheris Gray, 1842 were undertaken by Broadley (1996a, 1998a), but the taxonomic 
status of several entities remain to be established (Ernst and Rodel 2002). A re-examination of 
museum specimens is necessary to determine if Atheris squamigera (Hallowell, “1854” 1855) is a 
geographically variable species or a species complex (Broadley 1998a; B. Hughes in Ernst and 
Rodel 2002). 
12*E I^E 24°E 
Map 268. Distribution of Atheris squamigera in 
Angola. 
