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PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 
Series 4, Volume 65, Supplement II 
(Monard 1937b:146; Loveridge and Williams 1957:396). Namibe: “Capangombe” [-15.10000, 
13.15000] (Boeage 1895a:2; Loveridge and Williams 1957:396; Ceriaco et al. 2016a:54). Cuando 
Cubango: “Kakindo” [-15.45000, 17.05000] (Monard 1937b: 146; Loveridge and Williams 
1957:396). 
Taxonomic and distributional notes: There is some eonfusion surrounding the deseription of 
K. belliana (Bour 2006). Gray (1831a) in his Synopsis Reptilium. Cataphracta, the most frequent¬ 
ly cited source of the description (e.g.. King and Burke 1989), includes in his section “Additions 
and Corrections” a relatively detailed description headed "'Kinixys Belliana, Gray, Griffith, Trans. 
Animal Kingd.” In the other publication (Gray 1831b) a very short description is provided under 
the heading “Bell’s Tortoise. Test. {Kinixys) Belliana, Gray.” The latter work has a note by Gray 
dated “Oct. 1830,” whereas the first has a preface dated “Jan. 1831.” The title pages for both works 
are dated 1831. Thus it appears that the shorter description was written first, although this does not 
clarify which version was published first. We here cite the more extensive description in the Cat¬ 
aphracta (Gray 1831a). For some time all southern African hinged-back tortoises were treated as 
a single species, Kinixys belliana Gray, 1831 (Loveridge and Williams 1957). However, Pritchard 
(1979) suggested that a more detailed investigation might confirm the validity of some of the form 
described earlier as Kinixys spekii Gray, 1863. Broadley (1981b) divided K. belliana into two sub¬ 
species, the typical form being restricted to the coastal plain, with the depressed form K b. spekii 
replacing it inland. He later recognized the two as specifically distinct (Broadley 1989a, 1993). 
Broadley (1989a) provided a distribution map for K spekii depicting Angolan records previously 
assigned to K belliana. However, he later stated that K. belliana extends to Angola (Broadley 
1992a), whereas the presence of K. spekii had not yet been confirmed (Broadley 1993). Iverson 
(1986, 1992) placed K. spekii as a synonym of K. belliana and provided a map with all the known 
records from the country assigned to the latter. Mifsud and Stapleton (2014) and the Turtle Taxon¬ 
omy Working Group (2014) suggested that K. belliana occurs wiidely in Angola exclusive of the 
southeast and the true desert, whereas K. spekii is limited to the southeastern border areas with the 
Caprivi Strip and Zambia. On the other hand, Fritz and Havas (2007), Branch (2008) and Vetter 
(2011) considered K. belliana to be extralimital, with K. spekii widespread in Angola. 
A recent study of phylogeography, phytogeny and taxonomy of hinged-back tortoises (Kindler et 
al. 2012) found that the previously recognized savanna species K. belliana comprises three deeply 
divergent clades that are now treated as distinct species. Angolan populations are members of the 
clade that retains the name K. belliana, whereas West African and southeast African clades have 
been allocated to K. nogueyi (Lataste, 1886) and W zombensis Hewitt, 1931, respectively. Kindler 
et al. (2012) did identify K. spekii in Angola near the Namibian border, however, so it is highly like¬ 
ly that it occurs at least in southeastern Angola, although pending further investigation we list all 
K. belliana/K. spekii records together. 
Kinixys belliana Gray, 1831 Bell’s Hinge-Back Tortoise 
Kinixys Belliana Gray, 1831a:69. Holotype: BMNH 1979.919 (collector unknown). Type locality: “?” 
[“bought at Humphrey’s, West Africa” according the label on the type specimen fide King and Burke 
(1989)]. 
Cinixys belliana'. Boeage (1866a:40, 1895a:2), Boeage (1887c:209), Monard (1937b: 146). 
Kinixys belliana'. Schmidt (1933:4), Laurent (1950a: 13), Iverson (1986:158, 1992:271), King and Burke 
(1989:89), Broadley (1992a:12), Kindler et al. (2012:193), Turtle Taxonomy Working Group (2014:402), 
Ceriaco et al. (2016b:54). 
Kinixys belliana belliana'. Mertens (1937a:5, 1938a:430), Hellmich (1957a:32), Loveridge and Wiliams 
(1957:384), Loveridge (1957:170), Laurent (1964a:25), Broadley (1981b:208, 1992a: 13, 1993:47). 
Kinixys cf belliana'. Conradie et al. (2016:27). 
