Kinixys belliana (Bour, 2006)

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 : 169-172

Kinixys belliana and Kinixys spekii

The precise delineation of these two species of Kinixys in Angola remains unclear and genetic data will be necessary to establish species boundaries (U. Fritz, pers. comm.). To date only K. belliana has been genetically verified in Angola but it is highly likely, given the distribution of K. spekii elsewhere, that it is also present in the country (see notes below). Collectively the species are widespread, although absent from desert and forest habitats.

Ocurrences in Angola (Map 116): The following records combine both K. belliana and K. spekii (see K. spekii account below for the specific reference to its records in the cher-

sonyms list). Lunda Norte: “Muita (Luembe

E)” [-7.80000, 21.45000] ( Laurent 1950a:13;

Loveridge and Williams 1957:396). Lunda

Sul: “Alto Chicapa” [-10.88333, 19.23333]

( Laurent 1964a:25); “Lunda” [-10.96667,

20.06667] ( Monard 1937b:146; Loveridge and

Williams 1957:396). Moxico: “environs du lac

Calundo” [-11.80000, 20.86667] ( Laurent

1964a:25). Malanje: “Duque de Bragança ”

[-9.10000, 15.95000] ( Bocage 1866a:40,

1895a:2; Loveridge and Williams 1957:396);

“ Muata-Yamv, Lui River ” [-9.41667,

17.33333] (Loveridge and Williams 1957:396);

“Capanda” [-9.72841, 15.34585] (Ceríaco et al. MAP 116. Distribution of Kinixys belliana and Kinixys spekii in Angola.

2014b:670); “Cangandala National Park”

[-9.84606, 16.72233] (Ceríaco et al. 2016b:53). Bié: “Gauca” [-11.18333, 17.45000] ( Laurent 1964a:25; Loveridge and Williams 1957:396); “Chitau” [-11.43333, 17.15000] ( Schmidt 1933:4; Loveridge and Williams 1957:396); “Cuemba” [-12.15000, 18.08333] ( Kindler et al. 2012, supporting information:4); “Kuito” [-12.383599, 16.928029] ( Kindler et al. 2012, supporting information:4); “Cubando basin (13)” [-13.28061, 16.74722] ( Conradie et al. 2016 a:8-9, 27). Huambo: “Galanga” [-12.06667, 15.15000] ( Bocage 1895a:2; Loveridge and Williams 1957:396). Benguela: “Quissange” [-12.43333, 14.05000] ( Bocage 1887b:209, 1895a:2; Loveridge and Williams 1957:396); “Benguella” [-12.58333, 13.41667] ( Hellmich 1957a:32); “Ebanga” [-12.73333, 14.73333] ( Monard 1937b:146; Loveridge and Williams 1957:396); “Cubal” [-13.03333, 14.25000] ( Mertens 1937a:5, 1938a:430; Loveridge and Williams 1957:396). Huíla: “Caconda” [-13.73333, 15.06667] ( Bocage 1895a:2; Loveridge and Williams 1957:395); “Kuvangu” [-14.46667, 16.30000] ( Monard 1937b:146); “Osi” [-15.08333, 15.41667] ( Monard 1937b:146; Loveridge and Williams 1957:396). Cunene: “riv. Mbalé ” [-15.16667, 16.75000]

( Monard 1937b:146; Loveridge and Williams 1957:396). Namibe: “Capangombe” [-15.10000, 13.15000] ( Bocage 1895a:2; Loveridge and Williams 1957:396; Ceríaco 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 confusion surrounding the description of K. belliana ( Bour 2006) . Gray (1831a) in his Synopsis Reptilium. Cataphracta, the most frequently 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 Cataphracta ( 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 subspecies, 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 1989 a, 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 Taxonomy 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 Havaš (2007), Branch (2008) and Vetter (2011) considered K. belliana to be extralimital, with K. spekii widespread in Angola. A recent study of phylogeography, phylogeny 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 K. zombensis Hewitt, 1931 , respectively. Kindler et al. (2012) did identify K. spekii in Angola near the Namibian border, however, so it is highly likely 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 : Bocage (1866a:40, 1895a:2), Bocage (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), Ceríaco 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) .

Global conservation status (IUCN): Not Evaluated [Least Concern 1996] Global distribution: We here follow the distributions presented by Kindler et al. (2012) in their supporting information. The species ranges from the Red Sea coast of Sudan and the Horn of Africa south to the Kenya-Tanzania border area and thence southwest to central Angola.

Kinixys spekii Gray, 1863 SPEK’S HINGED-BACK TORTOISE

Kinixys spekii Gray, 1863a:381 . Holotype: BMNH 1936.5.3.117 (collector J. H. Speke). Type locality: “Central Africa” probably “northwestern Tanzania ” fide Broadley and Howell (1991). Kinixys belliana spekii : Broadley (1981b:211). “ Kinixys spekii ”: Broadley (1989a:52). Kinixys spekii : Fritz and Havaš (2007:286), Branch (2008:53), Vetter (2011:79), Kindler et al. (2012:193),

Ceríaco et al. (2014b:670), Turtle Taxonomy Working Group (2014:403), Ceríaco et al. (2016b:53).

Global conservation status (IUCN): Not Evaluated.

Global distribution: We here follow the distributions presented by Kindler et al. (2012) in their supporting information as modified in The Turtle Texonomy Working Group 2017). The species is known from southern Central Africa from Kenya and Uganda south to northern KwaZulu-Natal, South Africa (although not in the coastal plain and lowlands of southeastern Kenya, Tanzania and Mozambique) and west to the Caprivi Strip of Namibia and adjacent Cuando Cubango, Angola.

Kinixys erosa ( Schweigger, 1812) FOREST HINGE-BACK TORTOISE

Testudo erosa Schweigger 1812:321 . Syntypes: Two specimens originally in the Mannheim Museum and a specimen described and figured by Shaw (1802:59, pl. 13) (collectors unknown). Type locality: “America septentrionali” [in error].

Cinixys erosa : Bocage (1866a:40, 1895a:1), Peters (1877a:611).

Kinixys erosa : Laurent (1964a:24), Iverson (1986:159, 1992:272), Spawls et al. (2004:52), Fritz and Havaš 2007:285, Branch (2008:50), Vetter (2011:75), Kindler et al. (2012:193), Luiselli and Diagne (2014:084.1), Mifsud and Stapleton (2014:26), Turtle Taxonomy Working Group (2014:402).

Global conservation status (IUCN): Data Deficient.

Global distribution: A widespread silvi-

colous species in West Africa, preferring moist areas such as marshes and river banks from

Gambia eastward through the Democratic

Republic of Congo to Uganda, and south areas adjacent to the mouth of the Congo River .

Ocurrences in Angola (Map 117): This species is limited to extreme northern regions of the country, including Cabinda. Cabinda:

“Chinchoxo” [-5.10000, 12.10000] ( Peters

1877a:611); “ Cabinda ” [-5.55000, 12.18333]

( Bocage 1866a:40, 1895a:1; Mifsud and Stapleton 2014:31). Lunda Norte: “près de la R.

Camuálua (ou Camuáli), affl. de la rive gauche du Chicapa, à l’Ouest de Dundo, Posto de

Lóvua” [-7.31667, 20.15000] ( Laurent

1964a:24; Luiselli and Diagne 2014:084.6;

Mifsud and Stapleton 2014:31); “affl. non MAP 117. Distribution of Kinixys erosa in Angola. determine du Chicapa, à peu près dans la meme region que le precedent” (which represent the locality cited above: “affl. de la rive gauche du Chicapa, à l’Ouest de Dundo , Posto de Lóvua “) ( Laurent 1964a:24); “Cambulo, 70 km from the border with Democratic Republic of Congo, Kinshasa” [-7.73333, 21.26667] (Luiselli and Diagne 2014:084.6; Mifsud and Stapleton 2014:31) .

Taxonomic and distributional notes: Bour (in Iverson 1992) stated that the syntype figured by Shaw might have been in the Leverian Museum. This collection was sold at auction in 1806 and dispersed to a number of private and state-owned museums, but this specimen has not been located.