D.G.BROADLEY Natural History Museum, Bulawayo, Zimbabwe

Transcription

1 A REVIEW OF THE SOUTHERN AFRICAN SPECIES OF KINIXYS BELL (REPTILIA: TESTUDINIDAE) D.G.BROADLEY Natural Histry Museum, Bulaway, Zimbabwe Bradley, D. G., A review f the Suthem African species f Kinixys Bell (Reptilia: Testudinidae). Annals f the Transvaal Museum 36(6): Examinatin f all the Kinixys Bell material frm Suthem Africa suth f 22 S available in museum cllectins has shwn that fur species ccur in this regin. On the basis f its prly develped plastral hinge and gular lip, K. natalensis Hewitt is cnsidered the mst primitive species in the genus. It has a restricted distributin alng the eastern escarpment f Suth Africa frm the Transvaal suth t Natal. Kinixys spekii Gray seems t be the next mst primitive species, but its depressed carapace is an adaptatin that enables it t find refuge in rck crevices. It has a very extensive distributin n the central African plateau. Kinixys belliana Gray is a larger species with a mre cnvex carapace and a black radial pattern n the shields. Its distributin is restricted t the castal plain. Kinixys lbatsiana Pwer differs frm the ther savanna species, and resembles the frest species K. ersa (Schweigger) and K. hmeana Bell, in having the psterir marginal shields recurved and serrated. Its distributin is limited t the Transvaal Bushveld and sutheastem Btswana. Keywrds: Suthem Africa, systematics, Reptilia, Testudinidae, Kinixys. INTRODUCTION In the last cmprehensive revisin f the African hinged trtises f the genus Kinixys Bell Lveridge and Williams (1957) recgnized a single savanna species, K. belliana Gray. Bradley (1981) reviewed the ppulatins f Kinixys fund suth f 25 S and east f 30 E, reinstating K. natalensis Hewitt as a full species and reviving spekii Gray as a subspecies f K. belliana Gray. Subsequent examinatin f mre material f savanna Kinixys frm thrughut Africa, including mst f the material in United States museums, suggested that K. spekii shuld be treated as a full species and K. lbatsiana Pwer shuld als be resurrected as a full species (Bradley, 1989). Meanwhile, Byctt and Jacbsen (1988) cnfirmed the validity f K. natalensis and established that its range extends nrth int eastern Transvaal. In April 1991 I examined the large cllectin f Kinixys material in the Transvaal Museum. This firmly established the validity f K. lbatsiana as a full species with a restricted distributin and with n evidence f sympatry with any ther species. In Suth Africa I btained dry shells f K. lbatsiana and K. natalensis fr preparatin as skeletal specimens fr cmparisn with similar material f the ther species in the genus. This has helped t clarify relatinships. MATERIALS AND METHODS This study is restricted t ppulatins f Kinixys fund suth f 22 S, i.e., sutheastern Btswana, Suth Africa, suthern Mzambique and Swaziland (there are n recrds frm the Limpp valley n the Zimbabwe side). The mst useful diagnstic characters are the frm f the beak, the shape f the psterir marginals (especially the supracaudal), the drsal length f the epiplastrn in relatin t ttal plastrn length, the rati f shell height t shell length, and carapace pattern. The ratis used by Laurent (1956, 1962) in his diagnsis f K. belliana mertensiwere calculated fr the suthern frms. The measurements were taken with dial callipers. Gular length and width are maximum measurements, but measurements f mid-ventral sulci are average values in the many cases when shield margins are nt precisely juxtapsed. Specimens less than 75 mm in shell length have atypical prprtins and were nt used in the statistical analyses. Shell length t the nearest millimetre between blcks (with carapacial hinge clsed) was used as the standard measurement f length, as plastrn length is dependent n the very variable develpment f the gulars. Maximum shell height was als measured between blcks. The nmenclature f bnes and shields fllws Zangerl (1969). Tw hundred and fifteen specimens were examined frm the fllwing institutins (abbreviatins preceding catalgue numbers are indicated in parentheses): Albany Museum, Grahamstwn (AM); Field Museum f Natural Histry, Chicag (FMNH), Natural Histry Museum, Lndn (BM); McGregr Museum, Kimberley (MMK); Museum Annals f the Transvaal Museum. Vlume 36, Part 6 Annale van die Transvaal Museum. Band 36. Deel 6 June 1993 Junie 1993 ISSN

2 42 ANNALS OF THE TRANSVAAL MUSEUM N natalensis 4, I I I, spe k i i 15 I BE, b.belliana 19 I ~ lbatsiana 9 I, ~ %, Fig. 1 Variatin in the rati drsal epiplastrn length/plastrn midline length in Suthern African ppulatins f Kinixys. Hrizntal line = range, vertical line = mean, slid rectangle = ± ne standard deviatin; hllw rectangle = ± ne standard errr. d'histire Naturelle, Geneva (MHNG); Natal Museum, Pietermaritzburg (NM); Natural Histry Museum f Zimbabwe, Bulaway (~IMZB); Suth African Museum, Cape Twn (SAM); Transvaal Museum, Pretria (TM). A figure in parentheses after a catalgue number indicates the number f specimens in a series. CHARACTER AI\JALYSIS BEAK. Kinixys natalensis has a strngly tricuspid beak, thus differing frm all ther species in the genus, which have unicuspid beaks. SHAPE OF POSTERIOR MARGINALS. Kinixys lbatsiana has the psterir marginals strngly recurved and serrated like the frest species K. ersa and K. hmeana. In K. natalensis the psterir marginals are slightly reverted and feebly serrated (strngly serrated in juveniles). In K. spekii and K. b. belliana the lateral psterir marginals may be slightly recurved, but they are never serrated. SHAPE OF SUPRACAUDAL SHIELD AND PYGAL BONE. In the Natal ppulatins f K. natalensis, the supracaudal is usually divided, the tw halves being separated by a distinct ntch; in the Swaziland and Transvaal ppulatins the supracaudal is usually undivided. In K. spekiithe supracaudal (and underlying pygal bne) is usually distinctly narrwed mesially r prximally excavate. In K. b. belliana the supracaudal is nt narrwed mesially, but may have a shallw median depressin inferirly, widest prximally. In K. lbatsiana the supracaudal is widened mesially, with a distinct lngitudinal grve inferirly, widest prximally. => 2 u.. 0 ~I Fig ~ SPEKII B. BELl/ANA I 2 ANAL SULCUS - -. :~ 1'1 Scatter diagram f midline length f femral sulcus and anal sulcus fr tw Suthern African Kinixys species. The scale is in cm. DORSAL EPIPLASTRON LENGTH/PLASTRON MIDLINE LENGTH. In skeletal material, this rati readily distinguishes K. natalensis (10-15 %) frm K. spekii (15-19 %), K. b. belliana (18-25 %) and K. lbatsiana (18-22 %). The bserved range f variatin is shwn in Fig. 1. RATIO SHELL LENGTH/HEIGHT. The range f variatin is shwn in Table 1. This rati will usually distinguish K. spekii frm K. b. belliana, the ther tw species being intermediate.

3 BROADLEY: SOUTHERN AFRICAN KINIXYS 43 NATALENSIS 5 P E KII B. BE L L I A NA LOBATSIANA Fig.3 Histgrams shwing distributin f specimens by sex in each shell length size class fr Suthern African ppulatins f Kinixys. The hrizntal scale is in cm, e.g., Class 12 = mm. The smallest vertical divisin indicates ne specimen. CARAPACE PATTERN. Kinixys nata/ensis and K. spekii have a znary pattern n each drsal shield, the arelae are dark and are encircled by a pale zne, then a dark zne, ften with a ragged r radiate uter margin. The dark markings gradually disappear in adult male K. spekii and sme male K. nata/ensis. Kinixys b. belliana has a radiate pattern n each drsal shield, the dark arelae are surrunded by a pale zne, and brad black rays (usually six) extending t the margins f each shield. Kinixys /batsiana females usually have a radiate pattern n the drsal shields, but this is ften fragmented r vestigial, while adult males are usually devid f pattern. Btswana specimens f bth sexes are almst devid f markings. GULAR LENGTHIWIDTH RATIO. In K. nata/ensis the gular shields (tgether) are usually mre than twice as wide as lng, whereas in the ther three species they are less than twice as wide (Table 1). Kinixys spekii differs frm the ther species in frequently having the gulars frked anterirly, especially in males. PROPORTIONS OF PLASTRAL SHIELDS. Variatin in the ratis used by Laurent (1956, 1962) in his diagnsis f K. bel/iana mertensi (cnsidered a synnym f the typical frm) in nrtheastern Zaire and Uganda is shwn in Table 1: nne are diagnstic. Hwever, suthern ppulatins f K. b. belliana usually have the median femral sulcus lnger than the anal sulcus, whereas this relatinship is reversed in K. spekii (Fig. 2). SIZE Al\ID SEXUAL DIMORPHISM. Shell lengths fr the fur species are shwn in Fig. 3. Kinixys nata/ensis is the smallest species and K. belliana the largest. Kinixys nata/ensis females attain a larger size than males and this may als be the case in K. spekii. There is n bvius sexual dimrphism in K. belliana and K. /batsiana. CARAPACIAL HINGE. In K. nata/ensisthe hinge is restricted t the peripheral bnes, there is hardly any interventin f cartilage between the furth and fifth cstal bnes, even in the largest adults.

4 44 ANNALS OF THE TRANSVAAL MUSEUM Table 1 Variatin in prprtins f the shell and varius plastral shields in Kinixys ppulatins suth f 22" S. n Range Mean S.D. G.V. Shell lengthlheight rati nata/ensis 33 1,97-2,51 2,18 0,12 5,50 spekii 57 2,17-2,69 2,46 0,13 5,28 belliana belliana 40 1,82-2,34 2,10 0,12 5,71 /batsiana 68 1,95-2,59 2,28 0,15 6,58 Gular lengthlwldth rati nata/ensis ,06 10,12 23,50 spekii ,64 9,48 15,90 belliana belliana ,20 9,55 14,64 /batsiana ,39 10,68 18,61 Gular + humeral sulcus/plastrn width at humer-pectral sulcua (%) nata/ensis 30 56,0-81,1 68,40 6,40 9,36 spekii 58 54,5-90,6 70,26 8,19 11,66 belliana belliana 40 57,1-84,9 71,54 6,74 9,42 /batsiana 68 60,7-110,3 76,57 7,22 9,43 Pectral sulcus/gular + humeral sulci (%) nata/ensis 32 16,3-40,0 31,17 5,96 19,12 spekii 60 25,9-63,6 41,24 7,80 18,91 belliana belliana 40 28,6-64,7 45,35 9,21 20,30 Ibatsiana 68 18,6-57,1 32,30 7,55 23,37 Pectral sulcus/abdminal sulcus (%) nata/ensis 33 13,7-38,9 27,26 5,74 21,06 spekii 62 25,8-62,2 39,04 7,31 18,72 belliana belliana 40 30,0-62,9 43,56 7,85 18,02 /batsiana 66 16,3-58,5 32,89 6,87 20,89 In K. spekii the hinge is mderately develped, with cartilage rarely extending mre than half-way up the anterir edge f cstal 5. In K. belliana and K. lbatsiana the hinge is well develped, with cartilage extending alng the anterir edge f cstal 5 and ften arund the margins f neural 4. NUMBER OF MARGINALS. Kinixys natalensis usually has mre than 23 marginals, due t divisin f the supracaudal and/r insertin f extra marginals n each side. The ther species usually have 23 marginals, with rare variatins due t fusin r divisin f individual shields (Table 2). SYSTEMATICS KEY TO THE SOUTHERN AFRICAN SPECIES OF KINIXYS Beak tricuspid; carapacial hinge prly develped, nt extending beynd the marginals; gulars (tgether) usually at least twice as wide as lng; drsal epiplastrn length less than 15 % f plastrn midline length; marginals usually 24 r mre, with supracaudal frequently divided, the tw halves separated by a deep ventral ntch; plastrn nt cncave in males... natalensis Beak unicuspid; carapacial hinge well develped in adults; gulars (tgether) less than twice as wide as lng; drsal epiplastrn mre than 15 % f plastrn midline length; marginals usually 23, supracaudal nt divided; plastrn cncave in males Psterir marginals reverted and serrated, underside f supracaudal usually with a lngitudinal grve, widest anterirly... lbatsiana Psterir marginals nt reverted r serrated, underside f supracaudal usually smth r prximally excavate... 3

5 BROADLEY: SOUTHERN AFRICAN KINIXYS 45 3 Carapace distinctly depressed, shell height usually included in length mre than 2,3 times; gular lip usually cncave abve in crss-sectin, gular scutes ften frked; drsal epiplastrn length % f plastrn midline length; underside f supracaudal narrwed mesially r prximally excavate; median anal suture usually lnger than femral suture; shields f carapace with a basically znary pattern, adult males ften unifrm... spekii Carapace cnvex, shell height usually included in length less than 2,3 times; gular lip usually flat r cnvex abve in crss-sectin, gular scutes nt frked; drsal epiplastrn length % f plastrn midline length; underside f supracaudal nt narrwed mesially r prximally excavate; median femral suture usually lnger than anal suture; shields f carapace with a radial pattern... bel/iana be/liana Table 2 Number f marginal shields in Kinixys ppulatins suth f 22 S. n natalensis 41 spekii 61 bel/iana bel/iana 40 lbatsiana Kinixys natalensis Hewitt, Fig. 5 Kinixys belliana zuluensis (part) Hewitt, 1931: 475, pi. xxxviii, figs Kinixys natalensis Hewitt, Recrds f the Albany Museum 4: 353. pi. xxxv, figs 3-4 (type lcality: Jamesn's Drift, Tugela River. Natal); Bradley, 1981: 204, figs 7-B; Byctt and Burquin, 1988: 127. pi , fig. 28; Byctt, 1988: 90; Lambiris. 1988: 32; Byctt and Jacbsen. 1988: 93. figs 1--4; Branch, 1988: 31. pi. 5; Bradley, 1989: 60. fig. 15; Ernst and Barbur. 1989: 232; Crumly, 1989: 91. Kinixys belliana belliana (part): Lveridge and Williams. 1957: 384; Wermuth and Mertens, 1961: 180, 1977: 70. DIAGNOSIS. Beak tricuspid; carapacial hinge nt extending beynd marginals; gulars (tgether) usually at least twice as wide as lng; drsal epiplastrn length less than 15 % f plastrn midline length; marginals usually 24 r mre, with supracaudal frequently divided; plastrn nt cncave in males. DESCRIPTION. Beak tricuspid; prefrntallngitudinally divided; frntal usually entire, smetimes lngitudinally divided; frelimb cvered with subimbricate scales, a few enlarged and pinted; five claws n frnt ft. Carapace mderately cnvex (shell length/ height rati 1,97-2,51), ften with flat-tpped prtuberances n vertebrals 4 and 5; anterir margin nt expanded, psterir marginals slightly reverted and feebly serrated in adults (serrated in juveniles); drsal shields ften raised, with wellmarked grwth annuli; nuchal elngate in adults, ften minute drsally; vertebrals 5, rarely 6 (six specimens); pleurals 4 (five in a hatchling and n ne side f three specimens); marginals 23-26, supracaudal usually divided in Natal specimens, the tw halves separated by a distinct ntch. Frnt lbe f plastrn truncate anterirly, nt r slightly prjecting beynd anterir brder f carapace; gulars paired, usually wider than lng; pectrals with a narrw median sulcus; axillaries (usually three) small; inguinal large (absent nly in AM 6975 G), in cntact with r separated frm sixth marginal, in cntact with femral; rear lbe shrt and truncate, with a very shallw psterir ntch. Adult males withut a cncave plastrn. COLOURATION. Carapace with a znary pattern n each shield, the arelae brwn, fllwed by a brad range/yellw zne, then a blackish zne which may be brken up int shrt rays that may extend t the edges f the shield r be fllwed by anther range/yellw zne. The plastrn is yellw with symmetrical blackish markings, a pair f black rings n the abdminal shields being the

6 46 ANNALS OF THE TRANSVAAL MUSEUM mst prminent. The clur patterns becme brken up and ill-defined in large adults and may disappear cmpletely. SIZE. Largest male (TM 65799, Hedspruit, Transvaal) 127 mm lng, 89 mm wide, 57 mm high; largest female (AM 6975B, Jamesn's Drift, Natal) 155 mm lng, 113 mm wide, 70 mm high. HABITAT. Thrughut its range, K. natalensis is assciated with rcky areas alng the eastern escarpment at metres altitude. In the eastern Transvaal it has been fund amng granite utcrps. The habitat includes valley bushveld in the suth, muntain bushveld and muntain thrnveld in nrthern l'jatal, and arid bushveld and arid lwveld in the extreme nrth f its range (Byctt and Jacbsen, 1988). During the day these trtises have been fund under rcks n the summit f the Lebmb Range in Swaziland (Byctt, 1988). DISTRIBUTION. The eastern plateau slpes frm, the eastern Transvaal at Hedspruit suth t the Natal Midlands at Grey twn. Lambiris (1988) has recrded a specimen frm Imbali, near Pietermaritzburg, but he cncedes (in litt.) that it 'was prbably an intrduced specimen. It had been brught t the Natal Parks Bard by an Imbali resident, wh fund it n the radside, and there is n independent evidence f a natural ppulatin in the area.' LOCALITIES. (An * indicates sympatry with K. spekii, a # indicates sympatry with K. b. belliana.). SOUTH AFRICA. Transvaal: Bergplaats 25HU TM 69917; Hedspruit TM 65799; Mananga Kp TM 69915; Manyeleti Game Reserve* (Main Camp) TM 64809; Sabi-Sand/Kruger Nat. Park bundary, S f Tswariri River TM Natal: n lcality I\IMZB ; Dimane Stream AM - (4); Greytwn AM 77, NM 1130; Imbali (Lambiris, 1988); Impanza NM 1143 (2); Itala Game Reserve (Lambiris, 1988) TM 57520; Jamesn's Drift AM 6975( 12), BM ; ± 30 km S f Kranskp (Byctt and Jacbsen, 1988); Manaba# (Hewitt, 1935); Ntambanana (Hewitt, 1931); Othbthini# TM 19346, 69916; Uitkyk, Vryheid Dis!. TM 56755; Weenen Nature Reserve NMZB 11071, TM 50682; 'Zululand' NM SWAZI LAND. Big Bend FMNH ; Grenpan Farm (phts: J. Culverwell); Mbuluzi Game Reserve* and Mlawula Nature Reserve* (J. Culverwell in Byctt and Jacbsen, 1988); Tshaneni FMI\IH MOZAMBIQUE. N lcality TM 47208; 10 km SSE f Ressan Garcia* NMZB-UM Kinixys spekii Gray, Fig. 5 Kinixys spekii Gray, 1863, Annals and Magazine f Natural Histry(3)12: 381 (type lcality: Central Africa); Bradley, 1989: 52, fig. 12b. Kinixys australis Hewitt, 1931: 477, pi. xxxvi, figs 4-{) (type lcality: White River, E Transvaal). Kinixys belliana belliana (nt Gray): Lveridge and Williams, 1957: 384 (part); Wermuth and Mertens, 1961: 180(part); Pienaar, 1966: 127; Switak, 1971: 9; Wermuth and Mertens, 1977: 70 (part); Pienaar, 1978: 213. Kinixys belliana spekii: Bradley, 1981: 211, figs 11-12; Pienaar et al., 1983: 22, pi. 2; Byctt and Burquin, 1988: 124 (part), pi , fig. 27; Byctt and Jacbsen, 1988: 100; Branch, 1988: 30, pi. 5; Ernst and Barbur, 1989: 230. DIAGNOSIS. Beak unicuspid; carapacial hinge well develped; gulars (tgether) usually less than twice as wide as lng; drsal epiplastrn length % f plastrn midline length; marginals usually 23, the supracaudal undivided; carapace depressed, shell height usually included in length mre than 2,3 times; plastrn cncave in males; clur pattern f carapacial shields basically znary. DESCRIPTION. Beak unicuspid; prefrntal usually lngitudinally divided; frntal entire; frelimbs cvered with subimbricate scales, many enlarged and pinted; five claws n frnt ft. Carapace depressed (shell length/height rati 2,17-2,70), highest pint frequently n vertebral 1 r 2; drsal shields relatively flat, arelae f vertebrals usually with a weak lngitudinal keel; anterir margin nt expanded, lateral psterir marginals slightly reverted, nt serrated; nuchal elngate in adults; vertebrals 5 (six in AM 780 nly), pleurals 4 (three in TM and 65800; five in AM 780 and ne side f TM and 65699); marginals 23 (rarely 22 r 24). Frnt lbe f plastrn truncate anterirly, prjecting well beynd the anterir brder f the carapace in bth sexes; gulars paired, usually less than twice as brad as lng; pectrals with a mderate median sulcus; axillaries tw, f mderate size; inguinal large, usually in cntact with sixth marginal, in brad cntact with femral; hind lbe shrt and truncate, with a very shallw psterir ntch. Plastrn cncave in adult males. COLOURATION. Carapace with a znary pattern n each shield in juveniles and subadults. This may persist in adult males, but many becme unifrm live-brwn t buff. In females the dark znes tend t break up int ragged radiatins. The plastral shields usually have ragged circular radiating patterns in subadults and adult females, but these becme vestigial r absent in adult males.

7 BROADLEY: SOUTHERN AFRICAN KINIXYS 47 SIZE. Largest male (TM 41761, Maput Elephant Reserve, Mzambique) 181 mm lng, 124 mm wide, 79 mm high; largest female (AM 78C, White River, Transvaal: lecttype f K. australis) 183 mm lng, 115 mm wide, 70 mm high. HABITAT. This species has a wide range in trpical savannas, and suth f the Limpp it is fund in the lwveld areas, including Mpaniveld. Its depressed carapace enables this trtise t use rck crevices as refuges. DISTRIBUTION. In Suth Africa this species inhabits the lwveld areas f the nrthern and eastern Transvaal, extending suth thrugh Swaziland and adjacent Mzambique t Zululand. The suthernmst recrd frm nrth f Mehlwemamba, Hlabisa District, is based n the shell f a juvenile and needs verificatin. LOCALITIES. SOUTH AFRICA. Transvaal: n lcality ('N. Transvaal') TM 12768, ; Acrnhek TM 66326; Alldays TM 37638; Antich 240KT TM 64666; Bekaf 650MS TM 52485; De Hp 203JU TM 42393; Dwaalbm 217KP TM 69898; Hartbees TM 68243; Hilltp, Nelspruit TM 67469; Inkerman 10KQ TM 69899; Jerme 287MT TM 52217; Ka Mininginisi TM 52218; Kruger Natinal Park (lcalities frm Pienaar, 1966, fig. 32); Leamingtn 207KU TM 65775; Malep TM 6987; Manyeleti Game Reserve QDS 2431C2TM 65110, 65699, 66520, 69846, 69890, and QDS TM 65747, 65794, 65800, 65810; Merry Pebble Stream 246KU TM 69896; Mmableta Estate TM ; Mkeetsi-Duiwelsklf TM 21842; Nelspruit TM 67600; Nzulase TM 69894; Ostend 104KT TM 52161; Rlle 235KU TM 69900; SA Bantu Trust TM 69901; Seville 224KU TM 65563; The Glen 259 TM 67090; Uitkmst 769LS TM 69895; Vulcanus 584LS TM 69897; White River AM 78(5), Zululand: nrth f Mehlwemamba TM 64863; Ndumu Game Reserve TM SWAZILAND. Manzini TM 24205, 42735; 5 km NW f Siphfaneni TM 51023; Tambuji Estates TM 58100; Tshaneni BM , NMZB-UM MOZAMBIQUE. Maput Elephant Reserve TM 41761; Mamba TM ; 10 km SSE f Ressan Garcia NMZB-UM , 30455, 33010; 15 km SSE f Ressan Garcia NMZB-UM Kinixys belliana belliana Gray, Fig. 6 Kinixys Belliana Gray, 1831, Synpsis Reptilium: 69 (n lcality); Burquin, Vincent and Hitchins, 1971: 21, fig. 12. Cinixys belliana: Bulenger, 1905: 252, 1907: 428. Kinixys belliana zuluensis Hewitt, 1931: 471, pi. xxxviii, figs (type lcality: Richards Bay, Zululand); Hewitt, 1935: 350, pi. xxxv, figs 1-2. Kinixys belliana belliana: Lveridge and Williams, 1957: 384 (part); Wermuth and Mertens, 1961: 180, 1977: 70 (part); Pley, 1965: 54, pi. 11; Brutn and Haacke, 1980: 277; Bradley, 1981: 208, figs 9-10; Byctt and Burquin, 1988: 121, pi. 41; Lambiris, 1988: 32; Branch, 1988: 3D; Ernst and Barbur, 1989: 229. Kinixys belliana zmbensis: Bradley, 1989: 50. DIAGNOSIS. Beak unicuspid; carapacial hinge well develped; gulars (tgether) usually less than twice as wide as lng; drsal epiplastrn length % f plastrn midline length; marginals usually 23, supracaudal undivided; carapace cnvex, shell height usually included in length less than 2,3 times; plastrn cncave in males; clur pattern f carapacial shields radial. DESCRIPTION. Beak unicuspid; prefrntal entire (but usually cleft anterirly) r lngitudinally divided; frntal large, rarely lngitudinally divided; frelimb cvered with mre r less imbricate scales, many enlarged and pinted, usually five claws n freft (fur in TM and n ne side f TM and 55453; six n ne side f TM 29400). Carapace cnvex (shell length/height rati 1,82-2,34), highest pint usually n vertebral 3 r 4; anterir margin ften smewhat expanded, nt reverted; psterir marginals nt reverted r serrated; drsal shields with well-marked grwth annuli and deep sulci; nuchal elngate in adults; vertebrals 5, pleurals 4 (five n left side f hltype f K. b. zuluensis;) marginals usually 23. Frnt lbe f plastrn truncate anterirly, usually prjecting beynd anterir brder f carapace, especially in males; gulars paired, usually less than twice as brad as lng; pectrals with a mderate median sulcus; axillaries 2-3; inguinal large, usually in cntact with sixth marginal, in brad cntact with femral; hind lbe shrt and truncate with a very shallw psterir ntch. Plastrn distinctly cncave in adult males. COLOURATION. Carapace yellw, with a bld black radial pattern (4-6 rays n each vertebral and pleural shield). In females the black rays may be almst cnfluent as in TM (Hewitt, 1935; pi. xxxv, fig. 1), but at the ther extreme is TM (Hewitt, 1931, pi. xxxviii, fig. 22), with nly a few narrw black rays n the vertebrals and pleurals. In males the black pattern is usually less develped and may be bscure in ld individuals. The pi astral pattern is very variable. In juveniles the plastrn is largely black, except fr a brad median band f yellw extending alng the median sulci f the pectrals, abdminals and femrals. Thereafter the black is brken up by

8 48 ANNALS OF THE TRANSVAAL MUSEUM yellw intrusins until large adults retain nly vestigial black markings, usually in a fragmented radial pattern. SIZE. Largest male (TM 51378, Candver t Mkuze, Zululand) 211 mm lng, 137 mm wide, 90 mm high; largest female (TM 53989, Nkwaleni Valley, Zululand) 210 mm lng, 136 mm wide, 94 mm high. HABITAT. In the suthern part f its range K. b. bel/iana appears t be restricted t the castal plain (Castal Frest and Thrnveld), where it is cmmn. Haacke (in litt.) has a sight recrd f specimens in castal dune frest at Mapelane, suth f Lake St Lucia, and a midden f 34 carapaces was fund in castal dune frest near Mbibi (Brutn and Haacke, 1980). TM was fund n a rad thrugh savanna thrnveld west f the Lebmb Range. Occasinally encuntered n the marine beach (Brutn and Haacke, 1980). DISTRIBUTION. I have earlier (Bradley, 1989) assigned the eastern ppulatins f K. bel/iana t K. b. zmbensis Hewitt, 1931, with a range extending frm castal Kenya suth thrugh Tanzania, Mzambique, suthern Malawi and eastern Zimbabwe t Zululand. Hwever, examinatin f mre material makes it difficult t distinguish any subspecies except K. b. ngueyi (Lataste, 1886) f West Africa. The prprtins f the type specimen f K. bel/iana suggest that it riginated frm Ethipia and it just falls within the range f variatin fr K. b. mertensi Laurent 1956, (Bradley, 1989). The nrthern ppulatins f K. b. bel/iana range frm nrthwestern Smalia, thrugh Ethipia, suthern Sudan, western Kenya, Uganda and nrtheastern Zaire t the western Central African Republic and Camern (where there is intergradatin with K. b. ngueyi); the typical frm then extends suth thrugh western Zaire t Angla, where the presence f K. spekii has nt yet been cnfirmed (the Anglan recrds in Bradley, 1989: fig. 126, were speculative and based n literature recrds). LOCALITIES. SOUTH AFRICA. Zululand: n precise lcality AM -(2), NMZB ; Candver t Mkuze TM 51378; False Bay TM 63908; Futululu Frest Statin TM 65703; Ksi Bay (Rwe-Rwe, Murray and Daniel, 1968); Hluhluwe Game Reserve (Burquin et al., 1971); 20 km NNW f Hluhluwe TM ; 5 km N f Hluhluwe TM 55453; East Shre f Lake St Lucia TM ; Lake Sibaya Research Statin TM 48265; Lala Nek TM 47844; Manaba TM 14877; Manzengwenya TM ; Missin Rcks, Lake St Lucia TM 54562; Mkuze Game Reserve (Pley, 1965); Mtubatuba TM 13544, , 53940; Mtubatuba t Hluhluwe TM 57947; Ndumu Game Reserve TM ; l\jdumu Hill (Lambiris, 1988: 32); Nkwaleni Valley TM ; Othbthini TM 19363; Richards Bay NM 1203 (hltype f K. b. zuluensis;) St Lucia Bay AM - (2); Sdwana Bay TM 64227; Sdwana Bay t Umhlsinga NMZB 6044; Umflzi Game Reserve (Burquin et al., 1971); Umflzi Statin BM and 68. MOZAMBIQUE. Chimnz TM ; Maput NHMG 65311; Maput Elephant Reserve TM 41762; Zavra NMZB-UM Kinixys /batsiana Pwer, Figs 6, 7 Cinixys lbatsiana Pwer, 1927, Transactins f the Ryal SOCiety f Suth Africa 14: 410, pi. xix-xx (type lcality: Lbatse, Btswana) and 1931: 46, 49, figs 1-2. Kinixys lbatsiana: Hewitt, 1931: 488, pi. xxxvii, figs 12-13, text fig. 1e--f, 2; Bradley, 1989: 54, fig. 12b. Kinixys belliana belliana: (part) Lveridge and Williams, 1957: 384; Wermuth and Mertens, 1961: 180, 1977: 70; Jacbsen, 1977: 15. Kinixys belliana spekii(nt Gray): Auerbach, 1985: 17; Byctland Burquin, 1988: 124 (part), pi. 42; Branch, 1988: 30 (part). Kinixys belliana (part): Auerbach, 1987: 69, pi. 7/1. DIAGNOSIS. Beak unicuspid; carapacial hinge well develped; gulars (tgether) usually less than twice as wide as lng; drsal epiplastrn length % f plastrn midline length; marginals usually 23, supracaudal undivided, carapace mderately cnvex t distinctly depressed; plastrn cncave in males; clur pattern f carapacial shields radial in sme females, but ften fragmented, while males are usually unifrm. DESCRIPTION. Beak unicuspid; prefrntal lngitudinally divided; frntal large, entire (smetimes transversely divided: Hewitt, 1931, fig. 2); frelimb cvered with subimbricate scales, sme enlarged and pinted; five claws n frnt ft (fur n ne side f TM 4821). Carapace variable in depth (shell length/height rati 2,08--2,59 in 33 males: 1,95-2,51 in 32 females), usually level between the centres f vertebrals 1 and 4; drsal shields relatively flat; anterir margin slightly expanded, psterir marginals strngly reverted and serrated, the supracaudal widened mesially, with a lngitudinal grve belw; cervical elngate in adults; vertebrals usually 5 (six in TM 4805,34620,47207,69908 and 69914; seven in TM 69963); pleurals 4 (tw/three in TM 12760; five in TM 4805, 4826, and 69908); marginals usually 23 (rarely 19, 21,22,24 r 25).

9 BROADLEY: SOUTHERN AFRICAN KINIXYS 49 Frnt lbe f plastrn truncate anterirly, nt usually extending beynd the anterir brder f carapace; gulars paired, usually less than twice as brad as lng; pectrals with a shrt median sulcus; axillaries 2-3, f mderate size; inguinal mderate t large, usually in cntact with sixth marginal and femral; hind lbe shrt and truncate, with a shallw psterir ntch. COLOURATION. Subadults and females usually have a fragmented radiate pattern n each carapacial shield, which is red-brwn mesially. Males are usually devid f markings. The plastral shields have a sparse pattern f radiating dark streaks, which usually disappear in adult males. Transvaal trtises have mre distinct markings than thse frm sutheastern Btswana. SIZE. Largest male (TM 52259, Mamiaanshek, Waterberg, Transvaal) 172 mm lng, 119 mm wide, 74 mm high; largest female (TM 52258, Mamiaanshek) 200 mm lng, 135 mm wide, 86 mm high. HABITAT. This species inhabits the Transvaal Bushveld and Thrnveld (= Middleveld), extending int sutheastern Btswana. In the Nylsvley Nature Reserve this trtise was rarely seen and then mainly in the Burkea savanna (Jacbsen, 1977). The type series was taken near Lbatse in the klfs amng the hills (Pwer, 1931) and in the Transvaal they are mstly fund n rcky hillsides (Jacbsen, 1989). ".,.. 30" \ /' ".,.. 30" : Ii, ntlensis lbatsin ".,..... ::TIlllllllj speki i b. bellin BREEDING. A female frm the farm Rietfntein 214JR laid six eggs n 13 April and three f these hatched after 313 days (Jacbsen, 1989). DIET. One f the types was eating a mushrm when fund (Pwer, 1931). A specimen frm the Dasprt Range defaecated the fllwing: largely vegetable matter (leaves f a herb, seeds f a legume Elaphantrrhiza; tw species f tenebrinid beetles, a helicprid beetle and a millipede (Jacbsen, 1989). Specimens in captivity in Bulaway feed avidly n snails (Achatina and Helix) and millipedes, as d K. spekii and K. b. beliiana. DISTRIBUTION. The Transvaal bushveld, extending int sutheastern Btswana. Tw specimens frm utside this area require explanatin. An adult female frm Rchdale Farm, just nrth f the Sutpansberg, is prbably a waif that was swept thrugh the Waterprt by the Sand River in fld: it is far nrth f the nearest K. /batsiana lcalities. Fig. 4 Distributin f Suthern African species f Kinixys; tp: K. nata/ensisand K./batsiana" bttm: K. spekiiand K. b. belliana. Open symbls dente literature recrds; stars dente sympatry f species. The shell f an adult male was picked up in the Manyeleti Game Reserve in the eastern Transvaal lwveld (which has als Yielded nine K. spekii and ne K. natalensis) this specimen was almst certainly translcated frm the bushveld and released by a misguided turist. LOCALITIES. BOTSWANA. Gabrne (Auerbach, 1987); Lbatse AM -, BM ; MMK ,216-21,223-4, SAM 43464; Otse (Auerbach, 1987, pi. 7/1)NMZB SOUTH AFRICA. Transvaal: n precise lcality TM 12760, 67799, 69963; Bekenhutsklf Drift 286JR TM 64286; Buffelshek 351 KO TM 69904; Buffelsprt 421 KR TM 69906; Bulge River 198KO TM 69961; Dasprt Range TM 62795; Die Brn TM 50680;

10 50 ANNALS OF THE TRANSVAAL MUSEUM Gedehp 749KS TM 69913; Hennps River TM 28663; Klipheuwel 573KS TM 69945; Klppersdam 187JR TM 63774; Krmdraai I\JMZB 9877, TM 49863; Leeuwfntein 188clR TM 64362; Lskp Dam TM 48704; Lskp Nrd 12JS TM 69909; Ltteringskraal 243clP TM 34620; Malmaniesrivier TM 69983; Mamiaanshek TM ; 'Manyeleti Game Reserve' (translcatin) TM 65797; Marble Hall TM 22974; 58 km N f Marble Hall TM 42750; Miplaats 242JS TM 69914; Naauwprt FMI\JH 17179, TM , 13485, , 13548; Nylsvley Nature Reserve (Jacbsen, 1977); Orighstad area TM 21329; Pretria AM -, TM 62810, 65818; Pretria District NMZB , TM 4805, 34666, 34686, 47207; Rietintein 214JR TM 69962; Rietspruit TM 4821, 4826; Rchdale 700MS TM 36366; Schliderkrans 1041 LS TM 69908; Steynsdrift 145JS TM 69944; Syferfntein 178clP TM 69902; Tivli 98KP TM 69905; Tweefntein 523JQ NMZB 11076; Vlakplaats 354clR TM 69911; Vlakplaats 535KS TM 69946; Vlakplaats 723KS TM 69912; Waterval 297JR TM 69910; Witintein 521JR TM 69947; Zandspruit 189JR TM 69907; Znkll 473JR TM Fig. 5 Drsal lateral and ventral views f (left) Kinixys natalensis (AM -, lecttype female frm Dimane Slream, near Jamesn's Drift, Tugela Valley, Natal) and (right) K. spekii (AM 78C, lecttype female f K. australis Hewitt frm White River, Transvaal). Thick lines indicate sulci, thin lines indicate sutures between carapacial bnes; f = fntanelle. RELATIONSHIPS Until recently the relatinships f Kin/iys t ther trtise genera were bscure (Lveridge and Williams, 1957; Crumly, 1984), but the recent descriptin f the large trtise Impregnchelys pachytectis frm the Micene depsits f Rusinga Island in Kenya (Meylan and Auffenberg, 1985) has revealed a sister genus. Gaffney and Meylan (1988), in their phylgeny f turtles, have included these tw genera in an unnamed taxn that is mre derived than the tribe Megalchelyini (including Aldabrachelys, Megalchelys, Asterchelys, Chelnidis and Gechelne,) but less derived than Indtestud and the tribe Testudinini (including Testud, Malacchersus, Psammbates, Hmpus, Pyxis, Acinixys and Chersina.). Fig. 6 Drsal, lateral and ventral views f (left) Kinixys b. belliana (TM 34680, female frm Ndumu Game ReseNe, Zululand) and (right) K. lbatsiana (AM -, male frm Lbatse, Btswana). Cnventins as in Fig. 5.

11

12 52 ANNALS OF THE TRANSVAAL MUSEUM REFERENCES AUERBACH, R. D., The reptiles f Gabrne. Btswana Bk Centre, Gabrne. AUERBACH, R. D., The amphibians and reptiles f Btswana. Mkwepa Cnsultants, Gabrne. BOUlENGER, G. A., On a cllectin f batrachians and reptiles made in Suth Africa by Mr C. H. B. Grant, and presented t the British Museum by Mr C. D. RUdd. Prceedings f the Zlgical Sciety f Lndn 2: BOU lenger, G. A., Secnd reprt n the batrachians and reptiles cllected in Suth Africa by Mr C. H. B. Grant and presented t the British Museum by Mr C. D. Rudd. Prceedings f the Zlgical Sciety f Lndn 2: BOURQUIN, 0., VINCENT, J. and HITCHINS, P. M., The vertebrates f the Hluhluwe Game Reserve - Crridr (Stateland) - Umflzi Game Reserve Cmplex. LammergeyerN. 14: BOYCOTT, R. C., Kinixys natalensis: species accunt. In: BRANCH, W. R., ed., Suth African red data bk - reptiles and amphibians, pp Suth African Natinal Scientific Prgrammes Reprt N BOYCOTT, R. C. and BOURQUIN, 0., The Suth African trtise bk. Suthem Bk Publishers, Jhannesburg. BOYCOTT, R. C. and JACOBSEN, N. H. G., On the distributin, habitat and identificatin f Kinixys natalensis Hewitt, 1935 (Cryptdira: Testudinidae) in suthern Africa. Durban Museum Nvitates 14(5): BRANCH, W. R., Field guide t the snakes and ther reptiles f Suthem Africa. Struik, Cape Twn. BROADLEY, D. G., A review f the ppulatins f Kinixys (Testudinidae) ccurring in suth-eastern Africa. Annals f the Cape Prvincial Museums (Natural Histry) 13(14): BROADLEY, D. G., Species accunts fr Kinixys bel/iana, 'K. spekif, 'K. lbatsiana" and K. natalensis, pp , In: SWINGlAND, I. R. and KLEMENS, M. W., eds, The cnservatin bilgy f trtises. Occasinal Papers I.U.C.N. Species Survival Cmmissin NO.5. BRUTON, M. N. and HAACKE, W. D., The reptiles f Maputaland. In: BRUTON, M. N. and COOPER, K. H., eds, Studies n the eclgy f Maputaland, pp Rhdes University and Natal Branch, Wildlife Sciety f Suthern Africa, Durban. CRUMLY, C. R., The evlutin f land trtises (Family Testudinidae). Ph.D. thesis, Rutgers University, Newark, New Jersey. CRUMLY, C. R., Species accunt: Kinixys natalensis, In: KING, F. W. and BURKE, R. L., eds, Crcdifian, tuatara, and turtle species f the wrld, p. 91. Assciatin f Systematics Cllectins, Washingtn. ERNST, C. H. and BARBOUR, R. W., Turtles fthe wrld. Smithsnian Institutin Press, Washingtn. GAFFNEY, E. S. and MEYLAN, P. A., A phylgeny f turtles. In: BENTON, M. J., ed., The phylgeny and classificatin f the tetrapds, Vlume 1: Amphibians, reptiles, birds, pp The Systematics Assciatin, Oxfrd. HEWITT, J., Descriptins f sme African trtises. Annals f the Natal Museum 6(3): HEWITT, J., Sme new frms f batrachians and reptiles frm Suth Africa. Recrds f the Albany Museum 4: JACOBSEN, N. H. G., An anntated checklist f the amphibians, reptiles and mammals f the Nylsvley Nature Reserve. Suth African Natinal Scientific Prgrammes Reprt N. 21. JACOBSEN, N. H. G., A herpetlgical survey f the Transvaal. Ph.D. thesis: University f Natal (Durban). LAMBIRIS, A. J. L., Systematic list: Genus Kinixys, p. 32. Annual Reprt n the Lambiris Herptlgical Cl/ectin 8(1): LAURENT, R. F., Cntributin 11 I'Herpetlgie de la Regin des Grands lacs de l'afrique centrale. Annales du Muses ryal du Cng Beige, Serie in 8v, Sciences Zlgiques 48: laurent, R. F., On the races f Kinixys beffiana Gray. Brevira 176: 1-6. loveridge, A. and WilLIAMS, E. E., Revisin f the African trtises and turtles f the subrder Cryptdira. Bulletin f the Museum f Cmparative Zlgy (Harvard) 115(6): MEYLAN, P. A. and AUFFENBERG, W., New land trtises (Testudines: Testudinidae) frm the Micene f Africa. Zlgical Jumal f the Linnean SOCiety 86: PIENAAR, U. de V., The reptiles f the Kruger Natinal Park. Natinal Parks Bard, Pretria. PIENAAR, U. de V., The reptile fauna f the Kruger Natinal Park. Natinal Parks Bard, Pretria. PIENAAR, U. de V., HAACKE, W. D. and JACOBSEN, N. H. G., The reptiles f the Kruger Natinal Park. Natinal Parks Bard f Suth Africa, Pretria. POOLEY, A. C., A preliminary checklist f the reptiles fund within the Mkuzi and Ndumu Game Reserves in nrthem Zululand. Lammergeyer 3(2): POWER, J. H., On the herpetlgical fauna f the lbatsi Linkana area. Part I - lbatsi. Transactins f the Ryal Sciety f Suth Africa 14(4): POWER, J. H., On the herpetlgical fauna f the lbatsi Linkana area. Part II - Linkana. Transactins f the Ryal Sciety f Suth Africa 20(1): ROWE-ROWE, D. T., MURRAY, R. M. and DANIEL, P. J., Miscellaneus ntes f natural histry interest. 29. Trtise diet. LammergeyerN. 9: 53. SWITAK, K. H.,1971.ln Kruger Natinal Park. IntematinalTurtle and Trtise Sciety Jurnal 5(1): 7-9, 27. WERMUTH, H. and MERTENS, R., Schildkrten, Krkdile, Briickenechsen. G. Fischer, Jena. WERMUTH, H. and MERTENS, R., Liste der rezenten Amphibien und Reptilien. Testudines, Crcdylia, Rhynchcephalia. Tierreich 100: ZANGERl, R., The turtle shell. In: GANS, C., BElLAIRS, A. D'A. and PARSONS, T. S., eds, Bilgy f the Reptifia 1. Mrphlgy A, pp Academic Press, lndn & New Yrk. Pstal address: D. G. Bradley Natural Histry Museum P. O. Bx 240 Bulaway Zimbabwe