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Complete BEAST timetree. Bars at nodes indicate 95% HPD

A. namaquensis stat. nov. (FitzSimons, 1938) Distribution. Known from scattered localities in the Succulent Karoo Biome in the Little Namaqualand region of the Northern Cape Province (Bauer 2014 a) (Fig. 4). Remarks. Haacke (1965), in describing A. africana tirasensis considered A. namaquensis as a subspecies of A. africana, and it has maintained this rank since (Mertens 1971; Branch 1981, 1988, 1998; Onderstall 1984; Bauer 2014 a), although chiefly because this poorly known taxon has not been reviewed subsequently. Under modern species concepts, the differences between the supposed subspecies of A. africana— including precloacal pore counts, presence of internasal granules, gular scale counts, and color pattern (Haacke 1965) would generally be accepted as evidence of specific distinctness, especially in light of the large disjunctions between the forms and their likely low vagility (Mouton & Mostert 1985; Jacobsen 1997). On this basis we here recognize this taxon at the rank of full species.

A. hawequensis Mouton & Mostert, 1985 (Fig. 3 A) Distribution. Restricted to the fynbos habitats in Hawequa Mountains of the southwestern Western Cape Province, South Africa (Bauer 2014 c) (Fig. 4). Remarks. This species is the sister to all remaining congeners. It was previously considered a member of the A. africana group (Mouton & Mostert 1985). The species was previously listed in the South African Red Data Book as “Restricted” (Mouton 1988) and is currently considered as Near Threatened under IUCN assessment criteria (Bauer 2014 c).

A. langi (FitzSimons, 1930) (Fig. 5 C) Distribution. Lowveld savannah in northeastern Mpumalanga and northeastern Mpumalanga provinces and adjacent Mozambique (Visser 1984; Jacobsen 1990; Bauer 2014 e) (Figs. 4, 6). Remarks. This species was previously considered a subspecies of A. pondolia (e.g., Onderstall 1984) until raised to specific status by Jacobsen (1990, 1992a). Six of the nine new species described herein are related to A. langi and earlier references to this species outside of its currently recognized range may refer to one or more of these taxa., Published as part of Jacobsen, Niels H. G., Kuhn, Arianna L., Jackman, Todd R. & Bauer, Aaron M., 2014, A phylogenetic analysis of the southern African gecko genus Afroedura Loveridge (Squamata: Gekkonidae), with the description of nine new species from Limpopo and Mpumalanga provinces of South Africa, pp. 451-501 in Zootaxa 3846 (4) on page 483, DOI: 10.11646/zootaxa.3846.4.1, http://zenodo.org/record/250495, {"references":["FitzSimons, V. F. (1930) Descriptions of new South African Reptilia and Batrachia, with distribution records of allied species in the Transvaal Museum collection. Annals of the Transvaal Museum, 14, 20 - 48.","Jacobsen, N. H. G. (1990) A Herpetological Survey of the Transvaal. Unpublished Ph. D. Dissertation, University of Natal, Durban, South Africa, 1621 pp.","Bauer, A. M. (2014 e) Afroedura langi (FitzSimons, 1930). In: Bates, M. F., Branch, W. R., Bauer, A. M., Burger, M., Marais, J., Alexander, G. J. & de Villiers, M. (Eds.), Atlas and Red List of the Reptiles of South Africa, Lesotho and Swaziland. Suricata, 1. South African National Biodiversity Institute, Pretoria, pp. 93 - 94.","Onderstall, D. (1984) Descriptions of two new subspecies of Afroedura pondolia (Hewitt) and a discussion of species groups within the genus (Reptilia: Gekkonidae). Annals of the Transvaal Museum, 33, 497 - 509.","Jacobsen, N. H. G. (1992 a) Flat geckos (genus Afroedura) in the Transvaal. Journal of the Herpetological Association of Africa, 40, 22 - 25. http: // dx. doi. org / 10.1080 / 04416651.1992.9650313"]}

A. loveridgei Broadley, 1963 Distribution. Tete district, Mozambique (Onderstall 1984) (Fig. 4). Remarks. This species was originally described as a subspecies of A. transvaalica, but was elevated to specific status without comment by Branch (1998), although Kluge (2001) continued to recognize it as a subspecies., Published as part of Jacobsen, Niels H. G., Kuhn, Arianna L., Jackman, Todd R. & Bauer, Aaron M., 2014, A phylogenetic analysis of the southern African gecko genus Afroedura Loveridge (Squamata: Gekkonidae), with the description of nine new species from Limpopo and Mpumalanga provinces of South Africa, pp. 451-501 in Zootaxa 3846 (4) on page 467, DOI: 10.11646/zootaxa.3846.4.1, http://zenodo.org/record/250495, {"references":["Broadley, D. G. (1963) Three new lizards from South Nyasaland and Tete. Annals and Magazine of Natural History, series 13, 6, 285 - 288. http: // dx. doi. org / 10.1080 / 00222936308651355","Onderstall, D. (1984) Descriptions of two new subspecies of Afroedura pondolia (Hewitt) and a discussion of species groups within the genus (Reptilia: Gekkonidae). Annals of the Transvaal Museum, 33, 497 - 509.","Kluge, A. G. (2001) Gekkotan lizard taxonomy. Hamadryad, 26, 1 - 209."]}

A. nivaria (Boulenger, 1895 [1894]) Distribution. Drakensberg of the Eastern Cape and Free State provinces, South Africa. Its occurrence in Lesotho is suspected but not yet documented (Bates & Bauer 2014 b) (Fig. 4). Remarks. Loveridge (1947) synonymized A. amatolica with A. nivaria and this interpretation was long followed (e.g, Wermuth 1965; De Waal 1978; Branch 1981). The two are not sister taxa (Fig. 1) and A. amatolica was raised from synonymy by Visser (1984 b). Afroedura nivaria as currently construed is non-monophyletic (Makhubo et al. 2011; Makhubo 2013) and is currently under revision (Bates & Branch 1998)., Published as part of Jacobsen, Niels H. G., Kuhn, Arianna L., Jackman, Todd R. & Bauer, Aaron M., 2014, A phylogenetic analysis of the southern African gecko genus Afroedura Loveridge (Squamata: Gekkonidae), with the description of nine new species from Limpopo and Mpumalanga provinces of South Africa, pp. 451-501 in Zootaxa 3846 (4) on page 471, DOI: 10.11646/zootaxa.3846.4.1, http://zenodo.org/record/250495, {"references":["Boulenger, G. A. (1895 [1894]) On a gecko from South Africa. Proceedings of the Zoological Society of London, 1894, 608 - 609.","Loveridge, A. (1947) Revision of the African lizards of the Family Gekkonidae. Bulletin of the Museum of Comparative Zoology, 98, 1 - 469, pls. 1 - 7.","Wermuth, H. (1965) Liste der rezenten Amphibien und Reptilien, Gekkonidae, Pygopodidae, Xantusiidae. Das Tierreich, 80, xxii + 246 pp.","De Waal, S. W. P. (1978) The Squamata (Reptilia) of the Orange Free State, South Africa. Memoirs van die Nasionale Museum, 11, iii + 160 pp.","Branch, W. R. (1981) An annotated checklist of the lizards of the Cape Province, South Africa. Annals of the Cape Provincial Museums (Natural History), 13, 141 - 167.","Visser, J. 1984 b. Diksterte van die Amatolaberge. Akkedisse van Suider - Afrika 15. Landbouweekblad, 29 June 1984, 48 - 49, 51, 53.","Makhubo, B. G., Tolley, K. A. & Bates, M. F. (2011) Phylogenetic analysis of the Afroedura nivaria species complex in South Africa. 10 th Conference of the Herpetological Association of Africa, Cape Town, South Africa, 12 - 14 January 2011, Program and Abstracts, 61.","Makhubo, B. G. (2013) Phylogenetic and Morphological Analysis of the Afroedura nivaria (Reptilia: Gekkonidae) Species Complex in South Africa. Unpublished M. Sc. Thesis, University of Stellenbosch, xiii + 106 pp.","Bates, M. F. & Branch, W. R. (1998) Taxonomy and distribution of the Afroedura nivaria and A. karroica complexes (Sauria: Gekkonidae). Herpetological Association of Africa, Fifth Symposium, University of Stellenbosch, 14 - 16 September 1998, Programme and Abstracts. [2 unnumbered pages]"]}

A. karroica (Hewitt, 1925) (Fig. 3 D) Distribution. Karoo of the central-western Eastern Cape and adjacent areas of the Northern and Western Cape provinces (Bauer 2014 d) (Fig. 4). Remarks. Both A. bogerti and A. halli were previously treated as subspecies of A. karroica (Loveridge 1944, 1947; Wermuth 1965; Kluge 2001)., Published as part of Jacobsen, Niels H. G., Kuhn, Arianna L., Jackman, Todd R. & Bauer, Aaron M., 2014, A phylogenetic analysis of the southern African gecko genus Afroedura Loveridge (Squamata: Gekkonidae), with the description of nine new species from Limpopo and Mpumalanga provinces of South Africa, pp. 451-501 in Zootaxa 3846 (4) on page 470, DOI: 10.11646/zootaxa.3846.4.1, http://zenodo.org/record/250495, {"references":["Hewitt, J. (1925) On some new species of reptiles and amphibians from South Africa. Records of the Albany Museum, 3, 343 - 368, pls. xv - xix.","Loveridge, A. (1944) New geckos of the genera Afroedura, new genus, and Pachydactylus from Angola. American Museum Novitates, 1254, 1 - 4.","Loveridge, A. (1947) Revision of the African lizards of the Family Gekkonidae. Bulletin of the Museum of Comparative Zoology, 98, 1 - 469, pls. 1 - 7.","Wermuth, H. (1965) Liste der rezenten Amphibien und Reptilien, Gekkonidae, Pygopodidae, Xantusiidae. Das Tierreich, 80, xxii + 246 pp.","Kluge, A. G. (2001) Gekkotan lizard taxonomy. Hamadryad, 26, 1 - 209."]}

Afroedura pienaari sp. nov. (Figs. 11 B���C) Afroedura pondolia subsp. nov. Pienaar et al. 1983 Afroedura langi 'Waterpoort' Jacobsen 1992 a, 1997; Kirchhof et al. 2010 Afroedura langi 'Tshipise' Jacobsen 1992 a, 1997 Afroedura langi 'Shinokwen' Jacobsen 1992 a, 1997 Holotype. TM 81144, adult male, Farm Bristol 760 MS, 22 �� 59 ' S, 29 �� 37 ' E, Soutpansberg District (2229 DC), Limpopo Province, Republic of South Africa, collector R. E. Newbery, 25 July 1981. Paratypes. TM 81133 ���81139, 81141���81143, 81145���81146, 81160��� 81165, same data as holotype; TM 81140, same locality as holotype, collector N. H. G. Jacobsen, 2 February 1979; TM 81175, Farm Crimea 747 MS, Soutpansberg District, Limpopo Province (2229 DC), collector N. H. G. Jacobsen, 1 February 1979; TM 81147���81153, Farm Robertson 748 MS, Soutpansberg District, Limpopo Province (2229 DC), collector R. E. Newbery, 24 July 1981; TM 81167���81169, Farm Waterpoort 695 MS, Soutpansberg District, Limpopo Province (2229 DC), collector R. E. Newbery, 24 July 1981; TM 81170���81173, Farm The Moss 763 MS, Soutpansberg District, Limpopo Province (2229 DC), collector R. E. Newbery, 7 November 1985. Additional material examined (all from Limpopo Province). TM 81154 ���81157, 81174, Farm Zoutpan 459 MS, Soutpansberg District (2229 CD); TM 81107, Musekwas location 194 MT, Dzanani District (2230 CC); TM 81103 ���81104, 81106, 81111 Gumela, Sibasa District (2230 CB) TM 81102, 81105, 81110, Lavhalisa, Dzanani District (2230 CA); TM 30414, 35 km southwest of Tshipise, Soutpansberg District (2229 DD); TM 81342, Vhuswinzhe, Dzanani District (2230 CC); TM 81101, 81108��� 81109, near Shinokwenfontein, KNP (2231 AC); TM 77353���77355, Bobomeni, KNP (2231 BC); TM 77356���77360, Mashipange, KNP (2230 DB); TM 77336 ���77345, 3 km W of Baobab Hill, KNP (2231 AC); TM 77329���77335 Pafuri border gate, KNP (2231 AC); TM 77346 ���77352, 5 km NW of Hutwini, KNP (2231 AC); TM 77361 ���77362, 77363��� 77367, Madziringwe stream, S of the Trails Camp, KNP (2231 CA); TM 77368���77372 Nyala Drive, 4 km W of Bobomeni, KNP (2231 AC); TM 77309 ��� 77304, Shipudza Spring, KNP (2231 CA); TM 77315, 77318��� 77323, Shidzivani, KNP (2231 CA); TM 77324���77328, Xantangalani, KNP (2231 CA); TM 47314 Vivo area, Soutpansberg District (2329 AB); TM 81129���81132 Farm Calitzdorp 221 LS, Soutpansberg District (2329 BA). Etymology. The species is named for the late Dr. U. de V. Pienaar (1930���2011), former Chief Warden of Kruger National Park and former Chief Director of South African National Parks, in recognition of his work in furthering herpetofaunal knowledge of the Kruger National Park. Diagnosis. A medium-sized Afroedura (maximum SVL 53 mm) differing from all other congeners by the following combination of characters: two pairs of enlarged subdigital lamellae per digit; tail faintly verticillate and flattened near base, with four subcaudal rows and 6���7 supracaudal rows per verticil; dorsal scales smooth, 86���111 scale rows at midbody; internasal scales absent; 13���19 precloacal pores in males. Description. (based on holotype TM 81144) Adult male; SVL 48.0 mm; TailL 57.0 mm; mass before preservation 2.2 g. A medium sized, dorsoventrally depressed Afroedura, with a flat, ovate head, wider than the neck. Rostral 2.5 times wider than high; nostrils pierced between rostral, first upper labial and three nasal scales; nasorostrals raised and in broad contact behind rostral. Scales on snout variable, large and flattened, decreasing in size posteriorly, to being smallest on the crown of the head; 10 scales between nasals and eye and 16 between eye and ear; four supraciliary spines above eye; supralabials nine. Mental wedge-shaped, longer than wide; one postmental. Infralabials seven. Dorsal scales uniform, rounded, juxtaposed and flattened paravertebrally, becoming larger and oblique laterally. Midbody scales in 93 rows. Ventral scales flat, smooth, imbricate and irregular to hexagonal. Limbs robust; two pairs of enlarged scansors under each digit and four inferomedian scales under 4 th toe. Precloacal pores 18, arranged in a shallow ���V���-shape, with a single median poreless scale interrupting the series. Tail faintly verticllate, widening slightly posterior to the cloaca before tapering. Caudal verticils with seven dorsal and four ventral scale rows. Supracaudals almost square to slightly longer than broad, with a rounded to apical posterior margin; subcaudals almost square with a rounded posterior margin and imbricate; three post cloacal spurs at base of tail. Color. Gray-brown to brown dorsally with 6���7 irregular dark brown to blackish crossbars occasionally bordered posteriorly by a black edge, followed by a white marking, the latter sometimes evident only as a median spot. Dorsum variegated and mottled with brownish black and off-white. Head mottled dorsally; a dark stripe extending from the nostrils through the eye to the rear of the head, linking to the second crossbar in the neck region. Limbs brownish with pale and dark spotting and marbling. Original tail with 8���13 crossbars, with interstitial areas spotted or marbled. Venter whitish to pinkish mesially. Variation. There is significant variation in specimens of this species in several features (Tables 4, 5), in part reflecting the very large sample size and its relatively large geographic range. A single internasal is present in TM 81142. 10 ��� 14 scales between nasals and eye and 16���22 between eye and ear, 2���4 supraciliary spines. Supralabials 8���12. One or two postmental scales present. Infralabials 7���9. Midbody scale rows 86���98 in type series, up to 111 in other specimens examined. 3���7 enlarged inferomedian scales under 4 th toe. Precloacal pores in male paratypes in continuous or interrupted series of 14���18 (12 to 19 in other specimens examined, although 96 % of all male specimens have at least 14 pores); no precloacal pores in females. 0���4 postcloacal spurs on either side of tail base. Original tails 50.0��� 58.4 % of total length. 32.2 % of specimens have regenerated tails (n= 31). Distribution. Recorded from along the northern Soutpansberg and into the northwestern Kruger National Park, Limpopo Province (Fig. 6). Kirchhof et al. (2010) reported this species from Lajuma, on the southwestern Soutpansberg, in an area that would be expected to harbour A. broadleyi, however, their specimens had the typical precloacal pore counts of A. pienaari and on this basis we accept their identification. Natural history. Afroedura pienaari sp. nov. is a nocturnal, rupicolous gecko that inhabits crevices and fissures in and between boulders on rocky outcrops and under flakes of exfoliating rock beneath overhangs. It appears to be mostly solitary, but is occasionally found in pairs and rarely three indivduals may occupy the same crevice. Found on north and south-facing slopes in Makhuleke Sandy Bushveld (SVl 1), Limpopo Ridge Bushveld (SVmp 2), Soutpansberg Mountain Bushveld (SVcb 21) and Soutpansberg Summit Sourveld (GM 28) (Mucina & Rutherford 2006) at elevations of 800���1200 m a.s.l. Kirchhof et al. (2010) reported this species from cracks and fissures in vertical rock walls and from caves and found several individuals in an electrical junction box. Two soft shelled eggs are laid at a time. These are placed on the undersides of rocks, to which they adhere and then harden. Communal nesting takes place in favorable crevices. Remarks. Jacobsen (1990, 1992a) initially considered each of three discrete populations of this gecko to represent putative species. The western and eastern forms are similar in size but are separated by smaller geckos, with generally lower scale counts, in the central part of the range (Table 5). Genetic divergences between populations of Afroedura pienaari sp. nov. are also pronounced (Fig. 1). However, further sampling may reveal continuity between populations and existing populational differences in morphological features are largely overlapping, so we here treat all of the northern Soutpansberg/Limpopo Valley A. langi complex geckos as a single species. Waterpoort Tshipise KNP Midbody scales n = 31 n = 9 n = 66 86���107 86���102 86���111 93.9 + 4.71 90.67 + 4.8 96.44 + 5.29 Eye���Ear scales n = 33 n = 9 n = 62 13���19 15���17 13���19 16.58 + 1.15 15.78 + 0.67 16.05 + 1.45 Supralabials n = 38 n = 10 n = 59 8���12 8���10 8���11 9.6 + 0.8 9.4 + 0.7 9.6 + 0.81 Infralabials n = 42 n = 10 n = 62 7���10 7���9 6���10 8.21 + 0.68 7.7 + 0.67 8.02 + 0.78 Caudal verticils (supracaudals/subcaudals) 7 / 4���5 6���8 / 4 6���7 / 4 Precloacal Pores n = 16 n = 7 n = 27 14���19 12���15 14���19 16.94 + 1.53 14.14 + 1.21 16.89 + 1.40 Adult male SVL (mm) n = 16 n = 5 n = 27 51.0 42.5 48.0 44.56 + 3.08 39.60 + 2.16 43.96 + 2.61 Adult female SVL (mm) n = 16 n = 2 n = 31 52.0 39.0 53.0 47.12 + 3.72 34.5 + 6.36 44.47 + 4.76 This species may be distinguished from the geographically proximate A. broadleyi sp. nov. and from A. waterbergensis sp. nov. by its greater number of precloacal pores and from A. leoloensis sp. nov. by its lower number of pores. In addition, it is substantially larger than A. leoloensis sp. nov., A. langi, and A. waterbergensis sp. nov. (see Table 4) and has a significantly greater number of supralabials than A. langi (P A. broadleyi sp. nov. and A. langi (p A. granitica sp. nov., which is larger (58 versus 53 mm maximum SVL and even greater mean differences; Table 4). The great majority of male specimens, however, have 14 or more precloacal pores (up to 19), averaging more than 16, whereas A. granitica sp. nov. has a mean of only 14 pores. Afroedura pienaari sp. nov. also has fewer eye to ear scales than A. granitica sp. nov., although not statistically so., Published as part of Jacobsen, Niels H. G., Kuhn, Arianna L., Jackman, Todd R. & Bauer, Aaron M., 2014, A phylogenetic analysis of the southern African gecko genus Afroedura Loveridge (Squamata: Gekkonidae), with the description of nine new species from Limpopo and Mpumalanga provinces of South Africa, pp. 451-501 in Zootaxa 3846 (4) on pages 490-492, DOI: 10.11646/zootaxa.3846.4.1, http://zenodo.org/record/250495, {"references":["Pienaar, U. de V., Haacke, W. D. & Jacobsen, N. H. G. (1983) The Reptiles of the Kruger National Park. National Parks Board of South Africa, Pretoria, [viii] + 236 pp.","Jacobsen, N. H. G. (1992 a) Flat geckos (genus Afroedura) in the Transvaal. Journal of the Herpetological Association of Africa, 40, 22 - 25. http: // dx. doi. org / 10.1080 / 04416651.1992.9650313","Jacobsen, N. H. G. (1997) Family Gekkonidae, Genus Afroedura. In: Van Wyk, J. H. (Ed.), Proceedings of the FitzSimons Commemorative Symposium, South African Lizards: 50 years of Progress and Third H. A. A. Symposium on African Herpetology, Transvaal Museum, Pretoria, South Africa, 11 - 15 October 1993. Herpetological Association of Africa, Stellenbosch, pp. 34 - 39.","Kirchhof, S., Kramer, M., Linden, J. & Richter, K. (2010) The reptile species assemblage of the Soutpansberg (Limpopo Province, South Africa) and its characterisitcs. Salamandra, 46, 147 - 166.","Mucina, L. & Rutherford, M. C. (2006) The Vegetation of South Africa, Lesotho and Swaziland. Strelitzia, 19. South African National Biodiversity Institute, Pretoria, 807 pp.","Jacobsen, N. H. G. (1990) A Herpetological Survey of the Transvaal. Unpublished Ph. D. Dissertation, University of Natal, Durban, South Africa, 1621 pp."]}

Afroedura multiporis Group Diagnosis. Two pairs of enlarged scansors per digit, tail not, or only faintly verticillate, rounded to slightly flattened, dorsal scales keeled, one or more internasals between nasorostrals. Remarks. Onderstall (1984) mentioned the presence of keeled scales in members of this group but did not appreciate the phylogenetic value of this character. Jacobsen (1990, 1992a, 1997) identified A. multiporis ���Lebombo��� as another member of this species complex. This putative species remains known from a single juvenile specimen. Although apparently distinctive in several features, the lack of adult material and the imperfect state of the specimen prevent us from describing this form at this time. This form was noted as Afroedura sp. nov. ���Lebombo flat gecko��� by Bourquin (2004)., Published as part of Jacobsen, Niels H. G., Kuhn, Arianna L., Jackman, Todd R. & Bauer, Aaron M., 2014, A phylogenetic analysis of the southern African gecko genus Afroedura Loveridge (Squamata: Gekkonidae), with the description of nine new species from Limpopo and Mpumalanga provinces of South Africa, pp. 451-501 in Zootaxa 3846 (4) on page 473, DOI: 10.11646/zootaxa.3846.4.1, http://zenodo.org/record/250495, {"references":["Onderstall, D. (1984) Descriptions of two new subspecies of Afroedura pondolia (Hewitt) and a discussion of species groups within the genus (Reptilia: Gekkonidae). Annals of the Transvaal Museum, 33, 497 - 509.","Jacobsen, N. H. G. (1990) A Herpetological Survey of the Transvaal. Unpublished Ph. D. Dissertation, University of Natal, Durban, South Africa, 1621 pp.","Jacobsen, N. H. G. (1992 a) Flat geckos (genus Afroedura) in the Transvaal. Journal of the Herpetological Association of Africa, 40, 22 - 25. http: // dx. doi. org / 10.1080 / 04416651.1992.9650313","Jacobsen, N. H. G. (1997) Family Gekkonidae, Genus Afroedura. In: Van Wyk, J. H. (Ed.), Proceedings of the FitzSimons Commemorative Symposium, South African Lizards: 50 years of Progress and Third H. A. A. Symposium on African Herpetology, Transvaal Museum, Pretoria, South Africa, 11 - 15 October 1993. Herpetological Association of Africa, Stellenbosch, pp. 34 - 39.","Bourquin, O. (2004) Reptiles (Reptilia) in KwaZulu - Natal: 1 - Diversity and distribution. Durban Museum Novitates, 29, 57 - 103."]}

A. tembulica (Hewitt, 1926) Distribution. Queenstown region of the Eastern Cape province, South Africa (Bauer 2014 j) (Fig. 4). Remarks. Although considered poorly differentiated from A. amatolica and A. nivaria by Branch et al. (1988), this taxon has never been synonymized with any of its congeners., Published as part of Jacobsen, Niels H. G., Kuhn, Arianna L., Jackman, Todd R. & Bauer, Aaron M., 2014, A phylogenetic analysis of the southern African gecko genus Afroedura Loveridge (Squamata: Gekkonidae), with the description of nine new species from Limpopo and Mpumalanga provinces of South Africa, pp. 451-501 in Zootaxa 3846 (4) on page 471, DOI: 10.11646/zootaxa.3846.4.1, http://zenodo.org/record/250495, {"references":["Hewitt, J. (1926) Some new or little - known reptiles and batrachians from South Africa. Annals of the South African Museum, 20, 473 - 490."]}

Afroedura africana Group Diagnosis. Small to medium sized (SVL Remarks. The monophyly of this group is poorly supported and we recognize this group only tentatively. Additional genetic markers might help to support the union of A. karroica and the arid western taxa, or may reveal alternative placements of these species within the genus., Published as part of Jacobsen, Niels H. G., Kuhn, Arianna L., Jackman, Todd R. & Bauer, Aaron M., 2014, A phylogenetic analysis of the southern African gecko genus Afroedura Loveridge (Squamata: Gekkonidae), with the description of nine new species from Limpopo and Mpumalanga provinces of South Africa, pp. 451-501 in Zootaxa 3846 (4) on page 469, DOI: 10.11646/zootaxa.3846.4.1, http://zenodo.org/record/250495

A. bogerti Loveridge 1944 (Fig. 3 B) Distribution. Namibe Province, Angola and adjacent northern Kaokoveld of northern Namibia (Branch 1998; Griffin 2003) (Fig. 4). Remarks. This distinctive species was originally described by Loveridge (1944) as A. karroica bogerti, and was only elevated to specific status by Onderstall (1984). Afroedura bogerti and A. karroica possess the diagnostic features of different species groups and are separated from one another by a disjunction of approximately 2000 km; that they were previously considered conspecific is illustrative of the fact that herpetologists were reluctant to erect new specific level taxa for geckos of this genus, presumably because of the their general morphological conservatism (Jacobsen 1997). Barts & Haacke (2010) provided data about reproduction in captivity., Published as part of Jacobsen, Niels H. G., Kuhn, Arianna L., Jackman, Todd R. & Bauer, Aaron M., 2014, A phylogenetic analysis of the southern African gecko genus Afroedura Loveridge (Squamata: Gekkonidae), with the description of nine new species from Limpopo and Mpumalanga provinces of South Africa, pp. 451-501 in Zootaxa 3846 (4) on page 467, DOI: 10.11646/zootaxa.3846.4.1, http://zenodo.org/record/250495, {"references":["Loveridge, A. (1944) New geckos of the genera Afroedura, new genus, and Pachydactylus from Angola. American Museum Novitates, 1254, 1 - 4.","Griffin, M. (2003) Annotated Checklist and Provisional National Conservation Status of Namibian Reptiles. Namibia Scientific Society, Windhoek, [ii] + 169 pp.","Onderstall, D. (1984) Descriptions of two new subspecies of Afroedura pondolia (Hewitt) and a discussion of species groups within the genus (Reptilia: Gekkonidae). Annals of the Transvaal Museum, 33, 497 - 509.","Jacobsen, N. H. G. (1997) Family Gekkonidae, Genus Afroedura. In: Van Wyk, J. H. (Ed.), Proceedings of the FitzSimons Commemorative Symposium, South African Lizards: 50 years of Progress and Third H. A. A. Symposium on African Herpetology, Transvaal Museum, Pretoria, South Africa, 11 - 15 October 1993. Herpetological Association of Africa, Stellenbosch, pp. 34 - 39.","Barts, M. & Haacke, W. D. (2010) Afroedura bogerti Loveridge, 1944. Bogert's Flat Gecko. Reproduction. African Herp News, 50, 39."]}

A. tirasensis stat. nov. Haacke, 1965 Distribution. Farm Tiras, Lüderitz District, southern Namibia (Mouton & Mostert 1985; Griffin 2003) (Fig. 4). Remarks. Haacke (1965) originally described this form as a subspecies of A. africana, and it has maintained this rank since (Mertens 1971; Branch 1981, 1988, 1998; Onderstall 1984; Branch et al. 1988; Griffin 2003), although chiefly because this poorly known taxon has not been reviewed subsequently. Under modern species concepts, the differences between the supposed subspecies of A. africana (Haacke 1965) (see above) would generally be accepted as evidence of specific distinctness, especially in light of the large disjunctions between the forms and their likely low vagility (Mouton & Mostert 1985; Jacobsen 1997). On this basis we here recognize this taxon at the rank of full species.

A. major Onderstall, 1984 Distribution. Highveld and middleveld areas of western Swaziland (Boycott 1992, 2014) (Figs. 4, 6). Remarks. This species was originally described as a subspecies of A. pondolia and was elevated to specific rank by Branch (1998) without comment, although Kluge (2001) continued to list it as a subspecies. The species is considered Near Threatened (Monadjem et al. 2003; Boycott 2014)., Published as part of Jacobsen, Niels H. G., Kuhn, Arianna L., Jackman, Todd R. & Bauer, Aaron M., 2014, A phylogenetic analysis of the southern African gecko genus Afroedura Loveridge (Squamata: Gekkonidae), with the description of nine new species from Limpopo and Mpumalanga provinces of South Africa, pp. 451-501 in Zootaxa 3846 (4) on page 474, DOI: 10.11646/zootaxa.3846.4.1, http://zenodo.org/record/250495, {"references":["Onderstall, D. (1984) Descriptions of two new subspecies of Afroedura pondolia (Hewitt) and a discussion of species groups within the genus (Reptilia: Gekkonidae). Annals of the Transvaal Museum, 33, 497 - 509.","Boycott, R. C. (1992) An Annotated Checklist of the Amphibians and Reptiles of Swaziland. The Conservation Trust of Swaziland, Mbabane, 27 pp.","Boycott, R. C. (2014). Afroedura major Onderstall, 1984. In: Bates, M. F., Branch, W. R., Bauer, A. M., Burger, M., Marais, J., Alexander, G. J. & de Villiers, M. (Eds.), Atlas and Red List of the Reptiles of South Africa, Lesotho and Swaziland. Suricata, 1. South African National Biodiversity Institute, Pretoria, pp. 94.","Kluge, A. G. (2001) Gekkotan lizard taxonomy. Hamadryad, 26, 1 - 209.","Monadjem, A., Boycott, R. C., Parker, V. & Culverwell, J. (2003) Threatened Vertebrates of Swaziland. Swaziland Red Data Book: Fishes, Amphibians, Reptiles, Birds and Mammals. Ministry of Tourism, Environment and Communications, Swaziland, Mbabane, 256 pp."]}

Afroedura rupestris sp. nov. (Fig. 7 A���B) Afroedura multiporis ���Abel Erasmus��� Jacobsen 1992 a, 1997 Holotype. TM 81233, adult male, Farm Perkeo 223 KT, 24 �� 27 'S, 30 �� 35 'E, Sekukhuneland District (2430 BC), Limpopo Province, Republic of South Africa, collector R.E. Newbery, 28 June 1982. Paratypes. TM 81224 ���81228, 81230���81232, 81234��� 81235, same data as holotype. Additional material: TM 47504, 52120, 81229, Bourk���s Luck, Pilgrim���s Rest District, Limpopo Province, Republic of South Africa (2430 DB). Etymology. The species name refers to the rupicolous nature of this (and most) members of the genus Afroedura. Diagnosis. A large Afroedura (to 62 mm SVL) differing from all other congeners by the following combination of characters: two pairs of enlarged subdigital lamellae per digit; tail faintly verticillate with three subcaudal rows and usually six supracaudal rows per verticil; dorsal scales bluntly keeled to trihedral; usually a single internasal scale; 20���24 precloacal pores in males. Description. (based on holotype TM 81233) Adult male; SVL 52.0 mm; tail 61.0 mm; mass in life 3.4 g. A slender elongated, lightly pigmented gecko with a rounded crown to the head. Rostral twice as broad as high, heptagonal; nostril pierced between rostral, first upper labial and three nasal scales; nasorostrals separated by a granular scale behind rostral. Scales on snout larger than on crown with some keeled scales along canthus rostralis and above and slightly anterior to the eyes; 2���3 supraciliary spines. Scales between the nasals and anterior margin of eye 12 and from posterior margin of eye to ear 20. Supralabials nine. Mental wedge-shaped, pentagonal, longer than broad; postmentals two; infralabials eight. Dorsals granular, heterogeneous, smaller paravertebrally, larger dorsolaterally, bluntly keeled to rounded, conical and juxtaposed. Scales rows at midbody 87. Ventral scales smooth, larger than dorsals, imbricate. Digits slender with two pairs of enlarged scansors and some enlarged inferomedian scales under the fourth toe. Other digits lacking enlarged inferomedian scales. Precloacal pores in a continuous row of 24, arranged in a chevron; three postcloacal spurs on each side of tail base. Tail long, somewhat flattened proximally; caudal scales arranged in poorly developed verticils with six dorsal and three ventral rows per verticil; dorsal scales smaller than ventrals and subimbricate; ventrals imbricate. Color. Olive-brown above with six, rarely seven, blackish wavy and irregular crossbands extending from occiput to sacrum. Pale whitish spots sometimes present at posterior margin of crossbands but may be absent. Limbs pale olive-brown with reticulate darker patterns on the thighs or with wavy stripes and spots. Crown of head paler, mostly without distinct markings. A darkish stripe from nostril to the anterior margin of eye. Two stripes extend from posterior margin of eye to above ear opening. In some individuals a single stripe extends through the eye and is continuous with the occipital bar. Tail with up to 10 crossbars. Body venter whitish, tail venter brownish with darker speckling. Variation. Paratypes and other specimens agree with the holotype in most features of scalation (Table 4). Single internasal typically present but two in paratype TM 81232 and nasorostrals in contact (no internasals) in TM 81234. Scales from eye to ear 16���19. Supraciliary spines four and scales on snout not keeled except above and anterior to the eye in TM 81232. Rostral and mental scales variable in relative proportion and shape. Postmentals generally two, but three in TM 47504, 52120 and 81234 and one in TM 81232. Female specimens lack precloacal pores; males typically with 20���26 pores with the exception of TM 81231 and TM 81225 which have 12, with a series of poreless scales separating left and right pore series. Postcloacal spurs 1���4. Caudal verticils with 6���7 subcaudal rows each. Tail 51.76���54.31 % of total length. 45.45 % of individuals (n = 11) exhibit caudal autotomy. Distribution. Apparently limited to the Eastern Escarpment of Limpopo Province, between the Abel Erasmus Pass and Bourke���s Luck (Fig. 6). Natural history. An exclusively rupicolous species living in crevices between rocks and under flakes of exfoliating rock along sandstone cliffs in Ohrigstad Mountain Bushveld (SVcb 26) (Mucina & Rutherford 2006) at altitudes between 850���1200 m a.s.l. (Fig. 8 A). Also found in kloofs and on rocky Blackreef quartzite outcrops on top of cliffs. Remarks. Afroedura rupestris sp. nov. shares with A. haackei, A. multiporis, and A. major bluntly keeled to trihedral dorsal scales and although not sampled in our molecular phylogeny, presumably belongs to the same clade as these taxa (Jacobsen 1992 a, 1997). It differs from these species in typically having 20���24 preclocal pores in males (versus 16���17 in A. multiporis, generally 25���28 in A. haackei, and 18���19 in A. major). It may further be differentiated from A. multiporis and A. major by its lower number of supracaudal scale rows per vertical (6 versus 7 and 8, respectively). It is significantly larger (max SVL 62 mm) than A. haackei (max SVL 52 mm), from which it also has a significantly higher number of supralabials (p A. major (max SVL 76 mm)., Published as part of Jacobsen, Niels H. G., Kuhn, Arianna L., Jackman, Todd R. & Bauer, Aaron M., 2014, A phylogenetic analysis of the southern African gecko genus Afroedura Loveridge (Squamata: Gekkonidae), with the description of nine new species from Limpopo and Mpumalanga provinces of South Africa, pp. 451-501 in Zootaxa 3846 (4) on pages 474-475, DOI: 10.11646/zootaxa.3846.4.1, http://zenodo.org/record/250495, {"references":["Jacobsen, N. H. G. (1992 a) Flat geckos (genus Afroedura) in the Transvaal. Journal of the Herpetological Association of Africa, 40, 22 - 25. http: // dx. doi. org / 10.1080 / 04416651.1992.9650313","Jacobsen, N. H. G. (1997) Family Gekkonidae, Genus Afroedura. In: Van Wyk, J. H. (Ed.), Proceedings of the FitzSimons Commemorative Symposium, South African Lizards: 50 years of Progress and Third H. A. A. Symposium on African Herpetology, Transvaal Museum, Pretoria, South Africa, 11 - 15 October 1993. Herpetological Association of Africa, Stellenbosch, pp. 34 - 39.","Mucina, L. & Rutherford, M. C. (2006) The Vegetation of South Africa, Lesotho and Swaziland. Strelitzia, 19. South African National Biodiversity Institute, Pretoria, 807 pp."]}

A. pondolia (Hewitt, 1925) (Fig. 3 E) Distribution. Low to middle elevations of the eastern Eastern Cape Province and southern and central KwaZulu- Natal (Bauer 2014 i) (Fig. 4). Remarks. Afroedura pondolia was long considered polytypic. Onderstall (1984) recognized five nonnominotypic subspecies, all of which are here considered valid species: A. marleyi, A. major, A. multiporis, A. haackei, and A. langi, and this interpretation was long accepted by most authors (e.g., Branch et al. 1988; Kluge 1991, 1993, 2001). All of the new species described herein would also have been placed in A. pondolia under Onderstall���s concept of the species. To avoid confusion with the A. pondolia group of earlier authors, we have here used the term A. nivaria group for the clade to which A. pondolia belongs., Published as part of Jacobsen, Niels H. G., Kuhn, Arianna L., Jackman, Todd R. & Bauer, Aaron M., 2014, A phylogenetic analysis of the southern African gecko genus Afroedura Loveridge (Squamata: Gekkonidae), with the description of nine new species from Limpopo and Mpumalanga provinces of South Africa, pp. 451-501 in Zootaxa 3846 (4) on page 471, DOI: 10.11646/zootaxa.3846.4.1, http://zenodo.org/record/250495, {"references":["Hewitt, J. (1925) On some new species of reptiles and amphibians from South Africa. Records of the Albany Museum, 3, 343 - 368, pls. xv - xix.","Bauer, A. M. (2014 i) Afroedura pondolia (Hewitt, 1925). In: Bates, M. F., Branch, W. R., Bauer, A. M., Burger, M., Marais, J., Alexander, G. J. & de Villiers, M. (Eds.), Atlas and Red List of the Reptiles of South Africa, Lesotho and Swaziland. Suricata, 1. South African National Biodiversity Institute, Pretoria, pp. 97 - 98.","Onderstall, D. (1984) Descriptions of two new subspecies of Afroedura pondolia (Hewitt) and a discussion of species groups within the genus (Reptilia: Gekkonidae). Annals of the Transvaal Museum, 33, 497 - 509.","Kluge, A. G. (1991) Checklist of Gekkonid lizards. Smithsonian Herpetological Information Service, 85, 1 - 35. http: // dx. doi. org / 10.5479 / si. 23317515.85.1","Kluge, A. G. (1993) Checklist of Gekkonid Lizards. International Gecko Society, San Diego, 245 pp.","Kluge, A. G. (2001) Gekkotan lizard taxonomy. Hamadryad, 26, 1 - 209."]}

A. africana (Boulenger, 1888) Distribution. Erongo Region, west central Namibia (Mouton & Mostert 1985; Griffin 2003) (Fig. 4). Remarks. Since the work of Haacke (1965) A. africana has been considered a polytypic species (e.g., Branch et al. 1988; R��sler 2000; Kluge 2001; Griffin 2003). We here consider A. africana as a monotypic central Namibian endemic (see below). All three of our genetic samples originate from the Erongo Region of central Namibia and are thus presumably referable to A. africana sensu stricto. However, the levels of divergence among these samples (up to 19.34 %) are comparable to or greater than interspecific differences elsewhere in the genus (Fig. 1), suggesting that further study of this taxon is required., Published as part of Jacobsen, Niels H. G., Kuhn, Arianna L., Jackman, Todd R. & Bauer, Aaron M., 2014, A phylogenetic analysis of the southern African gecko genus Afroedura Loveridge (Squamata: Gekkonidae), with the description of nine new species from Limpopo and Mpumalanga provinces of South Africa, pp. 451-501 in Zootaxa 3846 (4) on pages 469-470, DOI: 10.11646/zootaxa.3846.4.1, http://zenodo.org/record/250495, {"references":["Boulenger, G. A. (1888) On new or little-known South-African reptiles. Annals and Magazine of Natural History, series 6, 2, 136 - 141. http: // dx. doi. org / 10.1080 / 00222938809460892","Mouton, P. le F. N. & Mostert, D. P. (1985) Description of a new species of Afroedura (Loveridge) (Reptilia: Gekkonidae) from the south - western Cape. South African Journal of Zoology, 20, 246 - 249.","Griffin, M. (2003) Annotated Checklist and Provisional National Conservation Status of Namibian Reptiles. Namibia Scientific Society, Windhoek, [ii] + 169 pp.","Haacke, W. D. (1965) Additional notes on the Herpetology of South West Africa with descriptions of two new subspecies of geckos. Cimbebasia, 11, 1 - 40.","Rosler, H. (2000) Kommentierte Liste der rezent, subrezent und fossil bekannten Geckotaxa (Reptilia: Gekkonomorpha). Gekkota, 2, 28 - 153.","Kluge, A. G. (2001) Gekkotan lizard taxonomy. Hamadryad, 26, 1 - 209."]}

Afroedura granitica sp. nov. (Fig. 10 A) Afroedura langi 'Lillie' Jacobsen 1992 a, 1997 Holotype. TM 81187, adult male, Farm Lillie 148 KT, 24 ��04' S, 30 �� 51 ' E, Letaba district (2430 BB), Northern Province, collector R. E. Newbery, 11 November 1985. Paratypes. TM 81239 -81240, 81243��� 81247, same data as for holotype; TM 81241���81242, same locality as for holotype, collector N. H. G. Jacobsen, 17 May 1980. Etymology. The specific epithet refers to the decomposing granite hills to which this gecko appears restricted. Diagnosis. A medium to large Afroedura (maximum SVL 58.0 mm) differing from all other congeners by the following combination of characters: two pairs of enlarged subdigital lamellae per digit; tail not obviously verticillate and weakly flattened, with four subcaudal rows and seven supracaudal rows per verticil; dorsal scales smooth, 99���107 scale rows at midbody; internasal scales typically absent, a deep furrow between nasorostrals; 13���16 precloacal pores in males. Description. (based on holotype TM 81187) Adult male; 55.0 mm SVL; 62.0 mm TailL; mass before preservation 3.0 g. Dorsoventrally depressed; head ovate, distinctly wider than neck. Forelimbs small, hindlimbs robust. Rostral more than twice as wide as high; nostril pierced between rostral, first upper labial and three nasal scales; nasorostrals in broad contact behind rostral, with a deep furrow between them. Scales on snout rounded and juxtaposed, much larger than granules on crown of head; ten scales between nasals and eye and 18 scales between eye and ear; three supraciliary spines; supralabials nine. Mental wedge-shaped, much longer than wide and in contact with a single postmental. Infralabials seven. Dorsal granules uniform, smooth, rounded and juxtaposed. Midbody scales 100. Ventrals lozenge-shaped and imbricate. Digits with two pairs of scansors and four enlarged inferomedian scales under the 4 th toe. Fifteen continuous precloacal pores arranged in a curved row. Tail flattened proximally and thickening about one verticil length posterior to the cloaca. Tail inconspicuously verticillate and tapering, with caudal scales in seven dorsal and four ventral rows per verticil; supracaudals subimbricate, squarish to rectangular, with a rounded posterior margin; subcaudals large, squarish to hexagonal with rounded posterior margins, subimbricate. Two postcloacal spurs at base of tail. Color. The crown of the head is pale brown with dark brown spots and stripes that are also found on the snout. A dark brown stripe extends from the nostril through the eye, and above the ear to communicate with the occipital bar. Pale gray-brown to brown dorsally with six wavy dark brown irregular crossbars between occiput and sacrum. Crossbars are edged with black posteriorly and in their posterior indentations have white or whitish spots, particularly along the vertebral column. Variable pale and dark spots and blotches present between crossbars. The limbs have a variable dark brown reticulate pattern. Original tails with at least 10 dark brown to blackish crossbands with half bands and speckling in the intervening areas. Venter white, subcaudals speckled. Variation. Paratypes and other specimens agree with the holotype in most features of scalation (Table 4). Paratypes TM 81244 and TM 81246 with nasorostrals separated by a single granular scale. Scales between nasals and eye 10���12, from eye to ear 18���22. Postmentals 1���2. Supralabials 7���10. Midbody scale rows 99���107. No enlarged inferomedian scales under the fourth toe in TM 81243. Precloacal pores in male paratypes 13���16, females lacking pores but female paratype TM 81243 with a row of elongate scales in precloacal position. Postcloacal spurs 1���2, absent in some females. Original tails 53.18���58.3 % of total length. Degree of distinctness of tail whorls variable; faint, but clearly evident in some specimens. Tail has been autotomized in only 20 % of the specimens examined. Distribution. Restricted to granite hills and outcrops in the eastern Lowveld near Mica, Limpopo Province (Fig. 6). Natural history. Apparently restricted to decomposing granite outcrops and hillsides on the farm Lillie 148 KT in veld type SVl 3 Granite Lowveld (Mucina & Rutherford 2006) at elevations of 600���800 m a.s.l. These geckos inhabit crevices formed by exfoliating sheets on the underside of boulders, as well as crevices between rocks. They are occasionally found occupying the same retreats as large flat rock scorpions (Hadogenes troglodytes). The very similar A. langi occurs on small schistose outcrops. Remarks. Afroedura granitica sp. nov. shares a suite of characters with A. langi and four other species described herein as new. The affinities of all but one of these, A. leoloensis sp. nov. have been confirmed by our molecular phylogeny (Fig. 1). In comparison with other members of this group A. granitica sp. nov. (maximum SVL 58 mm) is substantially larger than A. langi and A. waterbergensis sp. nov. (both reaching only 46 mm) and A. leoloensis sp. nov. (40.5 mm maximum SVL). In comparison with the last of these species it also has far fewer precloacal pores (13���16 versus typically 31���35), whereas compared to A. waterbergensis sp. nov. (4���7 pores) and A. broadleyi sp. nov. (8���12 pores) it has more. In addition, in comparison with the superficially similar A. langi, the nasorostrals of A. granitica sp. nov. form a deep furrow between them (versus weakly raised nasorostrals forming a shallow furrow). Afroedura granitica sp. nov. is most similar in the basic diagnostic features used herein to A. pienaari sp. nov., to which it is rather distantly related. It may be distinguished by its larger size (58 versus 53 mm maximum SVL and even greater mean differences; Table 4), lower number of male precloacal pores (mean ~ 14 versus ~ 16), and higher number of eye to ear scales (mean of 19.10 versus 16.20), although there is overlap in all characters., Published as part of Jacobsen, Niels H. G., Kuhn, Arianna L., Jackman, Todd R. & Bauer, Aaron M., 2014, A phylogenetic analysis of the southern African gecko genus Afroedura Loveridge (Squamata: Gekkonidae), with the description of nine new species from Limpopo and Mpumalanga provinces of South Africa, pp. 451-501 in Zootaxa 3846 (4) on pages 483-484, DOI: 10.11646/zootaxa.3846.4.1, http://zenodo.org/record/250495, {"references":["Jacobsen, N. H. G. (1992 a) Flat geckos (genus Afroedura) in the Transvaal. Journal of the Herpetological Association of Africa, 40, 22 - 25. http: // dx. doi. org / 10.1080 / 04416651.1992.9650313","Jacobsen, N. H. G. (1997) Family Gekkonidae, Genus Afroedura. In: Van Wyk, J. H. (Ed.), Proceedings of the FitzSimons Commemorative Symposium, South African Lizards: 50 years of Progress and Third H. A. A. Symposium on African Herpetology, Transvaal Museum, Pretoria, South Africa, 11 - 15 October 1993. Herpetological Association of Africa, Stellenbosch, pp. 34 - 39.","Mucina, L. & Rutherford, M. C. (2006) The Vegetation of South Africa, Lesotho and Swaziland. Strelitzia, 19. South African National Biodiversity Institute, Pretoria, 807 pp."]}

Afroedura leoloensis sp. nov. (Fig. 10 B) Afroedura pondolia langi (part) Visser 1984 a (fig. p. 61) Afroedura langi 'Leolo' Jacobsen 1992 a, 1997 Holotype. TM 81113, adult male, Farm Hendriksplaats 281 KT, 24 �� 38 ' S, 30 ��08' E, Lydenburg District, (2430 CA) Mpumalanga Province, Republic of South Africa, collectors R. E. Newbery and W. Petersen, 15 November 1985. Paratypes. TM 81115, TM 81125, 81127, same data as for holotype; TM 81119, 81124, Farm Maandagshoek 254 KT, Sekhukhuneland District, Limpopo Province (2430 CA) collector N. H. G. Jacobsen, 25 October 1980; TM 81112, 81116��� 81118, same locality as TM 81119, collector R.E.Newbery, 29 June 1982; TM 81126, Farm Kalkfontein 367 KT, Lydenburg District, Mpumalanga Provicne (2430 CC), collector N. H. G. Jacobsen, 23 April 1981; TM 81114, 81121, 81123, 81128, same locality as TM 81126, collectors R. E. Newbery and W. Petersen, 16 November 1985; TM 81122, Kgoloko lokasie, Sekhukhuneland District, Limpopo Province (2429 DB), collector R. E. Newbery, 27 October 1980; TM 81120, Farm De Grooteboom 373 KT, Lydenburg District, Mpumalanga Province (2430 CC), collector N. H. G. Jacobsen, 23 October 1981. Etymology. The specific epithet refers to the Leolo hills in Sekhukhuneland, Limpopo Province, where it was first discovered. Diagnosis. A small Afroedura (maximum SVL 40.5 mm) differing from all other congeners by the following combination of characters: two pairs of enlarged subdigital lamellae per digit; tail faintly verticillate and flattened near base, with four subcaudal rows and 6���7 supracaudal rows per verticil; dorsal scales smooth, 87���95 scale rows at midbody; internasal scales typically absent; 31���35 precloacal pores in males. Description. (based on holotype TM 81113) Adult male; 37.0 mm SVL; 45.0 mm TailL; mass before preservation 0.9 g. Body small and slender, dorsoventrally depressed; head oval, wider than the neck. Rostral approximately 2.5 times wider than high; nostril pierced between rostral, first upper labial and three nasal scales; nasorostrals in moderate contact behind rostral. Scales on snout hexagonal, flattened and much larger than scales on crown of head; nine scales between nasals and eye and 16 scales between eye and ear. Four supraciliary spines. Supralabials 10. Mental wedge-shaped, much longer than wide and in contact with two postmentals. Infralabials eight. Dorsal scales minute, more-or-less homogeneous, smooth, juxtaposed to subimbricate, rounded to slightly hexagonal. Midbody scales 89. Ventrals large, smooth and imbricate. Digits with two pairs of enlarged scansors and six enlarged inferomedian scales under the fourth toe. Precloacal pores in a continuous, almost straight row of 34. Tail broad and flattened near the base, tapering to a fine tip, faintly verticillate; caudal scales arranged in six dorsal and four ventral rows per verticil. Supracaudals subimbricate, almost rectangular; subcaudals as broad as long and imbricate. Two postcloacal spurs on either side of tail base. Color. Pale brown to brown dorsally with 7���8 dark brown irregular crossbands extending from the occiput to sacrum. Crossbands with darker posterior margins and, in most specimens, a white vertebral spot just posterior to each crossband. Limbs longitudinally striped or banded. The paler areas between the stripes are spotted dark brown. Crown of head pale brown with darker spotting and other variegations. Tail with 10 blackish crossbands from base to tip. Venter pinkish; tail brownish with darker markings. Variation. Paratypes and other specimens agree with the holotype in most features of scalation (Table 4). Nasorostrals in broad contact behind rostral but separated posteriorly by a single granule in TM 81120. Scales between nasals and eye 9���12, from eye to ear 16���18. Five supraciliary spines in TM 81115. Mental as long as broad in some specimens; postmentals three in TM 811141. Supralabials 8���10. Infralabials 6���10. Midbody scale rows 87���95. 0���8 enlarged inferomedian scales under fourth toe. Precloacal pores in male paratypes 31���35 (except for TM 81126, which has only 11), females lacking pores. Original tails 50.00��� 55.9 % of total length. Supracaudal scales in 6 or 7 rows per tail whorl. Tail has been autotomized in 54.5 % of the specimens examined (n= 17). Distribution. Endemic to the Leolo Hills and outcrops above the Steelpoort River on either side of the border between Mpumalanga and Limpopo Provinces (Fig. 6). Natural history. Exclusively rupicolous, apparently limited to norite and granitic formations. Afroedura leoloensis sp. nov. lives in narrow crevices under exfoliating rock. These are usually on the underside of boulders with the openings facing downwards, protecting them from rain. The species is found in Sekhukhune Mountain Bushveld (SVcb 28) and Leolo Summit Sourveld (GM 20) (Mucina & Rutherford 2006) at an elevation of 1200���1800 m a.s.l. Two eggs are laid at a time and measure 8.9 ���9.0 x 6.1���6.4 mm with a mass of 0.2 g. The eggs appear to be laid in midsummer and are initially soft-shelled, adhering to the rock and then hardening. Remarks. Afroedura leoloensis sp. nov. is a member of the A. langi clade (Figs. 1���2; see Remarks under A. granitica sp. nov.). This species exhibits the highest number of precloacal pores (31���35) not only in its clade, but in the genus as a whole, and may thus be distinguished from its congeners., Published as part of Jacobsen, Niels H. G., Kuhn, Arianna L., Jackman, Todd R. & Bauer, Aaron M., 2014, A phylogenetic analysis of the southern African gecko genus Afroedura Loveridge (Squamata: Gekkonidae), with the description of nine new species from Limpopo and Mpumalanga provinces of South Africa, pp. 451-501 in Zootaxa 3846 (4) on pages 486-487, DOI: 10.11646/zootaxa.3846.4.1, http://zenodo.org/record/250495, {"references":["Visser, J. 1984 a. Diksterte sukkel nie met blyplek. Akkedisse van Suider - Afrika 14. Landbouweekblad, 4 June 1984, 60 - 61, 63.","Jacobsen, N. H. G. (1992 a) Flat geckos (genus Afroedura) in the Transvaal. Journal of the Herpetological Association of Africa, 40, 22 - 25. http: // dx. doi. org / 10.1080 / 04416651.1992.9650313","Jacobsen, N. H. G. (1997) Family Gekkonidae, Genus Afroedura. In: Van Wyk, J. H. (Ed.), Proceedings of the FitzSimons Commemorative Symposium, South African Lizards: 50 years of Progress and Third H. A. A. Symposium on African Herpetology, Transvaal Museum, Pretoria, South Africa, 11 - 15 October 1993. Herpetological Association of Africa, Stellenbosch, pp. 34 - 39.","Mucina, L. & Rutherford, M. C. (2006) The Vegetation of South Africa, Lesotho and Swaziland. Strelitzia, 19. South African National Biodiversity Institute, Pretoria, 807 pp."]}

Afroedura waterbergensis sp. nov. (Fig. 11 A) Afroedura langi 'Waterberg' Jacobsen 1990, 1992a, 1997 Holotype. TM 81266, adult female, Farm Waterval 601 LQ, 23 �� 53 ' S, 27 �� 39 ' E, Waterberg District (2327 DC), Limpopo Province, collectors N. H. G. Jacobsen & R. E. Newbery, 20 January 1987. Paratypes. TM 81267, 81269, 81273, Farm Fancy 556 LQ, Waterberg District, Limpopo Province (2327 DC), collectors N. H. G. Jacobsen & R. E. Newbery, 22 January 1987. TM 81268, 81270, Farm Fourieskloof 557 LQ, Waterberg District, Limpopo Province (2327 DC), collector R. E. Newbery, 26 September 1979; TM 81271���81272, same data as for holotype. Etymology. The specific epithet refers to the Waterberg massif, Limpopo Province, to which the species is endemic. Diagnosis. A small Afroedura (maximum SVL 46 mm) differing from all other congeners by the following combination of characters: two pairs of enlarged subdigital lamellae per digit; tail moderately verticillate (semiverticillate) and flattened near base, with four subcaudal rows and 7 supracaudal rows per verticil; dorsal scales smooth, 92���99 scale rows at midbody; internasal scales absent; 4���7 precloacal pores in males. Description. (based on holotype TM 81266) Adult female, SVL 44.0 mm; TailL 53.0 mm; mass before preservation 2.2 g. A small to medium-sized, dorsoventrally depressed gecko. Head oval, neck thick, almost as wide as head. Rostral approximately 2.5 times wider than high; nostril pierced between rostral, first supralabial and three nasals; nasorostrals large and in contact behind rostral. Scales on snout rounded but not flattened, decreasing in size posteriorly to crown of head; 9���10 granular scales between nasals and eye and 18 from eye to ear; 3���4 supraciliary spines; supralabials 11. Mental longer than broad and not wedge-shaped; postmentals two; infralabials nine. Dorsal scales smooth, juxtaposed, uniform and rounded, becoming larger and oblique laterally. Midbody scales in 101 rows. Ventral scales hexagonal to rounded, smooth and juxtaposed. Hindlimbs robust, feet moderately enlarged. Digits with two pairs of enlarged scansors and 4 th toe with six enlarged inferomedian scales. Precloacal pores absent. Tail semi-verticillate, flattened and tapered, constricted at base and widening at the beginning of the first verticil. Caudal scales arranged in verticils with seven dorsal and four ventral rows. Supracaudals subimbricate, more or less square with a rounded posterior margin; subcaudals imbricate wider than or as wide as long. Two postcloacal spurs on either side of tail base. Color. Brown to pale brown above with 6���7 wavy dark brown to blackish crossbands extending from the occiput to the sacrum. Posterior margin of crossbands darker and with white spots in indentations. A vertebral row of white spots is most pronounced. Crown of head marbled with dark brown and a dark brown stripe extending from the nostril through the eye, merging into the occipital crossbar. Limbs striped with dark brown, with interstices spotted and a reticulate pattern on the thighs. Venter white to whitish pink. Tail with seven crossbands at regular intervals. Underside of tail brown with incomplete pale crossbars. Variation. Paratypes and other specimens agree with the holotype in most features of scalation (Table 4). Scales from eye to ear 16���18. Mental in some specimens only as long as wide. Supralabials 8���11. Infralabials 7���9. Midbody scale rows 92���99. Eight enlarged inferomedian scales under 4 th toe in TM 81272. Precloacal pores in males in continuous series of 4���7. 0���3 postcloacal spurs on either side of tail base. Original tails 50.0��� 56.4 % of total length. Supracaudal scales in 6���7 (rarely 8) rows per tail whorl. 33 % of specimens have regenerated tails (n= 6). Distribution. Endemic to the western Waterberg massif, Limpopo Province (Fig. 6). Natural history. Afroedura waterbergensis sp. nov. is a rupicolous gecko found in crevices between rocks on rocky outcrops and cliff faces of Waterberg sandstone. It inhabits both vertical and horizontal dry crevices. It has only been observed in Waterberg Mountain Bushveld (SVcb 17) (Mucina & Rutherford 2006) at an elevation of 1000 m a.s.l. Remarks. Afroedura waterbergensis sp. nov. is the westernmost member of its genus in Limpopo Province. It may be more widespread along the Waterberg than the current records indicate. It is a member of the A. langi complex and may be easily distinguished from all other members of this clade on the basis of its low number of precloacal pores (see Table 4)., Published as part of Jacobsen, Niels H. G., Kuhn, Arianna L., Jackman, Todd R. & Bauer, Aaron M., 2014, A phylogenetic analysis of the southern African gecko genus Afroedura Loveridge (Squamata: Gekkonidae), with the description of nine new species from Limpopo and Mpumalanga provinces of South Africa, pp. 451-501 in Zootaxa 3846 (4) on pages 488-490, DOI: 10.11646/zootaxa.3846.4.1, http://zenodo.org/record/250495, {"references":["Jacobsen, N. H. G. (1990) A Herpetological Survey of the Transvaal. Unpublished Ph. D. Dissertation, University of Natal, Durban, South Africa, 1621 pp.","Jacobsen, N. H. G. (1992 a) Flat geckos (genus Afroedura) in the Transvaal. Journal of the Herpetological Association of Africa, 40, 22 - 25. http: // dx. doi. org / 10.1080 / 04416651.1992.9650313","Jacobsen, N. H. G. (1997) Family Gekkonidae, Genus Afroedura. In: Van Wyk, J. H. (Ed.), Proceedings of the FitzSimons Commemorative Symposium, South African Lizards: 50 years of Progress and Third H. A. A. Symposium on African Herpetology, Transvaal Museum, Pretoria, South Africa, 11 - 15 October 1993. Herpetological Association of Africa, Stellenbosch, pp. 34 - 39.","Mucina, L. & Rutherford, M. C. (2006) The Vegetation of South Africa, Lesotho and Swaziland. Strelitzia, 19. South African National Biodiversity Institute, Pretoria, 807 pp."]}

Afroedura maripi sp. nov. (Fig. 9 A) Afroedura 'maripi' Jacobsen 1992 a, 1997 Holotype. TM 81262, adult male, Mariepskop, 24 �� 32 ' S, 30 �� 53 ' E, Pilgrim's Rest District (2430 DB), Limpopo Province, Republic of South Africa, collector R. E. Newbery, 25 June 1982. Paratypes. (all from same locality as holotype) TM 81248 ���81249, 81251���81255, 81257���81259, 81261, 81263���81264, same data as holotype; TM 81250, 81256, 81260, 81265, collector N. H. G. Jacobsen, 29 October 1980; TM 58031���58032, collector R. E. Newbery, 5 June 1984; TM 58118, collector R. E. Newbery, 21 November 1984; TM 62907���62909, collector R. E. Newbery, 21 February 1985; TM 64021, collector R. E. Newbery, 1 June 1984. Etymology. The species name refers to the local name for Mariepskop, the type locality. Diagnosis. A large Afroedura (to 63 mm SVL) differing from all other congeners by the following combination of characters: two pairs of enlarged subdigital lamellae per digit; a single internasal granule between nasorostrals; tail not obviously verticillate, with 4���5 subcaudal rows and 6���8 supracaudal rows per verticil; dorsal scales smooth, in 96���108 rows at midbody; a single internasal scale; 11���13 precloacal pores in males. Description. (based on holotype TM 81262) Adult male; SVL 57.0 mm; tail 61.0 mm (partly regenerated); mass in life 3.3 g. Head and body dorso-ventally depressed. Head oval and wider than neck, body broad and limbs relatively short, digits stout. Rostral twice as broad as high, more or less octagonal; nostril pierced between rostral, first upper labial and three nasal scales; nasorostrals separated by a single large granular scale. Scales on snout rounded to conical, larger than those on crown. Scales between nasals and anterior margin of eye 12, and from eye to ear 24. Four supraciliary spines at posterodorsal corner of eye. Supralabials 10. Mental wider than deep; postmentals three; infralabials eight. Dorsals smooth, uniform, each scale raised posteriorly and juxtaposed; scales at midbody 107. Ventral scales smooth, flattened and imbricate. Digits stout with two pairs of enlarged scansors and five enlarged inferomedian scales under fourth toe, not reaching the base. Precloacal pores 13, arranged in a continuous ���V���-shaped pattern. Tail regenerated and constricted at base. Two postcloacal spurs on either side of tail base. Color. Olive-brown to olive-gray above with variable blackish crossbars extending from occiput to sacrum. Crown of head variably marked, filigreed to reticulate. Interstices between bars are reticulate laterally. Limbs barred or variegated. Tail with approximately 10 black bars, becoming more irregular and diffuse posteriorly. Chin, throat, chest and abdomen whitish. Underside of tail grayish brown, faintly barred. Variation. Paratypes and other specimens agree with the holotype in most features of scalation (Table 4). Endolymphatic sacs distended in adult female paratype TM 81260. Scales between nasals and eye 11���13, from eye to ear 20���23. Supraciliary spines 2���4. Rostral and mental scales variable in relative proportion and shape. Postmentals 2���3. Supralabials 9���10, infralabials 7���8. Midbody scale rows 96���108. Inferomedian scales under the fourth toe 5���8. Precloacal pores in male paratypes 11���13, females lacking pores. Postcloacal spurs 2���3, rarely four. Original tails 52.1���53.9 % of total length and tapered. Tail with 6���8 (mostly seven) dorsal and four (rarely five) ventral scale rows per indistinct verticil. Supracaudals subimbricate with a rounded to pointed posterior margin; subcaudals broad and imbricate.Caudal autotomy evident in 76.3 % of individuals. Distribution. Restricted to the slopes of Mariepskop and God���s Window area of the Eastern Escarpment, in apparently two disjunct populations (Fig. 6). Occurrence in the latter area is verified by a photo voucher. Natural history. Usually associated with rocky outcrops on west-facing slopes of Mariepskop, frequently in the partial shade of pine plantations but also found on outcrops of Black Reef quartzites on top of the escarpment in the shade of indigenous montane forest. Preferred retreats include crevices under flakes of exfoliating rock and dark overhangs of large boulders. Found in Ohrigstad Mountain Bushveld (SVcb 26) and Northern Escarpment Quartzitic Sourveld (GM 23) (Mucina & Rutherford 2006) at elevations of 1700���1900 m a.s.l. data sample sizes, maxima and adult means �� 1 S.D. are given by sex f��r taxa newly described herein; maxima ��nly (regardless ��f sex) are pr��vided f��r previ��usly rec��gnized taxa. Internasal P��stmenal c��unts sh��w the character states rec��rded and, parenthetically, the number ��f specimens exhibiting each state. verticil c��unts are presented as supracaudal scale r��ws f��ll��wed subcaudal scale r��ws. All ��ther c��unts are presented as mean �� 1 S.D. Sample sizes f��r scale c��unts are the same as listed f��r Internasal (Intern) and P��stmental (P��stm) c��nditi��ns. ��ax��n SvL (mm) Mass (g) Intern (n) P��stm (n) Supralabials Infralabials Eye-Ear Midb��dy Scale verticils Precl��acal R��ws supra���sub P��res transvaalica gr��up transvaalica 64.0 5.4 0(35) 2 (28) 9.60 �� 0.80 8.81 �� 0.70 18.13 �� 0.81 109.13 �� 3.83 7-8 6.20 �� 0.83 1 (2) 3 (9) 5 nivaria gr��up pondolia 58.0 - 1 (8) 1 (2) 9.27 8.30 23.86 102.92 6-7 14.50 2 (5) 2 (14) 4 3 (4) multiporis gr��up haackei 52.0 2.3 0(3) 2 (16) 9.06 �� 0.68 8.38 �� 0.62 16.86 �� 0.86 90.36 �� 4.27 6-7 25.83 �� 1.64 1 (13) 3-4 major 76.0 - 2 (3) 2 (3) 11.00 9.33 22.17 106.50 8 18.50 4 multiporis 66.5 5.5 1 (11) 1 (1) 9.10 �� 0.70 8.60 �� 0.67 20.60 �� 1.51 99.60 �� 4.33 6-7 16.67 �� 0.58 2 (10) 3-4 rupestris sp. nov. 9 ♂, 3 ♀ 8 ♂, 3 ♀ 0(1) 1 (1) 9.79 �� 1.05 8.36 �� 0.63 18.77 �� 1.17 90.14 �� 4.90 6-7 22.12 �� 1.96 62.0♂,56.0♀ 5.8 ♂, 3.9 ♀ 1 (11) 2 (10) 3 54.5 + 4.0 ♂ 3.5 + 1.1 ♂ 2 (2) 3 (3) 54.7 + 1.5 ♀ 3.0 + 1.0♀ marleyi gr��up marleyi 36.0 1.2 1 (17) 2 (14) 8.47 �� 0.87 8.18 �� 0.64 19.1 �� 1.68 88.77 ��3.00 6-7 13.40 �� 0.89 3 (3) 4-5 maripi sp. nov. 7 ♂, 9 ♀ 7 ♂, 9 ♀ 1 (13) 2 (13) 9.59 �� 0.67 8.18 �� 0.73 21.81 �� 1.38 102.56 �� 3.81 7-8 12.29 �� 0.95 63.0♂,63.0♀ 3.8 ♂, 6.5 ♀ 3 (8) 4-5 56.7 + 5.2 ♂ 3.0 + 0.5 ♂ 56.7 + 5.1 ♀ 3.3 + 1.4 ♀ pongola sp. nov. 5 ♂, 5 ♀ 5 ♂, 3 ♀ 1 (12) 2 (8) 9.00�� 0.58 8.08 �� 0.28 20.77 �� 1.42 98.22 �� 3.07 6-7 22.40 �� 0.89 35.0♂,40.0♀ 1.0♂, 0.9 ♀ 2 (1) 3 (5) 4-5 34.1 + 0.6 ♂ 0.8 + 0.2 ♂ 38.0 + 1.4 ♀ 0.8 + 0.1 ♀ ������ continued on the next page Two eggs are laid at a time during August���October. They are soft when laid, adhering to the rock and hardening, as has been reported for A. halli (Power 1939). The eggs measure 9.5���10.9 x 7.8���9.3 mm and take approximately two months to hatch. Neonates measure 23.0 mm SVL, 21.0 mm TailL, with a mass of 0.25-0.30 g. Remarks. Afroedura maripi sp. nov. was included by Jacobsen (1990, 1992a, 1997) in his A. pondolia complex, but our molecular results suggest that while other members of this proposed complex��� A. marleyi, A. pongola sp. nov., and presumably A. rondavelica sp. nov. (also unsampled genetically), as well as A. maripi sp. nov. ���do form a monophyletic group, referred to as the A. marleyi group, A. pondolia is only distantly related. It may be distinguished from A. pongola sp. nov. and especially A. marleyi on the basis of its greater number of midbody scale rows (mean of 102.56 versus 98.22 and 88.77, respectively) and larger size (maximum SVL 63 mm versus 40 mm and 36 mm, respectively) and has substantially fewer precloacal pores than A. pongola sp. nov. (11���13 versus 21���23). It also has significantly more supralabial scales than A. marleyi (p versus 7���9) than A. rondavelica sp. nov., Published as part of Jacobsen, Niels H. G., Kuhn, Arianna L., Jackman, Todd R. & Bauer, Aaron M., 2014, A phylogenetic analysis of the southern African gecko genus Afroedura Loveridge (Squamata: Gekkonidae), with the description of nine new species from Limpopo and Mpumalanga provinces of South Africa, pp. 451-501 in Zootaxa 3846 (4) on pages 477-481, DOI: 10.11646/zootaxa.3846.4.1, http://zenodo.org/record/250495, {"references":["Jacobsen, N. H. G. (1992 a) Flat geckos (genus Afroedura) in the Transvaal. Journal of the Herpetological Association of Africa, 40, 22 - 25. http: // dx. doi. org / 10.1080 / 04416651.1992.9650313","Jacobsen, N. H. G. (1997) Family Gekkonidae, Genus Afroedura. In: Van Wyk, J. H. (Ed.), Proceedings of the FitzSimons Commemorative Symposium, South African Lizards: 50 years of Progress and Third H. A. A. Symposium on African Herpetology, Transvaal Museum, Pretoria, South Africa, 11 - 15 October 1993. Herpetological Association of Africa, Stellenbosch, pp. 34 - 39.","Mucina, L. & Rutherford, M. C. (2006) The Vegetation of South Africa, Lesotho and Swaziland. Strelitzia, 19. South African National Biodiversity Institute, Pretoria, 807 pp.","Power, J. H. (1939) A note on the habits, life history and distribution of Oedura halli Hewitt. South African Journal of Science, 36, 374 - 376.","Jacobsen, N. H. G. (1990) A Herpetological Survey of the Transvaal. Unpublished Ph. D. Dissertation, University of Natal, Durban, South Africa, 1621 pp."]}

Afroedura transvaalica Group Diagnosis. Two pairs of enlarged scansors per digit, tail strongly verticillate and flattened., Published as part of Jacobsen, Niels H. G., Kuhn, Arianna L., Jackman, Todd R. & Bauer, Aaron M., 2014, A phylogenetic analysis of the southern African gecko genus Afroedura Loveridge (Squamata: Gekkonidae), with the description of nine new species from Limpopo and Mpumalanga provinces of South Africa, pp. 451-501 in Zootaxa 3846 (4) on page 467, DOI: 10.11646/zootaxa.3846.4.1, http://zenodo.org/record/250495

Afroedura hawequensis Group Diagnosis. Large sized (maximum SVL 83 mm), three pairs of enlarged scansors on digits II���V, first digit with a single scansor, tail verticillate and flattened, precloacal pores> 25., Published as part of Jacobsen, Niels H. G., Kuhn, Arianna L., Jackman, Todd R. & Bauer, Aaron M., 2014, A phylogenetic analysis of the southern African gecko genus Afroedura Loveridge (Squamata: Gekkonidae), with the description of nine new species from Limpopo and Mpumalanga provinces of South Africa, pp. 451-501 in Zootaxa 3846 (4) on page 467, DOI: 10.11646/zootaxa.3846.4.1, http://zenodo.org/record/250495

A. marleyi (FitzSimons, 1930) (Fig. 5 B) Distribution. Coastal plain of Zululand, Kwazulu-Natal inland to the Lebombo Mountains and through Swaziland to southeastern Mpumalanga, and possibly adjacent Mozambique (Bauer 2014 f) (Figs. 4, 6). Remarks. This species was previously considered a subspecies of A. pondolia (Onderstall 1984) until elevated without comment by Branch (1998). Our samples come from a relatively limited area of the species��� range and we suspect that deeper sampling would reveal greater substructure within A. marleyi., Published as part of Jacobsen, Niels H. G., Kuhn, Arianna L., Jackman, Todd R. & Bauer, Aaron M., 2014, A phylogenetic analysis of the southern African gecko genus Afroedura Loveridge (Squamata: Gekkonidae), with the description of nine new species from Limpopo and Mpumalanga provinces of South Africa, pp. 451-501 in Zootaxa 3846 (4) on page 476, DOI: 10.11646/zootaxa.3846.4.1, http://zenodo.org/record/250495, {"references":["FitzSimons, V. F. (1930) Descriptions of new South African Reptilia and Batrachia, with distribution records of allied species in the Transvaal Museum collection. Annals of the Transvaal Museum, 14, 20 - 48.","Onderstall, D. (1984) Descriptions of two new subspecies of Afroedura pondolia (Hewitt) and a discussion of species groups within the genus (Reptilia: Gekkonidae). Annals of the Transvaal Museum, 33, 497 - 509."]}

A. amatolica (Hewitt, 1925) Distribution. Amatola Mountains, Eastern Cape Province, South Africa (Bauer 2014 b) (Fig. 4). Remarks. Loveridge���s (1947) synonymy of A. amatolica with A. nivaria was accepted by at least some authors as late as the 1980 s (Branch 1981), owing to the long-confused application of names to Eastern Cape Afroedura, but since 1984 it has generally be regarded as specifically distinct (Visser 1984 b)., Published as part of Jacobsen, Niels H. G., Kuhn, Arianna L., Jackman, Todd R. & Bauer, Aaron M., 2014, A phylogenetic analysis of the southern African gecko genus Afroedura Loveridge (Squamata: Gekkonidae), with the description of nine new species from Limpopo and Mpumalanga provinces of South Africa, pp. 451-501 in Zootaxa 3846 (4) on pages 470-471, DOI: 10.11646/zootaxa.3846.4.1, http://zenodo.org/record/250495, {"references":["Hewitt, J. (1925) On some new species of reptiles and amphibians from South Africa. Records of the Albany Museum, 3, 343 - 368, pls. xv - xix.","Branch, W. R. (1981) An annotated checklist of the lizards of the Cape Province, South Africa. Annals of the Cape Provincial Museums (Natural History), 13, 141 - 167.","Visser, J. 1984 b. Diksterte van die Amatolaberge. Akkedisse van Suider - Afrika 15. Landbouweekblad, 29 June 1984, 48 - 49, 51, 53."]}

A. halli (Hewitt, 1935) Distribution. Western Lesotho and adjacent southeastern Free State and Eastern Cape provinces, South Africa (Bates & Bauer 2014 a) (Fig. 4). Remarks. This species was previously considered a subspecies of A. karroica (e.g., Loveridge 1947; Wermuth 1965; Branch et al. 1988; Kluge 1991, 1993) but was elevated to specific status by Bates (1996), although without substantial discussion, and some later authors, especially outside of South Africa, have continued to include it within A. karroica (e.g., Kluge 2001). One subspecies, A. k. wilmoti, has been variously recognized as valid (FitzSimons 1943) or not (Loveridge 1947; Branch 1981). Its status is under review (Bates & Branch 1998). Our sampling included a single specimen resembling A. halli that was deeply divergent from all other samples (Fig. 1), supporting the findings of Makhubo (2013), who found multiple divergent clades within the species. Although the distinctiveness of this specimen in our two-gene analysis may be artificially high due to the absence of RAG 1 sequence (Table 2), the> 20 % ND 2 divergence from typical A. halli, which vary among themselves by 0.20���3.02 %, supports the view that this represents a different species., Published as part of Jacobsen, Niels H. G., Kuhn, Arianna L., Jackman, Todd R. & Bauer, Aaron M., 2014, A phylogenetic analysis of the southern African gecko genus Afroedura Loveridge (Squamata: Gekkonidae), with the description of nine new species from Limpopo and Mpumalanga provinces of South Africa, pp. 451-501 in Zootaxa 3846 (4) on page 471, DOI: 10.11646/zootaxa.3846.4.1, http://zenodo.org/record/250495, {"references":["Hewitt, J. (1935) Some new forms of batrachians and reptiles from South Africa. Records of the Albany Museum, 4, 283 - 357.","Bates, M. F. & Bauer, A. M. (2014 a) Afroedura halli (Hewitt, 1935). In: Bates, M. F., Branch, W. R., Bauer, A. M., Burger, M., Marais, J., Alexander, G. J. & de Villiers, M. (Eds.), Atlas and Red List of the Reptiles of South Africa, Lesotho and Swaziland. Suricata, 1. South African National Biodiversity Institute, Pretoria, pp. 91 - 92.","Loveridge, A. (1947) Revision of the African lizards of the Family Gekkonidae. Bulletin of the Museum of Comparative Zoology, 98, 1 - 469, pls. 1 - 7.","Wermuth, H. (1965) Liste der rezenten Amphibien und Reptilien, Gekkonidae, Pygopodidae, Xantusiidae. Das Tierreich, 80, xxii + 246 pp.","Kluge, A. G. (1991) Checklist of Gekkonid lizards. Smithsonian Herpetological Information Service, 85, 1 - 35. http: // dx. doi. org / 10.5479 / si. 23317515.85.1","Kluge, A. G. (1993) Checklist of Gekkonid Lizards. International Gecko Society, San Diego, 245 pp.","Bates, M. F. (1996) In search of the elusive flat geckos of the eastern Free State. Culna 51, 13 - 15.","Kluge, A. G. (2001) Gekkotan lizard taxonomy. Hamadryad, 26, 1 - 209.","FitzSimons, V. F. (1943) The lizards of South Africa. Memoirs of the Transvaal Museum, 1, i - xv + 1 - 528, + 24 pls + map.","Branch, W. R. (1981) An annotated checklist of the lizards of the Cape Province, South Africa. Annals of the Cape Provincial Museums (Natural History), 13, 141 - 167.","Bates, M. F. & Branch, W. R. (1998) Taxonomy and distribution of the Afroedura nivaria and A. karroica complexes (Sauria: Gekkonidae). Herpetological Association of Africa, Fifth Symposium, University of Stellenbosch, 14 - 16 September 1998, Programme and Abstracts. [2 unnumbered pages]","Makhubo, B. G. (2013) Phylogenetic and Morphological Analysis of the Afroedura nivaria (Reptilia: Gekkonidae) Species Complex in South Africa. Unpublished M. Sc. Thesis, University of Stellenbosch, xiii + 106 pp."]}

Afroedura broadleyi sp. nov. (Fig. 10 C) Afroedura langi 'Soutpansberg' Jacobsen 1992 a, 1997; Kirchhof et al. 2010 Afroedura 'Matlala' Jacobsen 1992 a, 1997 Holotype. TM 81316, adult male, Farm Peover 772 MS, 22 �� 59 ' S, 29 �� 44 ' E, Soutpansberg District (2229 DC), Limpopo Province, Republic of South Africa, collector R. E. Newbery, 6 November 1985. Paratypes. TM 81317 ���81322, 81341, same data as for holotype; TM 81338-81340, Farm Leek 769 MS, Soutpansberg District (2229 DC), Limpopo Province, collector R. E. Newbery 6 November 1985. Additional material examined (all localities in Limpopo Province). TM 81323-81326, Farm Outlook 789 MS, Soutpansberg District (2229 DD); TM 81327 ���81329, 81330- 81336, Farm Newgate 802 MS, Soutpansberg District (2229 DD); TM 81337, Mutshenzheni, Sibasa District (2230 CD); TM 81300 -81307, 81309- 81314, Farm Leipzig 264 LR, Bochum District (2328 BB); TM 81288 ���81299, 81308, Farm Urk 10 LS, Bochum District (2329 AA); TM 81188���81207 Matlala Mountain, Seshego District (2329 CC). Etymology. The species is named for our friend and colleague Dr. Donald G. Broadley in honor of his many contributions to African herpetology and in celebration of his 80 th birthday. Diagnosis. A mid-sized Afroedura (maximum SVL 56.0 mm) differing from all other congeners by the following combination of characters: two pairs of enlarged subdigital lamellae per digit; tail moderately verticillate (semi-verticillate) and flattened near base, with four subcaudal rows and 6���8 supracaudal rows per verticil; dorsal scales smooth, 88���106 scale rows at midbody; internasal scales typically absent; 8���12 precloacal pores in males. Description. (based on holotype TM 81316) Adult male; 45.5 mm SVL; 49.0 mm TailL (part regenerated); mass before preservation 2.3 g. Moderately dorsoventrally depressed; head oval, wider than neck; limbs robust with stout digits; tail depressed, long and tapered, distal portion regenerating. Rostral twice as broad as high; nostril pierced between rostral, first upper labial and three nasals; nasorostrals large and in contact behind rostral. Scales on snout large, heterogeneous and flattened, decreasing in size posteriorly, smallest on crown of head; 10 scales from nasals to eye and 21 scales from eye to ear; three supraciliary spines. Supralabials nine. Mental narrow, wedge-shaped, longer than wide and in contact with two postmentals. Infralabials seven. Dorsal scales relatively homogeneous, rounded,smooth, slightly flattened and juxtaposed; midbody scales 91. Ventral scales larger, imbricate and almost hexagonal. Digits with two pairs of enlarged scansors and one enlarged inferomedian scale under the 4 th toe. Precloacal pores 11, arranged in a shallow curve, separated (five and six) by a poreless scale. Tail semi-verticillate with scales arranged in whorls with six dorsal and four ventral rows per verticil. Supracaudals rectangular with a rounded posterior margin; subcaudals squarish with rounded posterior margins. Two to three cloacal spurs at base of tail. Color. Pale grey to buff or pinkish brown above with 6���7 dark brown to blackish, very wavy and irregular crossbands extending from occiput to sacrum. Crossbands darkest posteriorly, incompletely edged with white and with a median white spot. Crown of head heavily spotted or mottled with dark brown; a forward pointing darker ���V��� extending along the canthus rostralis, whereas a dark streak runs from the nostril through the middle of the eye to above the ear, joining the occipital band. Limbs spotted with white and dark brown. Original tail banded with 10 regular dark brown to blackish crossbars edged with black posteriorly, each followed by a narrow band of white. Venter pale whitish pink; underside of tail mottled and spotted with white and brown. Variation. Paratypes and other specimens agree with the holotype in most features of scalation (Table 4). Nasorostrals separated by a single granular scale in four (of 75) specimens examined. Scales between nasals and eye 10���13, from eye to ear 15���22. Mental from as long to twice as long as broad; one postmental in 18 of 70 specimens. Supralabials 7���10. Infralabials 6���9. Postmentals usually two, but occasionally 1 or 3 (18 and 2 of 72 examined, respectively). Midbody scale rows 88���106. 0���11 enlarged inferomedian scales under the fourth toe. Precloacal pores in males 8���12 (mostly 9���11) often with a median gap; females lacking pores, although exhibiting a row of somewhat enlarged scales in this position. Original tails 50.0��� 55.5 % of total length. Supracaudal scales in 6���7 (rarely 8) rows per tail whorl. 44.7 % of specimens have regenerated tails (n= 76). Distribution. Isolated populations are found on the Soutpansberg, Blouberg and Matlala inselberg, Limpopo Province (Fig. 6). Natural history. The species is nocturnal. It frequents crevices and fissures between rocks as well as under flakes on Waterberg sandstone outcrops, although the Matlala population occurs on granites. May be solitary or can occur in small (presumably family) groups in Soutpansberg Mountain Bushveld (SVcb 21) and Soutpansberg Summit Sourveld (GM 28) (Mucina & Rutherford 2006) at elevations of 1000���1700 m a.s.l. Two eggs at a time are laid during midsummer.The eggs adhere to the roof of the crevice before hardening. Communal nesting has been observed in the Soutpansberg where more than 20 eggs have been found under a rock. Remarks. Afroedura broadleyi sp. nov. shares a suite of characters with A. langi and four other species described herein as new. In comparison with other members of this group it differs most obviously in male precloacal pore counts, being much lower than A. leoloensis sp. nov. and having non-overlapping pore ranges with A. langi, A. pienaari sp. nov. and A. granitica sp. nov. (more pores) and A. waterbergensis sp. nov. (fewer pores). In addition it is substantially larger than A. leoloensis sp. nov., A. langi, and A. waterbergensis sp. nov. (see Table 4). Three populations are known, each of which appears to differ slightly from the others. However, there is significant overlap in morphological features and Jacobsen���s (1990, 1992a) initial recognition of two putative species (���Soutpansberg��� and ���Matlala���) cannot be substantiated given available data. Unfortunately, molecular data were available only from Soutpansberg samples. The Soutpansberg form is separated from the others by A. pienaari along the Waterpoort Gap, whereas those from Matlala are separated by 70 km from the nearest other population (on the Blouberg). The Matlala and Blouberg forms have the precloacal pores in a continuous row, only exceptionally subdivided by a poreless scale, as is frequently the case in Soutpansberg populations. Precloacal pores on Matlala and in the Blouberg are mostly eight, nine or 10 and on the Soutpansberg nine, 11 or 12. Individuals from the eastern Blouberg tend to be smaller and have a mean midbody scale count of 91.46 + 2.37 (n = 13), which is considerably lower than that from Soutpansberg individuals��� 96.91 + 4.32 (n = 22), western Blouberg��� 97.73 + 3.49 (n = 15) and Matlala��� 99.5 + 4.19 (n = 18)., Published as part of Jacobsen, Niels H. G., Kuhn, Arianna L., Jackman, Todd R. & Bauer, Aaron M., 2014, A phylogenetic analysis of the southern African gecko genus Afroedura Loveridge (Squamata: Gekkonidae), with the description of nine new species from Limpopo and Mpumalanga provinces of South Africa, pp. 451-501 in Zootaxa 3846 (4) on pages 487-488, DOI: 10.11646/zootaxa.3846.4.1, http://zenodo.org/record/250495, {"references":["Jacobsen, N. H. G. (1992 a) Flat geckos (genus Afroedura) in the Transvaal. Journal of the Herpetological Association of Africa, 40, 22 - 25. http: // dx. doi. org / 10.1080 / 04416651.1992.9650313","Jacobsen, N. H. G. (1997) Family Gekkonidae, Genus Afroedura. In: Van Wyk, J. H. (Ed.), Proceedings of the FitzSimons Commemorative Symposium, South African Lizards: 50 years of Progress and Third H. A. A. Symposium on African Herpetology, Transvaal Museum, Pretoria, South Africa, 11 - 15 October 1993. Herpetological Association of Africa, Stellenbosch, pp. 34 - 39.","Kirchhof, S., Kramer, M., Linden, J. & Richter, K. (2010) The reptile species assemblage of the Soutpansberg (Limpopo Province, South Africa) and its characterisitcs. Salamandra, 46, 147 - 166.","Mucina, L. & Rutherford, M. C. (2006) The Vegetation of South Africa, Lesotho and Swaziland. Strelitzia, 19. South African National Biodiversity Institute, Pretoria, 807 pp."]}

Altigekko [see Altiphylax] Comments: Junior subjective synonym of Altiphylax fide Sindaco & Jeremčenko (2008).

Mediodactylus (14 species) Distribution: southern Italy, Balkan Peninsula, Crimea (Ukraine), Turkey, Cyprus, Lebanon, Israel, Syria, Iraq, Iran, Pakistan, Afghanistan, Kyrgyzstan, Tajikistan, Turkmenistan, Uzbekistan, southern Khazakhstan, southern Russia, northern Xinjiang (China) Composition: M. amictopholis (Hoofien), M. aspratilis (Anderson), M. brachykolon (Krysko, Rehman, & Auffenburg), M. dehakroensis (Masroor), M. heterocercus (Blanford) (M. h. mardinensis (Mertens)), M. heteropholis (Minton, Anderson & Anderson, 1970), M. ilamensis (Fathinia, Karamiani, Darvishia, Heidari & Rastegar-Pouyani), M. kotschyi (Steindachner), (M. k. adelphiensis (Beutler & Gruber), M. k.bartoni (��t��p��nek), M. k. beutleri (Baran & Gruber), M. k. bibroni (Beutler & Gruber), M. k. bolkarensis R��sler, M. k. buchholzi (Beutler & Gruber), M. k. ciliciensis (Baran & Gruber), M. k. colchicus (Nikolsky), M. k. concolor (Bedriaga), M. k. danilewskii (Strauch), M. k. fitzingeri (��t��p��nek), M. k. fuchsia (Beutler & Gruber), M. k. kalypsae (��t��p��nek), M. k. karabagi (Baran & Gruber), M. k. lycaonicus (Mertens), M. k. maculates (Bedriaga), M. k. oertzeni (Boettger), M. k. orientalis (��t��p��nek), M. k. ponticus (Baran & Gruber), M. k. rumelicus (M��ller), M. k. saronicus (Werner), M. k. schultzewestrumi (Beutler & Gruber), M. k. skopjensis (Karaman), M. k. solerii (Wettstein), M. k. steindachneri (��t��p��nek), M. k. stepaneki (Wettstein), M. k. syriacus (��t��p��nek), M. k. tinensis (Beutler & Fr��r), M. k. unicolor (Wettstein), M. k. wettsteini (��t��p��nek)), M. narynensis Jerjomtschenko, (Zarinenko & Panfilow), M. russowii (Strauch) (M. r. zarudnyi (Nikolsky)), M. sagittifer (Nikolsky), M. spinicauda (Strauch), M. stevenandersoni (Torki), M. walli (Ingoldby) Comments: We here include the four species included in the analyses of Macey et al. (2000) and this paper, as well as the three species assigned to Carinatogecko by Sindaco & Jeremčenko (2008) and Torki (2011). We follow Červenka & Kratochv��l (2010) in assigning Cyrtopodion dehakroense to Mediodactylus, and Khan (2003 c, 2006, 2009) in assigning walli to this genus. The inclusion of Cyrtopodion brachykolon to Mediodactylus is based on the results presented in this paper., Published as part of Bauer, Aaron M., Masroor, Rafaqat, Titus-Mcquillan, James, Heinicke, Matthew P., Daza, Juan D. & Jackman, Todd R., 2013, A preliminary phylogeny of the Palearctic naked-toed geckos (Reptilia: Squamata: Gekkonidae) with taxonomic implications, pp. 301-324 in Zootaxa 3599 (4) on page 319, DOI: 10.11646/zootaxa.3599.4.1, http://zenodo.org/record/217999

Tropiocolotes (9 species) Composition: T. algericus Loveridge, T. bisharicus Baha el Din, T. nattereri Steindachner, 1901, T. nubicus Baha el Din, T. scortecci Cherchi & Spano, T. somalicus Parker, T. steudneri (Peters), T. tripolitanus Peters (T. t. occidentalis Parker, T. t. apoklomax Papenfuss), T. wolfgangboehmei Wilms, Shobrak & Wagner Distribution: North Africa from Mauritania to northern Somalia to the Mediterranean, Arabian Peninsula, Israel, Jordan Comments: Species from Iran through Rajasthan previously assigned to this genus are now recognized as members of the distantly related Microgecko (Gamble et al. 2012; this study). Both Asiocolotes (here not regarded as valid) and Microgecko were previously considered subgenera of Tropiocolotes., Published as part of Bauer, Aaron M., Masroor, Rafaqat, Titus-Mcquillan, James, Heinicke, Matthew P., Daza, Juan D. & Jackman, Todd R., 2013, A preliminary phylogeny of the Palearctic naked-toed geckos (Reptilia: Squamata: Gekkonidae) with taxonomic implications, pp. 301-324 in Zootaxa 3599 (4) on page 320, DOI: 10.11646/zootaxa.3599.4.1, http://zenodo.org/record/217999

Crossobamon (2 species) Composition: C. eversmanni (Wiegmann), (C. e. lumsdeni (Boulenger)), C. orientalis (Blanford) Distribution: northwest India, Pakistan, eastern Iran, northern Afghanistan, Turkmenistan, Uzbekistan, Tajikistan, southern Kazakhstan Comments: The status of Stenodactylus lumsdeni Boulenger is unstable. Anderson (1999) considered this as a synonym of Bunopus tuberculatus, but we follow Szczerbak & Golubev (1986, 1996) in regarding it as related to Crossobamon eversmanni. These authors regarded it as a valid subspecies of C. eversmanni and treated Stenodactylus maynardi Smith as its junior synonym. Having not examined specimens, we withhold judgement on its status and simply follow Szczerbak & Golubev (1986, 1996) pending further investigation., Published as part of Bauer, Aaron M., Masroor, Rafaqat, Titus-Mcquillan, James, Heinicke, Matthew P., Daza, Juan D. & Jackman, Todd R., 2013, A preliminary phylogeny of the Palearctic naked-toed geckos (Reptilia: Squamata: Gekkonidae) with taxonomic implications, pp. 301-324 in Zootaxa 3599 (4) on page 318, DOI: 10.11646/zootaxa.3599.4.1, http://zenodo.org/record/217999

Siwaligekko [see Cyrtodactylus] Comments: We synonymize Siwaligekko with Cyrtodactylus based on our phylogenetic results and those of Wood et al. (2012). Siwaligekko is applicable at a subgeneric level to those species most closely related to C. battalensis; however, given the uncertainty of relationships among basal Cyrtodactylus (recovered as paraphyletic in this paper), we suggest that Siwaligekko be retained as a subgeneric name for all members of the genus occurring in the western Himalayan region. Khan���s (2003) inclusion of Geckoella in Siwaligekko is not supported, but this genus is also embedded within Cyrtodactylus sensu lato (Wood et al. 2012)., Published as part of Bauer, Aaron M., Masroor, Rafaqat, Titus-Mcquillan, James, Heinicke, Matthew P., Daza, Juan D. & Jackman, Todd R., 2013, A preliminary phylogeny of the Palearctic naked-toed geckos (Reptilia: Squamata: Gekkonidae) with taxonomic implications, pp. 301-324 in Zootaxa 3599 (4) on page 319, DOI: 10.11646/zootaxa.3599.4.1, http://zenodo.org/record/217999