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Species-level phylogeny and especially phylogeography of African chelonians is a comparatively under-studied field of research. We review the current knowledge of phylogeny and phylogeography, highlight congruence of spatial phylogeographic patterns amongst chelonians and other taxa and suggest future research directions to address gaps in knowledge. Our review shows that phylogeographic and phylogenetic investigations have led to unexpected findings. For example, for Pelomedusa, a putatively wide-ranging monotypic terrapin genus, cryptic diversity was revealed, with more than ten species being uncovered. The formerly recognized tortoise genus Homopus sensu lato was found to be paraphyletic with respect to Chersina. To resolve this situation, Homopus was restricted to the four-toed species H. areolatus and H. femoralis and the genus Chersobius was resurrected for the five-toed species C. boulengeri, C. signatus, and C. solus. Three previously recognized taxa were shown to be invalid, viz. the putatively extinct terrapin species Pelusios seychellensis and the tortoise subspecies Chersobius signatus cafer and Stigmochelys pardalis babcocki. Together with taxonomy, the knowledge of phylogeographic structuring sets a solid foundation for conservation measures and allows the identification of Management and Conservation Units. However, the current legislation, in particular the enforcement of the Nagoya Protocol under the Convention of Biological Diversity (CBD), has largely halted research on widely distributed taxa and turned the well-intended concept of Access and Benefit Sharing into a major impediment for conservation and research. The current situation leads for many species to a continued usage of outdated and incorrect taxonomic classifications resulting in an error cascade of conservation decisions. This is counterproductive to the aims of the CBD, that is, the protection of biodiversity. Sequencing historical DNA from museum specimens using aDNA approaches could be a short-term approach to mitigate, but not solve, this impediment.

Nucras broadleyi sp. nov. Angolan Sandveld Lizard urn:lsid:zoobank.org:act: C82E3A75-96FF-4D2A-9B52-3ABF4B58BC2B (Figures 4–6) Chersonymy. Nucras tessellata var. taeniolata (Bocage 1895: 30), Nucras tessellata var. taeniolata (Boulenger 1910: 474), Nucras tessellata var. holubi (Boulenger 1917: 210), Nucras intertexta var. holubi (Boulenger 1920: 20), Nucras tessellata (Monard 1937: 73; Laurent 1964: 56), Nucras ornata (Broadley 1965: 23), Nucras tessellata (Broadley 1972: 30; Ceríaco et al. 2016: 56; Burger 2014: 171), Nucras aff. tessellata (Marques et al. 2018: 221; Branch et al. 2019: 317). Type material. The type series is comprised of the three most recently collected specimens, which are housed in PEM and TM. Holotype. A subadult male (PEM R 24005, AG 018), 10 km west of Lola, edge of Bentiaba River valley, Namibe Province, Angola (-14.29028, 13.53056, WGS 84, 802 m asl). Collected by W. R. Branch, P. Vaz Pinto, and J.S. de Almeida on 2 November 2015. Paratypes (2). a) A subadult female (PEM R 24157, AG 166), 8.8 km southwest of Farm Mucungo, Namibe Province, Angola (-14.80167, 12.41917, WGS 84, 385 m asl). Collected by W. R. Branch, P. Vaz Pinto, and J. S. de Almeida on 8 November 2015. b) An unsexed adult (TM 40392), “ 34 km S of Moçâmedes to Porto Alexandre, Angola, 1512 Ca ” (= 34 km S Namibe to Tômbwa), Namibe Province, Angola (approx. -15.48220, 12.18289). Collected by W.D. Haacke on 30 March 1971. Additional referred material: The following additional material was used to expand the description of variation within the species: a) an adult male (MD 1967, Laurent 1964), “km 34 de la route de Moçâmedes à Sa da Bandeira” (= 34 km from Namibe on Lubango road, -15.03333, 12.41667), collected 24 October 1949, b) MBL 646, 647 a, 647b (Bocage 1895: 30) from Maconjo (approx. -15.01667, 13.20000), c) BM 1970.6.29.10–11 (Boulenger 1910: 474) from Ponang Kuma (= Donguena, approx. -17.01667, 14.71667), and d) MHNC 91.0524 (Monard 1937) from Capelongo (approx. -14.88333, 15.083333), collected April 1933. Etymology. The specific epithet is a patronym in honor of Donald G. Broadley for his numerous contributions to the herpetofauna of Africa. Don (as most of us knew him) was the first to recognize the Angolan population as a separate species (Broadley 1972). The name is constructed in the masculine genitive. Diagnosis. Assigned to Nucras due to a well-defined collar (absent in Ichnotropis), toes not serrated or fringed (versus serrated or fringed in Meroles), subdigital lamellae smooth (versus keeled in Pedioplanis and Heliobolus), subocular bordering lip, the nostril is pierced between two nasals, nasal well separated from upper labial, and dorsal scales small, smooth, and juxtaposed. * anterior subocular (posterior subocular), ** total (largest number, t = truncated) The new species can be diagnosed from other Nucras species based on a combination of the following characters: series of transversely enlarged plates present under forearm (versus absent or only feebly enlarged in Nucras lalandii), a small series (0–6) of small granules present between supraciliaries and supraoculars (versus mostly absent in N. boulengeri and N. lalandii), 23–29 lamellae under 4 th toe (versus less than 22 in N. lalandii), dorsum with a series of longitudinal pale stripes (versus dark cross bands present in N. lalandii and N. scalaris or a series of pale vertebral spots, sometimes forming irregular transverse bands in N. intertexta or lack of any dorsal patterns in N. aurantiaca), four pale stripes on nape with outer stripes forming a. Bloemfontein Museum Province Province Province Province Province Province Province Province Province National Cape Cape Cape Cape Cape Cape Cape NMB – , , Northern Western, Northern,, Northern Northern, Western,, Western Kamanjab, KwaZulu-Natal Cape , Western Museum Locality South Africa Africa South South Africa South Africa South Africa GenBank Africa South South Africa Namibia, South Africa Africa South Elizabeth Port – 1 005257 951543 951540 951542 951541 632229 871208 547691 871206 871207 547718 RAG PEM HG HF HF HF HF EF DQ HF DQ DQ HF, Science 4 005232 951535 951536 751398 547725 547732 547733 547759 of ND HG — HF — HF — FR HF HF HF HF Academy 005211 951559 951556 951558 951557 080358 871152 547772 871148 871149 547800 California S 16 HG HF HF HF HF AF DQ HF DQ DQ HF CAS – : number 206725 R 18745 R 16873 R 11574 R 16872 209602 R 18376 abbreviations Museum accession CAS PEM PEM NMB PEM NA NA NMNW CAS PEM Museum. accession 5584 08 069 - 20650 21061 20687 0531 08 6001 6067 049 analysis Field ID AMB KTH MB MB MB MH GW AMB AMB SVN genetic in Samples used Species tessellata tessellata tessellata tessellata tessellata longicaudata australis australis capensis capensis suborbitalis. ) continued 2 Table (Genus Nucras Nucras Nucras Nucras Nucras Outgroup Latastia Australolacerta Australolacerta Ichnotropis Ichnotropis Meroles continuous light stripe with the outer edges of the parietals (similar to Broadley’s (1972) N. tessellata tessellata var. “ T;” differs from N. livida and N. tessellata where the outer stripes often do not form a continuous light stripe with the outer edges of the parietals; differs from N. caeiscaudata and N. ornata where there are only three longitudinal stripes present on nape and sometimes the vertebral ones are absent), well defined occipital scale separating parietals (versus reduced or absent in northern Namibia N. holubi, which is referred to as N. intertexta damarana Parker; as well as absent in N. caesicaudata), parietal foramen absent (often present in all other species except N. taeniolata), and postnasals separated (usually fused in N. taeniolata). In the phylogenetic analysis, the uncorrected p -distances show that this clade differs by>8% for 16S,>14% for ND4, and>1% for RAG 1 sequence divergence from other members of the N. tessellata clade. Description of Holotype (Fig. 4). Body relatively slender (SVL approx. 4.5 times the head length, tail truncated), with hindlimbs larger than forelimbs (femur of hind limb equal to length of tibia); head narrow and elongated (56% longer than wide) with narrow pointed but blunt snout, that is slightly longer than distance from back of eye to rear of ear opening. Rostrum protruding and visible from below. Nasals paired and in contact (0.2 mm suture length), not swollen, nostril directed backwards separating postnasals. Frontonasal single, wider than long (1.1 × 1.8 mm). Prefrontals paired and in broad median contact with one another (0.6 mm suture length), wider than long (1.1 × 1.2 mm). Frontal entire, longer than wide (2.7 × 1.9 mm). Two large rounded supraoculars, both in contact with the frontal, with anterior supraocular preceded by a single large scale in contact with prefrontal, frontonasal, and posterior loreal, with posterior supraocular bordered by a single large scale in contact with parietal and frontoparietal. Paired frontoparietal in broad contact (1.3 mm suture length), nearly as wide as long (1.7 × 1.5 mm). Parietals twice as long as wide (3.1 × 1.8 mm), fully separate by a large, pentagonal interparietal (2.5 × 1.2 mm) that is twice as long as wide, slightly shorter than frontoparietals and nearly equal to length of frontonasal and prefrontal combined. Small subtriangular occipital (0.5 × 0.7 mm). Two loreals, second much larger than first. Six supraciliaries on each side, 1 st is the longest. A single minute granule scale between supraocular and supracilliares on right side, none on left side. Four supralabials anterior to subocular and three supralabials posterior to subocular, on both sides. Subocular slightly elevated medial and bordering the lip, its lower border being shorter than the upper. Three temporal scales, first longer than others, smooth. Tympanic shield as wide as long, border of ear opening. No ear lobes. Lower eyelid with transparent brille formed by five larger scales, surrounded by numerous smaller scales. Lower eyelid separated from subocular and enlarged temporal scales by a series of 10 smaller scales. Small scale above 3 rd supralabial separating the posterior loreal and subocular. Enlarged scale bordering 1 st post subocular, supralabial, and the subocular. Six infralabials on both sides, with 3 rd being longest; four enlarged pairs of chin shields, last largest and first three in broad contact. Twenty-four gular scales in a straight line between symphysis of chin shields and median collar plate, equal in size except last 4–5 larger. Collar free, comprising seven enlarged plates (median subtriangular) and extending slightly onto side of neck as a crease, bordered by 2–3 smaller scales. Dorsal scales small, juxtaposed, granular, smooth, larger on sides toward ventrals. Midbody scales 42. Ventral plates eight longitudinal and 28 transverse rows (from collar to groin), plates of the innermost rows longer than broad, with outer row notably smaller than other rows, transverse row of ventrals across chest just behind collar longer than broad; preanal scales irregular, median ones larger. Scales on upper surface of forearm large, smooth or slightly keeled. Scales on lower surface of forearm with eight enlarged plates, at least twice the width of scales on upper forearm. Scales on upper surface of tibia rhombic, subimbricate, smooth, and much larger than dorsal scales. Tibia below with a series of large plates. Subdigital lamellae under fourth toe 23 R / 25L. Femoral pores 13 R / 15L. Dorsal scales on tail oblique, strongly keeled diagonally, and truncate behind, ventral scales on tail obtusely keeled. Coloration. Dorsum with eight pale cream to white dorsolateral longitudinal stripes, separated by dark brown to black stripes. These stripes are more boldly patterned anteriorly, fading posteriorly. No light vertebral stripe. The two pale paravertebral stripes are separated by a very narrow strip of darker scales that starts on the interparietal through the occipital scale and fades posteriorly onto body and tail. The dorsolateral stripe extending along outer borders of parietals continues onto the tail. It is followed by the upper lateral stripe extending from posterior of the eye onto the head through the mid-temporal with a brief break above the ear opening, and continues onto the tail. The lower lateral stripe starts at the subocular, through the ear opening, broken briefly above the arm, after which it continues all the way onto the tail. Ventrum white and lower limbs oblique white. Fore limbs upper surface black with scattered pale blotches. Hind limbs light brown with pale blotches. Upper surface of tail red-brown, similar to hind limbs. Scales bordering the orbit are black edged. Variation (Figs. 5–6). Meristic and escalation data are summarized in Table 1. The largest specimen examined is (BM 1907.6.29.10) 74 + 144 mm (tail regenerated). Regarding coloration, there seem to be three main variations among material examined: 1) 8–9 longitudinal stripes as in holotype (in PEM R 24005, MBL 647a, 647b, MHNC 91.0524–5), 2) 4–5 pale longitudinal stripes broken up posteriorly with flanks spotted (in BM 1970.6.29.10–11, TM 40392, MD 1967), and 3) broken paravertebral stripes, continuous dorsolateral line and barred flanks (in PEM R 24157), similar to N. intertexta. Distribution. Found only in semi-arid south-western Angola, throughout much of Namibe Province and extending onto the escarpment of southern Hu í la and Cunene Provinces (Fig. 1). Known localities include: Maconjo (Bocage 1895: 30), Ponang Kuma (=Donguena) (Boulenger 1910: 472), 34 km from Namibe on Lubango road (Laurent 1964: 56), 34 km south of Tombwa (TM 40397), 8.8 km southwest of Farm Mucungo (this study), 10 km west of Lola (this study), and Capelongo (Monard 1937: 73). The locality of Caconda (Bocage 1895) extends the species distribution further north into Huíla Province, but the specimens could not be critically evaluated by Broadley (1972) and are now presumably lost. Habitat. The species appears to be associated with mopane woodlands, dry savannas, and semi-desert shrublands (Barbosa 1970). The new material was found in sandy plains with scattered low granite outcrops, with varying degrees of short grass cover and scattered bushes. Vegetation included Colophospermum mopane, Ficus sp., Senegalia (=Acacia) mellifera, Commiphora sp., Boscia foetida, and Salvadora persica. The confirmed historical records were also obtained within the dry woodland zone, even though the possible occurrence of the species in Caconda would place the species above 1,500 m asl and well into the mesic conditions of Brachystegia habitats (Barbosa 1970)., Published as part of Branch, William R., Conradie, Werner, Pinto, Pedro Vaz & Tolley, Krystal A., 2019, Another Angolan Namib endemic species: a new Nucras Gray, 1838 (Squamata: Lacertidae) from south-western Angola, pp. 82-95 in Amphibian & Reptile Conservation 13 (2) on pages 86-91, DOI: 10.5281/zenodo.3731977, {"references":["Bocage JVB. 1895. Herpetologie d'Angola et du Congo. Ministerio da Marinha e das Colonias, Lisbonne, Portugal. 203 p., 20 pls.","Boulenger GA. 1910. 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Proceedings of the California Academy of Sciences Series 4, 65 (Supplement II): 1 - 501.","Branch WR, Baptista N, Vaz Pinto P, Conradie W. 2019. The reptiles of Angola - History, updated checklists, endemism, hot spots, and future directions for research. Pp. 283 - 326 In: Biodiversity of Angola. Science and Conservation: A Modern Synthesis. Editors, Huntley BJ, Ferrand N, Russo V, Lages F. Springer Open, Cham, Switzerland. 552 p.","Barbosa LAG. 1970. Carta Fitogeografica de Angola. Instituto de Investigacao Cientifica de Angola, Luanda, Angola. 323 p."]}

Branch, William R., Conradie, Werner, Pinto, Pedro Vaz, Tolley, Krystal A. (2019): Another Angolan Namib endemic species: a new Nucras Gray, 1838 (Squamata: Lacertidae) from south-western Angola. Amphibian & Reptile Conservation 13 (2): 82-95, DOI: http://doi.org/10.5281/zenodo.3731977, {"references":["Aljanabi SM, Martinez I. 1997. Universal and rapid saltextraction of high quality genomic DNA for PCR based techniques. Nucleic Acids Research 25: 4,692- 4,693.","Arevalo E, Davis SK, Sites JJW. 1994. Mitochondrial DNA sequence divergence and phylogenetic relationships among eight chromosome races of the Sceloporus grammicus complex (Phrynosomatidae) in central Mexico. Systematic Biology 43: 387-418.","Barbosa LAG. 1970. Carta Fitogeografica de Angola. Instituto de Investigacao Cientifica de Angola, Luanda, Angola. 323 p.","Bauer AM, Childers JL, Broeckenhoven C, Mouton PLN. 2019. A new Nucras Gray, 1838 (Squamata: Lacertidae) from the Strandveld of the Western Cape, South Africa. 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Kinyongia mulyai sp. nov. Synonomy: Kinyongia adolfifriderici Tilbury 2010 Holotype. PEM-R 19199 (CT 426 — Fig. 11), a sub-adult male with partially everted hemipenes, collected by Colin Tilbury and Isak Hattingh on 21 st March 2010, from path side vegetation at 1700 metre a.s.l., Mount Nzawa, Moba District, Katanga Province, Democratic Republic of the Congo (6 ° 51.07 ’ S; 29 ° 35.87 ’ E). Diagnosis. Within the genera Kinyongia and Trioceros, the ontogenetic development of body crests and rostral process’s—whilst not developed to the same degree as seen in adults—is still evident in sub-adult specimens, and can be reliably used to determine the presence or absence of these characters. Although Kinyongia mulyai sp. nov. is known from only one sub-adult specimen, it is likely that the morphological characters are sufficiently well developed to provide reliable criteria on which to base a morphological determination. Kinyongia mulyai sp. nov. is distinguishable from the closely-related species K. adolfifriderici, K. gyrolepis (Greenbaum et al. 2012) and K. excubitor (Barbour 1911) by the absence of a dorsal crest in the male. It is also distinguished from K. adolfifriderici by having a narrower and more elongate head (Fig. 10), with a head length/inter-orbital width ratio of 4 (versus 3 for K. adolfifriderici), and from both K. gyrolepis and K. adolfifriderici by having a higher number of upper labial scales (17 vs. a maximum of 15 and 14 respectively; Greenbaum et al. 2012). Distinguished from all the other Kinyongia congeners by the absence of any form of naso-rostral ornamentation in males. Description of the Holotype. A small chameleon with a snout-vent length of 44mm and tail measuring 63mm. Head length from casque tip to snout 13.7mm and the inter-orbital width at mid orbit 2.8mm. Casque low, barely elevated above the nape. Parietal crest consists of a row of 3 enlarged tubercles at the apex of the casque. This forks anteriorly into two short rows each consisting of 2 low tubercles. Supra-orbital ridges more or less smooth. The supra-orbital ridges are separated from each other across the mid-orbital point by three flattened tubercles. The canthal ridge is adorned with a row of 4 prominent rounded tubercles—the most anterior the largest. A temporal crest composed of 4 similarly sized enlarged tubercles arises from the mid posterior orbital rim; this crest ascends along the posterior rim of the casque to its apex. The top of the casque and the zone below the temporal crest are covered with similar-sized, flattened tubercles. The nares open infero-posteriorly about midway between the anterior orbital rim and the snout. A single row of tubercles separates the inferior orbital rim and the upper labials. There is no rostral process. Seventeen upper labials and 17 lower labials are present from the tip of the snout to the posterior margin of the orbital rim. Two tubercles separate the upper labials from the end of the canthal ridge. There is no trace of a gular crest, or ventral crest. The sides of the body are clad with somewhat heterogeneous, flattened polygonal tubercles, with larger tubercles showing a tendency to rosette formation on the lower flanks. A network of thin interstitial grooves extends between the tubercles. There is no row of enlarged flank tubercles. The dorsal vertebral crest is represented by a single small cone at the nape behind the casque, followed by a second much smaller cone. Thereafter the dorsal keel is smooth. The tail is likewise smooth. The outer surfaces of the limbs are covered in enlarged flattened tubercles while the inner surfaces of the limbs are clad with small regular tubercles. Colour in Life. Head—Top of head olive green with lighter green tubercles on casque. Skin of eyeballs light brown with two thin horizontal stripes running through the middle and extending posteriorly along the temporal crest and anteriorly just below the canthal ridge. The zone below the temporal crest is powder blue scattered with greenish tubercles. A small triangular area between the mouth line, nares and the orbital rim is blue green. The gular region is suffused with pale orange and yellow extending posteriorly to between the front limbs. A pale line extends from below the eye to just beyond the commissure of the mouth. Background colouration of body is olive green. Interstitial skin between the tubercle rosettes on lower flanks forms a network of dark interstitium. The dark interstitium of the flanks is broken by two broad zones of pale interstitium extending from the dorsal ridge down over the flanks to the middle of the body. There is no white line on the abdomen. Outer side of limbs with light green tubercles, skin on inner sides of limbs off white. Tail olive green with many thin dark bands. Habitat. The Holotype was collected in closed canopy Afrotemperate montane forest at 1800 metres a.s.l. from path side vegetation, perched at approximately 2.5 metres from the ground. One other chameleon (not collected) was observed occupying a sleeping perch on an exposed liana vine over 20 metres above the ground. Etymology. Named for Mr. Jules Mulya, whose energetic support and assistance with smoothing away mountains of obstacles to the progress of the Mt. Nzawa expedition led to the discovery of both new species.

Rhampholeon (Rhinodigitum) hattinghi sp. nov. Synonymy: Rhampholeon boulengeri Tilbury 2010. Holotype. PEM-R 19194, an adult male with everted hemipenes, collected by Colin Tilbury and Isak Hattingh on 21 st March 2010, from path side vegetation at 1700 metres a.s.l., Mount Nzawa, Moba District, Katanga Province, Democratic Republic of the Congo (6 �� 51.07 ��� S; 29 �� 35.87 ��� E). Paratypes. PEM-R 19193, adult male; PEM-R 19195, adult male, PEM-R 19196, adult female; PEM-R 19197, adult female; PEM-R 19198, adult female; all collected on the same date and locality as the holotype. Diagnosis. Due to its markedly bicuspid claws, amelanotic parietal peritoneum and acalyculate bag-like hemipenes with dual apical horns, this taxon can be placed within the sub-genus Rhampholeon (Rhinodigitum) Matthee et al. 2004. In spite of the remarkably conserved external morphology within this genus, this new taxon is easily distinguishable from congeners by virtue of the following identifying characteristics: The consistent absence of deep mite pockets (or pits) in the inguinal flexure distinguishes this taxon from Rh. beraduccii Mariaux & Tilbury and the six species of the Rh. platyceps G��nther complex (Branch et al. 2014), including Rh. platyceps, Rh. chapmanorum Tilbury, Rh. maspictus Branch et al., Rh. tilburyi Branch et al., Rh. bruessoworum Branch et al. and Rh. nebulauctor Branch et al.; the presence of deep pits in the axillae of this taxon distinguishes it from Rh. nchisiensis Loveridge and Rh. acuminatus Mariaux & Tilbury; the relatively smooth supra-orbital and canthal crests distinguish this taxon from Rh. boulengeri, Rh. uluguruensis Tilbury & Emmrich and Rh. moyeri Menegon et al., in all of which a peaked cluster of tubercles forms distinct protuberances above the eye and the nasal aperture (Fig. 5). Description of Holotype. Snout-vent 56mm, tail 11mm. Body habitus leaf like - typical of all other Rhampholeon (Rhinodigitum) species. Head short, casque flattened, top of head shallowly concave. The lateral crests are studded with several prominent tubercles. Parietal crest indistinct, indicated by a short row of 3 marginally enlarged tubercles. The supra-optic ridge gathers into a low cluster of tubercles anteriorly above each eye, but without forming a supra-optic horn. The two supra-orbital ridges are connected to each other by a series of 18 inter-orbital tubercles arranged in a shallow V across the top of the head. The canthal ridges are formed by a row of enlarged, relatively smooth tubercles which terminate anteriorly at the base of a small, finely tuberculated stump-like process that barely projects off the anterior end of the snout. The nares open posteriorly, from a small bulge at a point roughly one third of the distance between the anterior orbital rim and the front of the maxilla. A distinct temporal crest arises from the mid post-orbital rim and consists of 5 tubercles on the right and six tubercles on the left, of which the most posterior is the largest. Three large sub-conical tubercles are spaced along the inferoposterior rim of the orbit. A prominent tubercle is situated just above the mouth in line with the posterior orbital rim. The dorsal vertebral line is weakly crenulated, fading to smooth over the pelvic region. The dorsal margin of the tail is likewise smooth. Deep axillary mite pockets (or pits) are present, whilst the inguinal flexures are smooth. The background scalation of the body, tail, limbs and belly is composed of tightly packed sub-homogeneous, stellate-edged tubercles. The gular region of the throat extending to between the front limbs is studded with many small conical tubercles. These also are found on the belly, underside of the tail and the underside and particularly on the outer-surface of the limbs. The sides of the body have regularly-spaced, enlarged conical tubercles, with three particularly enlarged cones situated in a row at midbody, the first sited above the shoulder, the second at midbody, and the last just before the pelvis. Claws are markedly biscuspid, several prominent palmar and plantar accessory spines are seen at the base of the toes, and the outer soles of the feet are relatively spinose whilst the inner soles are smooth with a cobblestone appearance. Colour in life. Adult male (Figs 6 & 7). Top of head and snout dark brown extending to cover the sides of the snout above the line of the mouth and to a lesser extent the eyeball and superior temporal zone. Eyeball with dark radiations especially prominent over the upper half of the eyeball. Gular region off-white. Body light brown mottled with olive green. Five dark blotches are spaced along the dorsal vertebral ridge, and a broad dark blotch is also located over the dorsum of the tail. Two thin dark green stripes, originating from the second and third vertebral blotches, extend infero-posteriorly over the flanks. Three dark spots are arrayed along the flank, each spot centred over a prominent conical tubercle. The inner sides of the fore and hind limbs pale orange. A prominent white spot is located on each side of the tail. Adult female stress pattern (Fig. 8). Dorsal ridge orange from the tip of the casque to the tip of the tail. Flanks speckled with small orange spots and three large white spots on the flanks, each surround an enlarged conical tubercle. Sides of head and gular region flecked in white, sometimes forming short vertical stripes. Eyeball dark and speckled with light blue tubercles. Lateral flank stripes dark but heavily flecked with light blue. Hemipenes. Short bag-like, truncus acalyculate; a pair of short, broad-based apical horns incurving above the sulcus spermaticus (Fig. 9). The upper surface of the horns is covered in closely packed thorn-like papillae, much like a pineapple skin. Fourteen papillae are arranged in a proximal or basal cluster terminating in a distal row of two papillae. Inferior to each apical horn, a prominent balloon-like membranous evagination billows anteriorly. Sexual Dimorphism. Apart from the prominent hemipenal bulges there is little sexual dimorphism evident in the type series (Table 4). In males the tail comprises between 16���23 % of the total length whilst in females it is between 14���18 % of total length. Number Sex Length Rp Casque/snout Sub-orbital Axil pit Inguinal pit (mm) (mm) rows *= Holotype, measurements in millimetres; s-v = snout-vent length; Io tub = inter-orbital tubercles; Io diam = interorbital diameter; Length Rp = length of rostral process; Sub-orbital rows = number of rows of tubercles between the inferior orbital rim and the upper labials; na = not measured. Variation. In the paratypes the parietal crest was either absent, or merely indicated as in the holotype. The supra-orbital crest is relatively smooth in all specimens, only forming a low cluster of tubercles in 2 specimens at the point where the inter-orbital tubercles terminate above the eye. The soles of the feet may vary from relatively smooth to a sub-acuminate field of tubercles. The hemipenes of PEM-R 19193 and PEM-R 19195 do not differ in any substantial way from that of the holotype. Reproduction. The parietal peritoneum in the abdominal cavity of the largest female (PEM-R 19198) was unpigmented and the fallopian tubes each held a single egg measuring 6.4mm x 5.6mm. Habitat. All specimens were found in closed canopy Afrotemperate montane forest on low vegetation alongside a path. Perch heights varied from a few centimetres up to 50 cm from the ground. Etymology. Named for Isak Hattingh, who enthusiastically but unwittingly agreed to accompany the first author on a trip which tested many boundaries. He never complained once., Published as part of Tilbury, Colin R. & Tolley, Krystal A., 2015, Contributions to the herpetofauna of the Albertine Rift: Two new species of chameleon (Sauria: Chamaeleonidae) from an isolated montane forest, south eastern Democratic Republic of Congo, pp. 345-364 in Zootaxa 3905 (3) on pages 355-359, DOI: 10.11646/zootaxa.3905.3.2, http://zenodo.org/record/240513, {"references":["Tilbury, C. R. (2010) The Chameleons of Africa, an Atlas including the chameleons of Europe, the Middle East and Asia. Edition Chimaira, Germany, 831 pp.","Matthee, C. A., Tilbury, C. R. & Townsend, T. (2004) A phylogenetic review of the African leaf chameleons: genus Rhampholeon (Chamaeleonidae): the role of vicariance and climate change in speciation. Proceedings of the Royal Society of London Series B, Biological Sciences, 271, 1967 - 1976. http: // dx. doi. org / 10.1098 / rspb. 2004.2806","Branch, W. R., Bayliss, J. & Tolley, K. A. (2014) Pygmy chameleons of the Rhampholeon platyceps complex (Squamata: Chamaeleonidae): Description of four new species from isolated ' sky islands' of northern Mozambique. Zootaxa, 3814 (1), 1 - 36. http: // dx. doi. org / 10.11646 / zootaxa. 3814.1.1"]}

Tilbury, Colin R., Tolley, Krystal A. (2015): Contributions to the herpetofauna of the Albertine Rift: Two new species of chameleon (Sauria: Chamaeleonidae) from an isolated montane forest, south eastern Democratic Republic of Congo. Zootaxa 3905 (3): 345-364, DOI: http://dx.doi.org/10.11646/zootaxa.3905.3.2

Rhampholeon (Rhinodigitum) chapmanorum Tilbury, 1992 Chapmans��� Pygmy Chameleon Synonomy: Rhampholeon chapmani Tilbury 1992 Rhampholeon chapmanorum Klaver & B��hme 1997 Rhampholeon (Rhinodigitum) chapmanorum Matthee, Tilbury & Townsend 2004. Holotype: Adult male (BMNH 1988.399), collected by Colin and Sarah Tilbury. Type locality: in evergreen forest at 940 m asl., above the Chididi Police post, on top of Malawi Hills, southern Malawi. Variation in adult coloration in an adult male and an adult female are illustrated in Fig. 4 D and Fig. 4 E, respectively. Note also the pronounced nasal and supraorbital processes on the head of the male. For full details of morphology, hemipenial structure, biology and habitat see Tilbury (2010)., Published as part of Branch, William R., Bayliss, Julian & Tolley, Krystal A., 2014, Pygmy chameleons of the Rhampholeon platyceps compex (Squamata: Chamaeleonidae): Description of four new species from isolated ' sky islands' of northern Mozambique, pp. 1-36 in Zootaxa 3814 (1) on page 17, DOI: 10.11646/zootaxa.3814.1.1, http://zenodo.org/record/286211, {"references":["Tilbury, C. R. (1992) A new dwarf forest chameleon (Sauria: Rhampholeon) from Malawi, Central Africa. Tropical Zoology, 5, 1 - 9. http: // dx. doi. org / 10.1080 / 03946975.1992.10539176","Klaver, C. J. J. & Bohme, W. (1997) Liste der rezenten Amphibien und Reptilien-Chamaeleonidae. Das Tierreich, 112, i - xiv, 1 - 85.","Matthee, C. A., Tilbury, C. R. & Townsend, T. (2004) A phylogenetic review of the African pygmy chameleons: genus Rhampholeon (Chamaeleonidae): the role of vicariance and climate change in speciation. Proceedings of the Royal Society of London, Series B, 271, 1967 - 1976.","Tilbury, C. R. (2010) Chameleons of Africa. An Atlas including the Chameleons of Europe, the Middle East and Asia. Edition Chimaira, Frankfurt am Main, 831 pp."]}

Rhampholeon (Rhinodigitum) bruessoworum sp. nov. Mount Inago Pygmy Chameleon Synonymy: Rhampholeon sp. Bayliss et al. 2010, p 17. fig. 13. Etymology. The specific epithet honours the contributions of the brothers Carl and Darren Bruessow to the protection of wildlife in southern Malawi, particularly via the Mount Mulanje Conservation Trust. Types. The type series comprises three specimens, including: Holotype.- An adult female (PEM R 20375; Fig. 9 A) collected by J. Bayliss, 5 September 2009, in a small patch of wet forest at the base of a granite inselberg of Mt. Inago, Zambézia Province, Mozambique (15 ˚04' 51 ”S 37 ˚ 23 ' 37 ”E, ca 1478 m a.s.l.). Allotype. An adult male (PEM R 20376, Fig. 9 B), same collecting details as holotype. Paratype. An adult female (PEM R 20374), same collecting details as holotype. Meristics. Measurements of the type series of Rhampholeon bruessoworum sp. nov. are summarized in Table 9. Diagnosis. The Mt. Inago Pygmy Chameleon is referable to Rhampholeon (subgenus Rhinodigitum) by possessing an unpigmented parietal peritoneum, claws that are strongly bicuspid, smooth plantar surfaces, and a rostral process. It can be distinguished from most other species in Rhampholeon (Rhinodigitum) by having deep inguinal (absent or indistinct in Rh. boulengeri, Rh. nchisiensis, Rh. uluguruensis, and Rh. moyeri) and axillary pits (also absent in Rh. nchisiensis). It differs from Rh. platyceps and Rh. maspictus sp. nov. in its small size (Rh. chapmanorum in having a relatively large rostral process in males (small in both sexes in Rh. chapmanorum), and from all other members of the Rh. platyceps complex in Mozambique (i.e. Rh. maspictus sp. nov., Rh. nebulauctor sp. nov. and Rh. tilburyi sp. nov.) in having a relatively longer tail in both sexes. From all other Rhampholeon it is also genetically well differentiated, and all chameleons examined form a monophyletic clade. Description of Holotype. Adult female, viscera exposed by a single ventral incision. Head: Dorsum of head flattened, with no upward flexure of the snout; casque flat, edged with weakly-defined lateral crests that are mainly restricted to the posterior region of the casque; temporal crest weakly-developed, comprising a single, interrupted row of large tubercles; parietal crest almost absent, composed of a few enlarged tubercles in the mid-line; supraorbital ridges reduced to a few scattered enlarged scales but with a very small multiscaled process forming a ‘soft horn’ at each end of the inter-orbital ridge that passes across the crown and is composed of 10 small granular tubercles, and that demarcate the posterior edge of a slight frontal depression; inferior orbital rim with 4 (right) and 5 (left) enlarged tubercles; snout bordered on each side by moderately developed rostral crests, that fuse together at the tip of the snout which is adorned with a very small, flattened rostral process (1.6 mm) which underneath is only slightly free from the rostral, and is four small tubercular scales long and four tubercular scales wide at its base; nares opening posterio-ventrally; no gular or mental appendages; scales on throat homogenous, more conical but smaller than those on crown of head and subequal in size to those on the belly. Body: Dorsal crest very weakly developed, reduced to 9 crenulations of enlarged, but not obviously spinose scales; crenulations most strongly developed over mid-body, reduced in size over on neck and which are present on the tail in more reduced form; deep axillary and inguinal pits are present; flank scalation heterogeneous, composed of small, stellate granules with few scattered, enlarged spinose tubercles, the largest at the shoulder and in two clusters; chest, belly and lower surface of tail smooth; limb scalation more tubercular, with a few enlarged, spinose tubercles on the forearms; claws strongly bicuspid; accessory planter spines on the soles of the fore and hind feet are present, but reduced to very small, soft, spinose scales at the base of the claws; tail flattened laterally, flexing downward on the distal third. Colour in life (based on two images of holotype, JB; Fig. 9 A): Mid-body mottled brown with two narrow, oblique purple-brown lateral stripes; dorsal surface of fore-body, neck, top of head and upper surfaces of limbs darker brown; Throat lightly mottled cream extending onto chin and labials, which have a light yellow flush; belly and base of tail pale brown. Colour in preservative: Body mottled brown with two narrow oblique reddish-brown bars on mid-flanks; lower surface of neck, belly, base of tail, soles of feet, and lower limbs pale brown. Description of Allotype (as for holotype, unless noted): Adult male, incision along base of tail with hemipenal muscle removed; hemipenes not everted. Head: Parietal crest almost absent, composed of a four enlarged tubercles in the mid-line; five enlarged tubercles on both sides of the inferior orbital rim; rostral process six small tubercular scales long and four tubercles wide at base. Body: Dorsal crest very weakly developed, reduced to nine crenulations of enlarged but not obviously spinose scales, subequal in development to those of the female holotype; tail relatively long (30.5 % SVL), with a prominent hemipeneal bulge. Colour in life (based on two images of allotype; Fig. 9 B): Mid-body mottled brown with small orange brown blotches along dorsal crest, separated by 2–3 crenulations; throat, belly and lower surfaces of limbs and tail pale brown. Colour in preservative: After preservation body pale brown with vestiges of two narrow oblique stripes on the flanks; pale brown below. Paratype variation (as for holotype, unless noted): Adult female with a large ventral incision. The ‘soft horn’ on the supraorbital ridge is very small and hardly protrudes; only four enlarged tubercles on inferior orbital rim; plantar spines very reduced to small, soft, spinose scales. Size. Presumably a small species, as all three specimens in the type series were sexually mature, with turgid testes or developing ova. Largest male - PEM R 20376 (allotype) 39.0 + 17.1 = 56.1 mm; largest female—PEM R 20374 (holotype) 47.5 + 15.3 = 62.7 mm. Distribution. Restricted to the type locality; Mt. Inago, Zambézia Province, northern Mozambique. Habitat. The Inago Massif shows habitat zonation, and is surrounded by Brachystegia woodland at the base. The type series were collected at night in mid-altitude (~ 1500 m) evergreen forest at the base of a granite inselberg (Fig. 9 C). This forest type comprised relatively large trees between 20-30 m high, with the upper canopy layer composed of species such as Drypetes natalensis, Schefflera umbellifera and Newtonia buchananii, whilst the midcanopy layer (Chrysophyllum gorungosum, Myrianthus holstii, bridelia sp. and Garcinia sp. These remnant forest patches are greatly reduced in size (et al. 2010).

Rhampholeon tilburyi sp. nov. Mount Namuli Pygmy Chameleon Synonomy: Rhampholeon platyceps Branch & Ryan 2001, p 282. Rhampholeon tilburyi Timberlake et al. 2010, p 57 nomen nudum Rhampholeon nov. sp. (tilburyi) Timberlake et al. 2010, p 61 nomen nudum Rhampholeon platyceps (tilburyi) Timberlake et al. 2010, p 61 nomen nudum Rhampholeon Portik et al. 2013, p 416. Etymology. The specific epithet is a patronym for our good friend Dr. Colin Tilbury for his outstanding contributions to knowledge of African chameleons, most recently exemplified in his magnum opus, the Chameleons of Africa (Tilbury 2010). For the last 30 years his travels throughout Africa in search of chameleons, combined with his stunning photography, have been a source of inspiration. His studies have done much to enhance our understanding of African chameleons, and it is with great pleasure that we name this chameleon in his honour. Types. The type series comprises eight specimens, including: Holotype. An adult female (PEM R 14921; Fig. 8 D) collected by a local guide, 30 November 1998, in the Ukalini Forest that nestles under the south face of the main Namuli peak, Namuli Massif, Zamb��zia Province, Mozambique (15 ˚ 22 'S 37 ˚04'E, ca 1550 m a.s.l.). Allotype. An adult male (PEM R 17132, Fig. 8 A) with everted hemipenes, collected by K.A. Tolley and S. van Noort, 26 May 2006, at night perched on a dead branch 50 cm above ground in a patch of forest dominated by overgrown tea (Camellia sinensis) on the SDZ Cha Sarl Tea Estate at Guru��, Namuli Massif, Zamb��zia Province, Mozambique (15 ˚ 26 ' 51.6 ���S, 37 ˚00���, 32.6 ���E, ca 839 m a.s.l.). Paratypes. Six specimens, comprising two females (PEM R 17134, Fig. 8 B, same collecting details as allotype; PEM R 17135, perched on a tea bush in an overgrown patch of tea, on SDZ Cha Sarl Tea Estate at Guru��, 15 ˚ 26 ' 42.8 ���S, 37 ˚00���, 19.2 ���E); three males (PEM R 17131, SDZ Cha Sarl Tea Estate at Guru��, 15 ˚ 26 ' 49 ���S, 37 ˚00���, 29 ���E; PEM R 17133, same collecting details as allotype; PEM R 20372, Muretha Plateau, Namuli Massif, Zamb��zia Province, Mozambique (15 ˚ 23 ' 26 ���S, 37 ˚02���,03���E, ca 1804 m a.s.l.), collected by J. Bayliss 27 May 2007; and a very small juvenile (PEM R 20373, same details as previous specimen). Meristics. Measurements for the type series of Rhampholeon tilburyi sp. nov. are summarized in Table 8. Diagnosis. The Mt. Namuli pygmy chameleon is referable to the Rhampholeon (subgenus Rhinodigitum Matthee et al. 2004) by possessing a short hemipenis that is almost bag-like, acalyculate and adorned with a pair of simple apical ���horns��� with a variable number of thorn-like papillae arranged on the outer aspect of the horn; having an unpigmented parietal peritoneum, claws that are strongly bicuspid, smooth plantar surfaces, a rostral process, and short tail (Rhampholeon (Rhinodigitum) by having deep inguinal (absent or indistinct in Rh. boulengeri, Rh. nchisiensis, Rh. uluguruensis, and Rh. moyeri) and axillary pits (also absent in Rh. nchisiensis). It differs from Rh. platyceps and Rh. maspictus sp. nov. by its smaller size (Rh. platyceps complex, including the populations from Mt Mabu, Mt Chiperone and Mt Inago, by retaining in adult males a prominent flexure (> 32 ˚) of the snout in front of the orbit (flat or Rh. platyceps and Rh. maspictus sp. nov.; Rh. nebulauctor sp. nov. and Rh. bruessoworum sp. nov.), and a narrower head (HW/HL% 49.4 %; 53.2���59.8 % in all other species). Finally, the species is also genetically well differentiated from all other Rhampholeon, and all specimens examined form a monophyletic clade from Mt. Namuli. Description of Holotype. Adult female, viscera exposed by a single ventral incision and two lateral incisions on left flank. Head: Flattened, with only a small upward flexure of the snout at the front of the orbit (angling upwards at approximately 28 ˚ from a line between tip of snout and top of casque); casque weak, flat with small, conical tubercular edges; temporal crest moderate, forming a straight ridge of 7 raised, rounded tubercles, 3 rd and last largest; lateral crests weak, composed of small conical tubercles that demarcate the edge of the casque; temporal crest moderate with a single row of interrupted large tubercles; parietal crest almost absent, composed of a few enlarged tubercles in the mid-line; supraorbital ridge well marked but without obvious clusters of tubercles (vaguely present at the posterior edge), and with a very small ���soft horn��� of enlarged tubercles at the junction with the shallow interorbital ridge; interorbital ridge composed of 12 small granular tubercles that demarcate the posterior edge of a prominent frontal depression; two enlarged tubercles on inferior orbital rim; rostral ridges very well marked, and fusing at tip of the snout which bears a rostral process (1.5 mm, in fixation flattened against rostrum), about 7 small granules long and 5-6 granules wide at base; nares opening posterio-ventrally; no gular or mental appendages; scales on throat homogenous, more stellate than those on crown of head and subequal in size to those on the belly. Body: Dorsal crest weak, composed of a double row of enlarged, rounded tubercles along the backbone; the crest undulates between 13 equally spaced clusters of enlarged, spinose tubercles that are largest on the middle of the body, flatter and smooth over the forelimb and sacral regions, and only vaguely visible on the tail; deep axillary and inguinal pits are present; flank scalation heterogenous, composed of small, stellate granules with scattered, enlarged spinose tubercles, the largest occurring over the shoulder; chest, belly and lower surface of tail smooth; limb scalation more irregular, with numerous larger, spinose tubercles on forearms; soles of feet sub-spinose; claws strongly bicuspid; accessory planter spines at base of claws very small, almost absent; tail gently tapering, dorsoventrally flattened, 21.7 % of total length, and covered with homogenous conical tubercles. Colour in preservative: Body mottled brown with reddish brown reticulation on flanks; head darker above, blotched with dark brown; belly and inner surfaces of limbs paler with darker specking; tail dark brown above and below. Description of Allotype (as for holotype, unless noted). Adult male, with everted hemipenes and viscera exposed by a single, ragged ventral incision and liver tissue excised for DNA analysis. Head: A very prominent angular flexure of the snout at the front of the orbit (approx 40 ˚ from line between tip of snout and casque); supraorbital ridge well marked with clusters of tubercles at posterior and anterior level of orbit, the latter forming a ���soft horn��� two scales high; rostral ridge fusing anteriorly with an obvious rostral process (1.3 mm), that is rounded in profile and composed of about 6 small granules long and 5-6 granules wide at base. Body: Claws strongly bicuspid; accessory planter spines at the base of claws weakly developed; tail 24.0% of total length. Coloration in life (Fig. 8 A): Head, body and tail tan to brown with orange flush along midflank and greenish flush over back and outer surfaces of hind limbs; two prominent diagonal lines on flanks, formed from three darkish spots joined by grey infusion; belly, throat, inner surfaces of limbs and underside of tail light tan; outer skin of limbs darker brown; snout with wide orange band running diagonally through nostril; skin of orbit and temporal region darker brown with scattered, intense green-blue individual tubercles; iris orange-red. Coloration after preservation: Body very dark, almost blackish; all normal coloration lost due to fixation in concentrated formalin. Paratype variation. The tail varies from 18.3���21.7 % of total length in females, and from 24.0��� 25.8 % in males; the rostral process varies from 7.9���10.4 % of head length in females, and from 15.9���16.6 % in males. In life the iris of a male paratype (R 17131) was reddish and the snout had an indistinct brown stripe rather than the orange band in the allotype. In addition, a few scattered green-blue tubercles extended onto the snout, as well as on to the orbit and temporal region. Prominent ���leaf vein��� dark lines on the flank were also well developed. Size. Largest male���PEM R 17132 (allotype) 39.6 + 12.5 = 52.1 mm; largest female���PEM R 14921 (holotype) 55.3 + 15.4 = 70.7 mm. The smallest male (SVL 33.2 mm) still has well-developed hemipenes, indicating that it is sexually mature (although testicular activity was not determined). Sexual Dimorphism. Whilst the type series is limited in number, there are indications of sexual dimorphism. Females grow considerably larger than males, and the three females in the type series have an average SVL of 51.67 mm, compared with only 34.67 mm for the three males. In addition, males have a more angular head shape (slope 39-45 ˚, Fig 9 B) than females (slope 22-28 ˚), and also proportionately longer tails and a longer rostral process (see above). However, the small number of specimens precludes statistical confirmation. More subtle difference in scalation details cannot be assessed until larger series are available. Hemipenis. (based on everted hemipenes of all three males in type series). Hemipenis short with unadorned basal and apical sections, except for a pair of curved horns that emerge prominently from the expanded, flattened crown, and which bear well developed spines on their upper surfaces; the folds of the sulcus spermaticus flare to drain into the flattened apical region. Distribution. Restricted to the type locality; Mt. Namuli, Zamb��zia Province, northern Mozambique, in both the evergreen forests on Manho and Ukalini forests on the Muretha Plateau (1804 m a.s.l.) at the base of the main peaks, respectively, and also in fallow areas of tea plantations at lower elevations (838 m a.s.l). Habitat. The Namuli Massif shows habitat zonation and is surrounded by Brachystegia woodland at the base. Grassland and scrub with forested river valleys cover the slopes, with grassy plateaus and patches of dense moist evergreen forest near the summit (Fig. 8 E). The largest surviving blocks include the Manho Forest (c. 1,000���1,100 ha) and the Ukalini Forest (c. 100 ha), the latter lodged against the base of the Namuli dome. Dominant emergent trees include Cryptocarya liebertiana, Faurea wentzeliana and Olea capensis (Timberlake et al. 2009). These forests are under high anthropogenic threat as they are cleared for potato cultivation by surrounding rural communities. The allotype and most paratypes were collected at night on perches from 30-50 cm above ground in forest adjacent to a mountain stream in a fallow areas of an operational tea estate that have overgrown into a forest. Portik et al. (2013) noted that adults and juveniles were found sheltering on branches of small trees or on the fronds of epiphytic ferns within Ukalini Forest. This contrasts with perches heights of 1.3���5 m recorded for Rh. chapmanorum (Tilbury 1992). Reproduction. The female holotype was gravid with seven almost spherical eggs (largest diameter 4.1-4.5 mm, four in the left oviduct and three in the right oviduct) that lack obvious signs of embryonic development. Diet. The stomach of the holotype contained insect fragments, including coleopteran elytra. Predation. A small chameleon, possibly referable to this species, was seen being carried by a Crowned Hornbill (Tockus alboterminatus) that was flying over the canopy of Ukalini Forest, 29 November 1998 (Branch & Ryan 2001)., Published as part of Branch, William R., Bayliss, Julian & Tolley, Krystal A., 2014, Pygmy chameleons of the Rhampholeon platyceps compex (Squamata: Chamaeleonidae): Description of four new species from isolated ' sky islands' of northern Mozambique, pp. 1-36 in Zootaxa 3814 (1) on pages 25-28, DOI: 10.11646/zootaxa.3814.1.1, http://zenodo.org/record/286211, {"references":["Branch, W. R. & Ryan, P. G. (2001) Additions to the Mozambique Herpetofauna: Two new lizards from the Namuli Massif, Mozambique. Herpetological Review, 32 (4), 281 - 282.","Bayliss, J., Monteiro, J., Fishpool, L., Congdon, C., Bampton, I., Bruessow, C., Matimele, H., Banze, A. & Timberlake, J. R. (2010) Biodiversity and Conservation of Mount Inago, Mozambique. Report produced under Darwin Initiative Award 15 / 0 36. Mulanje Mountain Conservation Trust, Malawi, 32 pp.","Portik, D. M., Travers, S., Bauer, A. M. & Branch, W. R. (2013 b) A new species of Lygodactylus (Squamata: Gekkonidae) endemic to Mount Namuli, an isolated ' sky' island of northern Mozambique. Zootaxa, 3710 (5), 415 - 435. http: // dx. doi. org / 10.11646 / zootaxa. 3710.5.2","Tilbury, C. R. (2010) Chameleons of Africa. An Atlas including the Chameleons of Europe, the Middle East and Asia. Edition Chimaira, Frankfurt am Main, 831 pp.","Timberlake, J. R., F. Dowsett-Lemaire, F., Bayliss, J., Alves, T., Baena, S., Bento, C., Cook, K., Francisco, J., Harris, T., Smith, P. & de Sousa, C. (2009) Mt. Namuli, Mozambique: Biodiversity and Conservation. Report produced under the Darwin Initiative Award 15 / 036. Royal Botanic Gardens, Kew, London, 114 pp.","Tilbury, C. R. (1992) A new dwarf forest chameleon (Sauria: Rhampholeon) from Malawi, Central Africa. Tropical Zoology, 5, 1 - 9. http: // dx. doi. org / 10.1080 / 03946975.1992.10539176"]}

Rhampholeon (Rhinodigitum) maspictus sp. nov. Mount Mabu Pygmy Chameleon Synonomy: Rhampholeon (Rhinodigitum) chapmanorum Tilbury 2010, p 179. Rhampholeon sp. Timberlake et al. 2012, p 45; Bayliss et al. 2014, p 179. Etymology. The specific epithet derives from a combination of mas (L. = man) and pictus (L. = painted), alluding to the unusual bright colours of most males, which are often retained for long periods, even when sleeping at night. Types. The type series comprises ten specimens all preserved in 50 % propanol (except for one hatchling in 90 % ethanol), including: Holotype. An adult male (PEM R 18072; Fig. 5 A, 6 A) collected by a W.R. Branch, J. Bayliss & W. Conradie, 27 May 2009, in the vicinity of the main forest base camp, Mt. Mabu, Zamb��zia Province, Mozambique (16 �� 17 ' 10.1 "S, 36 �� 24 '02.2"E; 967 m a.s.l.). Allotype. An adult female with a small ventral incision (PEM R 18061, Fig. 5 B, 5 A), same collecting details as holotype. Paratypes. Eight specimens, comprising four males (PEM R18059, 18073 (Fig. 6 B), 18074 - 75), same collecting details as holotype; three females (PEM R 18069 -70, 18076), same collecting details as holotype; and a hatchling (PEM R 18068, Fig. 6 C) Additional material (used in analysis but not forming part of the type series): 13 specimens, all from forest Camp region, Mt. Mabu, Zambezia Province, Mozambique (16 ˚ 17 ' 12 "S, 36 ˚ 24 ' 14 "E; 1000m a.s.l.); PEM R 17130 (J. Bayliss, January 2006), PEM R 17911 - 12 (J. Bayliss, 16 April 2009), PEM R 18057 -58, 18060, 18062 -67, 18071, same collecting details as holotype. Meristics. Measurements for the type series of Rhampholeon maspictus sp. nov. are summarized in Table 6. Diagnosis. The Mt. Mabu Pygmy Chameleon is referable to the Rhampholeon (subgenus Rhinodigitum Matthee et al. 2004) by possessing a short hemipenis that is almost bag-like, acalyculate and adorned with a pair of simple, curved apical ���horns��� with a variable number of thorn-like papillae arranged on the outer aspect of the horn; claws that are strongly bicuspid, smooth plantar surfaces, a rostral process, and short tail (Rhampholeon (Rhinodigitum) by having deep inguinal (absent or indistinct in Rh. boulengeri, Rh. nchisiensis, Rh. uluguruensis, and Rh. moyeri) and axillary pits (also absent in Rh. nchisiensis). It differs from all other members of the Rh. platyceps complex by the bright green male breeding coloration, including blue flanks and side of head, and yellow throat, snout and eye ring (all of which may be retained even at night). It shares with Rh. platyceps, but differs from all other populations of the complex from Mt Chiperone, Mt Namuli and Mt Inago, its large size (> 60 mm SVL) in both sexes, lack of male dwarfism, well-developed dorsal crenulations, and reduced rostral and supraocular processes; it differs from Rh. platyceps by its more dorsoventrally flattened habitus (more rounded in Rh. platyceps), and very weak or even absent accessory plantar spines (present but small in Rh. platyceps). Finally, the species is also genetically well differentiated from all other Rhampholeon, and all chameleons examined from Mt. Mabu form a monophyletic clade. Description of Holotype. Adult male, viscera exposed by a single ventral incision in the chest region; both hemipenes everted. Head: Dorsum of head flattened, with no upward flexure of the snout; casque flat with weakly define lateral crests; temporal crest moderate with a single row of interrupted large tubercles; parietal crest almost absent, composed of a few enlarged tubercles in the mid-line; supraorbital ridge present but with only few raised conical tubercles; three enlarged tubercles on inferior orbital rim; supraorbital connected in middle by shallow interorbital ridge composed of 11 small granular tubercles that demarcate the posterior edge of a prominent frontal depression; rostral ridge very well marked, forming a small bump over the nostrils and fusing anteriorly and adorned with a small rostral process (2.1 mm, in fixation flattened against rostrum); rostral process 4 small tubercular scales long and 4 tubercles wide at base; nares opening posterio-ventrally; no gular or mental appendages; scales on throat homogenous, more conical than those on crown of head and subequal in size to those on the belly. Body: Dorsal crest moderately developed, comprised of 9 weak crenulations each comprised of a cluster of enlarged spinose scales, crenulations most strongly developed over mid-body, reduced over base of neck and tail; crenulations continue onto tail comprised of 9 small groups that are largest over the distal half of the tail; deep axillary and inguinal pits are present; flank scalation heterogeneous, composed of small, stellate granules with scattered, enlarged spinose tubercles, the largest occurring over the shoulder; chest, belly and lower surface of tail smooth; limb scalation more tubercular, with a few enlarged, spinose tubercles on the forearms; claws strongly bicuspid; accessory plantar spines very reduced or almost absent. Colour in Life: Male coloration stunningly beautiful (Figs. 5 A); body, tail and limbs leaf green, slightly mottled and also darker along the dorsal crest of body, tail and upper surfaces of fore- and hindlimbs; central region of flanks light blue, with two oblique, broad yellow-green stripes; light blue of flanks extends onto the side of head and around eye, where it becomes darker and more intense; front of chest, throat, labials, rostral process and upper surface of snout to the level of the orbit, dirty yellow; bright yellow eye ring. When stressed (e.g. in artificially close proximity to another chameleon) the body coloration changes dramatically, the body darkening to greenbrown, and the blue flank region becomes paler and infused with brown blotches (Fig. 6 A); the green orbital region, yellow eye ring and dirty yellow throat colour are retained, albeit in subdued form. Colour in preservative: Body dark brown with 3 faint oblique bars of lighter brown on flanks running from backbone to lower flank; belly, tail and limbs uniform dark brown, except light brown on inner surface of forelimbs. Description of Allotype (as for holotype, unless noted): adult female, with viscera exposed by a single, ragged ventral incision in the belly, and liver tissue excised for DNA analysis. Head: Temporal crest weakly developed comprised of a disrupted row of slightly enlarge tubercles; parietal crest absent; subocular ridge weakly pronounced with few enlarged tubercles; interorbital ridge with 13 weakly enlarged tubercles Body: Dorsal crest very weakly developed, comprising 7 crenulations of tubercles that are only slightly enlarged; crenulations most strongly developed over mid-body and almost absent on the tail; flank scalation heterogeneous, composed of small, stellate granules with very few scattered, enlarged spinose tubercles, the largest at the shoulder. Coloration in Life: Less intensely coloured than male holotype (Figs. 5 B); sides of body dorsally dull green with two vague, oblique, dark olive stripes that extend from dorsal crest, where they are broader, onto lower flank; upper flanks with large, irregular light brown blotches that coalesce and cover upper surface of tail; lower flanks brighter green with scattered pale blotches, extending as pale green onto lower surface of tail; upper surfaces of limbs olive, paler green below; upper surface of casque pale brown, extending at a narrow, almost white line along the dorsal crest; front and sides of snout mottled in light green and brown giving a dull green-brown appearance; throat cream with numerous light blue tubercles that coalesce to form irregular blotches; orbital area olive green with scattered pale blue tubercles and a dull red eye ring. When stressed the irregular light brown blotches on the upper flanks become paler, and the sides of head become blotched, especially in the labial region, giving an appearance of alternating pale and dark bands radiating from the eye (Fig. 6 A). Colour in preservative: Body and head uniform dark brown above and below. Paratype variation. All paratype males have well-everted hemipenes. In life the orbital scalation of a male paratype (R 17130) was light green, with a conspicuous yellow ring around the iris; the pale brown body had two prominent rust red ���leaf vein��� narrow lines that ran from the dorsal crest obliquely to the lower flank; there is a prominent enlarged, pale yellow-cream spinose tubercle on the fore-flank in the shoulder region; the crown of the casque and dorsal crest on the body and tail was medium brown (Fig. 6 B). A very small specimen (PEM R 18068, SVL 19.6 mm) has a mottled reddish brown head, body and limbs, with two vague, narrow oblique red-brown stripes that run from the dorsal crest to the lower flanks; the dorsal crest forms a narrow cream stripe that extends onto the casque, where it becomes disrupted into small blotches; the limbs are darker brown with scattered bright yellow tubercles of the upper surfaces; the orbital region is dull brown with scattered small blue tubercles and a dull red eye ring (Fig. 6 C). Size. Largest male���PEM R 18074 (paratype) 65.2 + 20.1 = 85.3 mm; largest female���PEM R 18069 (paratype) 64.4 + 16.2 = 80.6 mm. The smallest specimen���PEM R 18068 (paratype, unsexed) 19.9 + 4.9 = 24.8 mm) appears to be newly hatched. Sexual Dimorphism. The average total size of males (65.3 mm, n = 11) and females (64.4 mm, n = 11) is similar and suggests no sexual size dimorphism. However, males do have proportionately longer tails (TL as = % of SVL 24.0% in males, but only 18.6 % in females). Sexual dichromism in pygmy chameleons remains poorly known, and has not been studied in the Rh. platyceps complex. Tilbury (1992) noted the emergence of bright colours (pale powder blue head and neck, and bright white eyelids) in male Rh. chapmanorum in male-male interactions. The bold colours of large male Rh. maspictus sp. nov. (Fig. 5 A) was noted in three different males at the type locality, including two specimens sleeping at night on low vegetation. No female was observed showing the bright green body and blue and yellow head of adult males. Hemipenis. (based on everted hemipenes of five males in the type series). Hemipenis short with unadorned basal and apical sections, except for a pair of curved horns that form the lateral edges of a prominent, flat-topped crown; 3-4 weakly-developed spines occur on the upper surfaces of the horns; the folds of the sulcus spermaticus flare to drain into the flattened apical region. Distribution. Known only from the type locality; Mt. Mabu, Zamb��zia Province, central Mozambique, but expected to occur throughout much of the extensive forest (+ 7880 ha) remaining on the massif (Bayliss et al. 2014). Habitat. The Mabu massif is a complex of granitic inselbergs formed of syenite, an igneous intrusion of the younger Precambrian Namarroi series dating from 850 ���1100 Mya (Instituto Nacional de Geologia 1987). Although the Mabu Massif shows vegetational zonation and a significant extent (+ 7880 ha) of mid-altitudinal forest, it lacks the large area of upland plateau that occurs on Mt. Namuli. Mt. Mabu is surrounded by Brachystegia woodland at the base, much of which is replaced to the south by the Cha Madal tea estate. The paratype series were collected at night on perches from 30-100 cm above ground in forest clearings and adjacent to a mountain stream (Fig. 6 D). No specimens were collected on perches as high as 1.3 m to 5 m noted for Rh. chapmanorum (Tilbury 1992). This may be related to the presence of an additional chameleon (N. baylissi) which utilizes this forest stratum on Mt. Mabu, and that has not been recorded on Mt. Chiperone. Reproduction. The female allotype was gravid with seven almost spherical eggs (largest diameter 4.1-4.5 mm, four in the left oviduct and three in the right oviduct) that lack obvious signs of embryonic development., Published as part of Branch, William R., Bayliss, Julian & Tolley, Krystal A., 2014, Pygmy chameleons of the Rhampholeon platyceps compex (Squamata: Chamaeleonidae): Description of four new species from isolated ' sky islands' of northern Mozambique, pp. 1-36 in Zootaxa 3814 (1) on pages 17-22, DOI: 10.11646/zootaxa.3814.1.1, http://zenodo.org/record/286211, {"references":["Tilbury, C. R. (2010) Chameleons of Africa. An Atlas including the Chameleons of Europe, the Middle East and Asia. Edition Chimaira, Frankfurt am Main, 831 pp.","Timberlake, J. R., Bayliss, J., Dowsett-Lemaire, F., Congdon, C., Branch, B., Collins, S., Dowsett, R. J., Fishpool, L., Francisco, J. Harris, T., Kopp, M. & da Sousa, C. (2012) Mount Mabu, Mozambique: Biodiversity and Conservation. Report produced under the Darwin Initiative Award 15 / 036: Monitoring and Managing Biodiversity Loss in South-East Africa's Montane Ecosystems. Royal Botanic Gardens, Kew, London. pp. 94.","Bayliss, J., Timberlake, J., Alves, T., Branch, W. R., Bruessow, C., Collins, S., Congdon, C., De Sousa, C., Dowsett-Lemaire, F., Fishpool, L., Harris, T., Herrmann, E., Giogiardis, S. Liggitt, B., Monadjem, A., Patel, H., Spottiswoode, C., Taylor, P., Wilcocks, S., Ribeiro, D. & Smith, P. (2014) The discovery, biodiversity, and conservation of Mabu forest-the largest mid-altitude rainforest in southern Africa. Oryx, 48 (2), 177 - 185. http: // dx. doi. org / 10.1017 / s 0030605313000720","Tilbury, C. R. (1992) A new dwarf forest chameleon (Sauria: Rhampholeon) from Malawi, Central Africa. Tropical Zoology, 5, 1 - 9. http: // dx. doi. org / 10.1080 / 03946975.1992.10539176"]}

Branch, William R., Bayliss, Julian, Tolley, Krystal A. (2014): Pygmy chameleons of the Rhampholeon platyceps compex (Squamata: Chamaeleonidae): Description of four new species from isolated ' sky islands' of northern Mozambique. Zootaxa 3814 (1): 1-36, DOI: http://dx.doi.org/10.11646/zootaxa.3814.1.1

Rhampholeon (Rhinodigitum) nebulauctor sp. nov. Mount Chiperone Pygmy Chameleon Synonomy: Rhampholeon champmanorum (sic.) Timberlake et al. 2007, p 20. Etymology. The specific epithet is a noun in apposition and derived from nebula (L. = cloud, mist) and auctor (L. = maker), i.e. ���cloudmaker���, alluding to the ���Ciperoni���, the local name for the cold drizzle that comes to the Shire Highlands of southern Malawi as moist air from the Indian Ocean is forced to rise over Mt. Chiperone. Types. The type series comprises five specimens, including: Holotype.- An adult female (PEM R 17278) collected by J. Bayliss, 1 December 2008, in the shrub understorey of evergreen forest on the southeast slopes of Mt. Chiperone Massif, Zamb��zia Province, Mozambique (16 ˚ 30 ' 25.9 ���S, 35 ˚ 43 ' 33.4 ���E, ca 1000 m a.s.l.). Allotype. An adult male (PEM R 17281), collected by J. Bayliss on 27 November 2008; same locality details as holotype. Paratypes. Three specimens, comprising an adult female (PEM R 17277) and two subadult females (PEM R 17279 - 80), all collected by J. Bayliss between 26 November and 3 December 2008, same locality details as holotype. Meristics. Measurements for the type series of Rhampholeon nebulauctor sp. nov. are summarized in Table 7. Diagnosis. The Chiperone Pygmy Chameleon is referable to the Rhampholeon (subgenus Rhinodigitum Matthee et al. 2004) by having an unpigmented parietal peritoneum, claws that are strongly bicuspid, smooth plantar surfaces, a rostral process, and short tail (Rhampholeon (Rhinodigitum) by having deep inguinal (absent or indistinct in Rh. boulengeri, Rh. nchisiensis, Rh. uluguruensis, and Rh. moyeri) and axillary pits (also absent in Rh. nchisiensis). It is geographically closest to Rh. chapmanorum, but differs from that species by its smaller size, the presence of a relatively large rostral process in males, and accessory planter spines that are very poorly developed in both sexes. It is well differentiated from Rh. platyceps and Rh. maspictus sp. nov. by its smaller size (SVL Rhampholeon, and all chameleons examined from Mt. Chiperone form a monophyletic clade. Description of Holotype. Adult female, viscera exposed by a large ventral incision. Head: Dorsum flattened, with no upward flexure of the snout; casque flat, edged with weakly-defined lateral crests that are mainly restricted to the posterior region of the casque; temporal crest weakly-developed, comprising a single, interrupted row of large tubercles; parietal crest almost absent, composed of a few enlarged tubercles in the mid-line; supraorbital ridges reduced to a few scattered enlarged scales but with a very small multi-scaled process forming a ���soft horn��� at each end of the inter-orbital ridge that passes across crown and is composed of 11 small granular tubercles, and that demarcate the posterior edge of a slight frontal depression; inferior orbital rim with 4 (right) and 4 (left) enlarged tubercles; snout bordered on each side by moderately developed rostral crests, that fuse together at the tip of the snout which is adorned with a very small, flattened rostral process (1.5 mm), and is three small tubercular scales long and three tubercular scales wide at its base; nares opening posterio-ventrally; no gular or mental appendages; scales on throat homogenous, more conical but smaller than those on crown of head and subequal in size to those on the belly. Body: Dorsal crest very weakly developed, reduced to 7 crenulations of enlarged, but not obviously spinose scales; crenulations most strongly developed over mid-body, reduced in size over on neck and tail; crenulations continue onto tail but in more reduced form; deep axillary and inguinal pits are present; flank scalation heterogeneous, composed of small, stellate granules with few scattered, enlarged spinose tubercles, the largest at the shoulder and in a single cluster; chest, belly and lower surface of tail smooth; limb scalation more tubercular, with a few enlarged, spinose tubercles on the forearms; claws strongly bicuspid; accessory planter spines on the soles of the fore and hind feet are present, but reduced to a few very small, soft, spinose scales at the base of the claws; tail flattened laterally, flexing slightly downward on the distal third. Colour in life (Fig. 7 A, B): Mid-body mottled brown with two vague oblique lateral stripes; fore-body, neck, head and upper surfaces of limbs with extensive bright green tubercles, with scattered light blue tubercles on temporal region of head and throat and on front of belly; scales around orbit dark blue, with a red rim to the iris; tip of casque and dorsal crest cream; enlarged scales of the dorsal crest crenulations orange, that is more extensive and intense along the top of the tail. Colour in preservative. Body mottled brown with no obvious lateral stripes; lower surface of neck, belly, base of tail, soles of feet, and lower limbs pale brown, except on throat and sides of head which are dark brown. Description of Allotype (as for holotype, unless noted): adult male, very large ventral incision; hemipenes not everted. Head: Supraorbital ridge reduced to a few scattered enlarged scales but with a distinct multi-scaled process forming a ���soft horn��� at the end of the inter-orbital crest that is slightly more pronounced than in female holotype; interorbital ridge, shallow and composed of 12 small granular tubercles; rostral process six tubercular scales long and five wide at base. Body: Dorsal crest very weakly developed, reduced to eight crenulations of enlarged but not obviously spinose scales, which are subequal in development to those of the female holotype; tail relatively long (26.5 % SVL), with a prominent hemipenal bulge; distal third of tail flexes strongly downward. Colour in preservative: Body mottled brown with no obvious lateral stripes on the flanks. Pale brown below. Paratype variation. In the largest female paratype (PEM R 17277) the ���horn��� on the supraorbital ridge is reduced to a very small cluster of about 5 spinose tubercular scales; rostral process is very small, 3 scales long and 4 wide at base. Size. Presumably a small species, as all except the smallest female were reproductively active. Largest male���PEM R 17281 (allotype) 32.3 + 11.6 = 43.9 mm; largest female���PEM R 17277 (paratype) 48.9 + 13.2 = 62.1 mm. Reproduction. The three adult females contain enlarged ova, albeit without signs of embryonic development: PEM R 17277, Four ova ca. 3.4 mm dia.; PEM R 17278, three ova largest measuring 5.0 mm x 3.5 mm; PEM R 17279, two small ova (ca. 3.1 mm dia.). The smallest female, PEM R 17280, has small developing follicles. The only male (PEM R 17281) has well-developed testes despite its small size (32.3 mm SVL). Distribution. Restricted to the type locality; Mt. Chiperone, Zamb��zia Province, northern Mozambique. Habitat. The Chiperone Massif shows habitat zonation and is surrounded by Brachystegia woodland at the base. Chameleons were found on the southeast side of the mountain at 1000 m in wet forest dominated by Khaya anthotheca, Strombosia schefflerii, Rawsonia burtt-davyi, and Drypetes arguta (Fig. 7 C.), Published as part of Branch, William R., Bayliss, Julian & Tolley, Krystal A., 2014, Pygmy chameleons of the Rhampholeon platyceps compex (Squamata: Chamaeleonidae): Description of four new species from isolated ' sky islands' of northern Mozambique, pp. 1-36 in Zootaxa 3814 (1) on pages 22-24, DOI: 10.11646/zootaxa.3814.1.1, http://zenodo.org/record/286211, {"references":["Timberlake, J. R., Bayliss, J., Alves T., Baena, S., Francisco, J., Harris, T. & da Sousa, C. (2007) The Biodiversity and Conservation of Mount Chiperone, Mozambique. Report produced under the Darwin Initiative Award 15 / 036: Monitoring and Managing Biodiversity Loss in South-East Africa's Montane Ecosystems. Royal Botanic Gardens, Kew, London. pp. 33."]}

Rhampholeon (Rhinodigitum) platyceps G��nther, 1892. Mount Mulanje Pygmy Chameleon Synonomy: Brookesia platyceps Loveridge 1933 Brookesia platyceps platyceps Loveridge 1953 Brookesia platyceps carri Loveridge 1953 Rhampholeon (Rhinodigitum) platyceps Matthee, Tilbury & Townsend 2004. Holotype: Adult female (BMNH 1946.8.21.73), collected by A. White. Type locality (by subsequent designation, Stevens 1974): Mt. Mulanje, southern Malawi. Variation in adult coloration of an adult male (Fig. 4 A) and adult female (Fig. 4 B), and evergreen forest habitat in the Lichenya plateau region, are shown (Fig. 4 C). For full details of morphology, hemipenial structure, biology and habitat see Tilbury (2010)., Published as part of Branch, William R., Bayliss, Julian & Tolley, Krystal A., 2014, Pygmy chameleons of the Rhampholeon platyceps compex (Squamata: Chamaeleonidae): Description of four new species from isolated ' sky islands' of northern Mozambique, pp. 1-36 in Zootaxa 3814 (1) on pages 15-16, DOI: 10.11646/zootaxa.3814.1.1, http://zenodo.org/record/286211, {"references":["Gunther, A. (1892) Report on a collection of reptiles and batrachians transmitted by Mr H. H. Johnson from Nyasaland. Proceedings of the Royal Society of London, 1982, 555 - 558.","Loveridge, A. (1953) Zoological results of a fifth expedition to East Africa. III Reptiles from Nyasaland and Tete. Bulletin of the Museum of Comparative Zoology, 110, 141 - 322.","Matthee, C. A., Tilbury, C. R. & Townsend, T. (2004) A phylogenetic review of the African pygmy chameleons: genus Rhampholeon (Chamaeleonidae): the role of vicariance and climate change in speciation. Proceedings of the Royal Society of London, Series B, 271, 1967 - 1976.","Stevens, R. A. (1974) An annotated check list of the amphibians and reptiles known to occur in southeastern Malawi. Arnoldia (Rhodesia), 6 (30), 1 - 22.","Tilbury, C. R. (2010) Chameleons of Africa. An Atlas including the Chameleons of Europe, the Middle East and Asia. Edition Chimaira, Frankfurt am Main, 831 pp."]}

Meroles Gray 1838 Type species. Meroles knoxii (Milne-Edwards 1829) Content. Meroles anchietae (Bocage 1867), Meroles ctenodactylus (Smith 1838), Meroles cuneirostris (Strauch 1867), Meroles knoxii (Milne-Edwards 1829), Meroles micropholidotus Mertens 1938, Meroles reticulatus (Bocage 1867), Meroles squamulosus (Peters 1854), Meroles suborbitalis (Peters 1869) Characterization and diagnosis. The inclusion of M. squamulosus requires the genus to be redefined. Head shields normal and usually smooth (rugose in squamulosus), but occipital often very small or absent; nostril pierced between three nasals and widely separated from 1 st upper labial; subocular not bordering mouth; lower eyelid scaly, without window; collar distinct (absent in squamulosus); gular fold absent; dorsal scales granular, juxtaposed or subimbricate, (but rhombic, strongly keeled and imbricate in squamulosus); ventral plates smooth, not or feebly imbricate, posterior borders straight; digits subcylindrical, compressed or depressed (feebly compressed in squamulosus), laterally serrated, denticulated or fringed (except in squamulosus); subdigital lamellae smooth or keeled (pluricarinate and spinolose in squamulosus), femoral pores present; parietal foramen present (absent or feebly marked in squamulosus); and tail long and cylindrical (in knoxii, suborbitalis and squamulosus) or depressed basally and feebly compressed distally. Remark. As the gender of Meroles is masculine the specific ending of squamulosa must be adjusted accordingly to squamulosus., Published as part of Edwards, Shelley, Branch, William R., Vanhooydonck, Bieke, Herrel, Anthony, Measey, G. John & Tolley, Krystal A., 2013, Taxonomic adjustments in the systematics of the southern African lacertid lizards (Sauria: Lacertidae), pp. 101-114 in Zootaxa 3669 (2) on page 110, DOI: 10.11646/zootaxa.3669.2.1, http://zenodo.org/record/221968

Vhembelacerta Edwards, Herrel, Vanhooydonck, Measey, Tolley& Branch, gen. nov. Type species. Lacerta rupicola FitzSimons 1933 Content. Vhembelacerta rupicola (FitzSimons 1933) Characterization and diagnosis. The monophyly of the monotypic genus Vhembelacerta is established on the basis of a suite of mitochondrial and nuclear markers (Edwards et al. 2012; this study). Morphologically similar to Australolacerta (differences noted in brackets below), it can be distinguished from all other lacertids by the following combination of characteristics (FitzSimons 1943; Jacobsen 1989; Branch 1998;Kirchhof& Richter 2009): size small, (SVL ~ 52mm), maximum snout-vent length (SVL) 70mm, tail somewhat depressed basally, cylindrical distally, nearly 1.5 x SVL (up to 2 x SVL); upper head shields smooth; nostril pierced between a supranasal, 2 postnasals and narrowly separated from first upper labial (in contact with first upper labial); supranasals in contact behind rostral; frontonasal much broader than long; prefrontals in contact; frontal hexagonal; supraoculars 4, 1st smallest and separated from frontal; parietals in contact with 4 th supraocular; shallow parietal foramen present (absent); rostral not entering nostril; 5 (4) upper labials anterior to subocular, whose lower border is not distinctly narrowed (lower border much shorter than upper) and only feebly keeled; lower eyelid scaly but with a about 3 enlarged and elongate scales in the middle (no enlarged scales in lower eyelid); elongate tympanic shield on upper anterior border of large, exposed ear-opening; five pairs of chin shields, first smallest, first 3 in contact in midline; gular fold distinct (present, but not strongly marked); collar present, straight, free, composed of 7���8 scales; dorsal scales flat, hexagonal, faintly keeled posteriorly and in 34���43 rows across midbody (small, granular, non-keeled and about 68 across midbody); ventral plates quadrangular, feebly imbricate and in 6 longitudinal and about 26 (28) transverse rows; a very large preanal plate, bordered by smaller plates (enlarged preanal preceded by two smaller ones); femoral pores 15���19; subdigital lamellae smooth, about 26 below 4 th toe (23���25); and the adpressed hindlimb reaches the armpit (to collar). Coloration: top of head and dorsum dark brown, back with paired narrow reddish brown vertebral stripes, and a white dorsolateral stripe that extends from the eye to the tail base (head and dorsum olive-green, body with numerous spots that are yellow dorsally and white on flanks, demarcated by a dorsolateral series of orange spots). Distribution. Endemic to the Soutpansberg mountain range in Limpopo Province, South Africa (Branch 1998). Etymology. The species is endemic to the Vhembe region of Limpopo Province, South Africa, after which the genus name is partially constructed. The second part of the name ���lacerta��� (L. lizard) also retains the historical link to the genus Lacerta to which the single species was originally referred., Published as part of Edwards, Shelley, Branch, William R., Vanhooydonck, Bieke, Herrel, Anthony, Measey, G. John & Tolley, Krystal A., 2013, Taxonomic adjustments in the systematics of the southern African lacertid lizards (Sauria: Lacertidae), pp. 101-114 in Zootaxa 3669 (2) on page 108, DOI: 10.11646/zootaxa.3669.2.1, http://zenodo.org/record/221968

Australolacerta Arnold 1989 Type species. Lacerta australis Hewitt 1926 Content. Australolacerta australis (Hewitt 1926) Characterization and diagnosis. With the transfer of Lacerta rupicola to Vhembelacerta, a re-diagnosis of Australolacerta is required. The monophyly of the monotypic genus Australolacerta is established on the basis of a suite of nuclear and mitochondrial markers (Edwards et al. 2012). Morphologically closest to Vhembelacerta, it can be distinguished from all other lacertids by the following combination of characteristics (FitzSimons 1943; Branch 1998): head moderately depressed, body feebly so; SVL 50���65mm, maximum 70mm; adpressed hindlimb reaches collar; tail cylindrical; head shields normal with upper head shields smooth, occipital region flat; snout pointed, shorter than postocular part of head; nostril pierced between the nasal and one or two postnasals, and 1 st upper labial, with nasals in contact with one another behind rostral; frontoparietals paired in contact; parietals in contact with the 4 th of four supraoculars and separated from the postoculars; parietal foramen absent; interparietal about twice as long as broad, in good contact with occipital; a series of 9 granules between supraoculars and supraciliaries; 4 upper labials anterior to subocular, which has a strongly-marked keel along upper border and a lower border that is much shorter than upper; elongate temporal shield posterior to the subocular, followed by 3 smaller rounded ones; temporal scales small and granular, similar to dorsal scales; ear-opening large, exposed, bordered anteriorly by an elongate tympanic shield and with no auricular denticulation; lower eyelid scaly, lacking vertically-enlarged scales in the middle; 6 lower labials and five pairs of enlarged chin-shields, 1 st smallest, 4 th largest, and 1 st three pairs in median contact with one another; gular fold present, but not strongly marked; collar composed of 8 plates, straight, free, and even-edged; dorsal scales small, granular, smooth, similar to laterals and about 68 across midbody; ventral plates quadrangular, feebly imbricate, in 6 longitudinal and 28 transverse series; preanal plate enlarged, preceded by two smaller scales; Forelimb with small granular scales on upper surface of forearm and a series of strongly enlarged, smooth and imbricate plates along anterior surface of humerus; hindlimb with granular scales on upper surface of tibia; a series of much enlarged and vertically elongate plates run along anterior surface of thigh and on the lower surface of tibia;. 16���19 femoral pores; sub-digital lamellae smooth; scales on tail enlarged, quadrangular, elongate; more or less smooth dorsally, becoming keeled distally, and below scales smooth basally, more strongly keeled and bluntly mucronate distally. Coloration: head and dorsum dark brown to olive, with numerous pale spots arranged in more or less regular longitudinal series that are yellow on back, white on flanks, and separated by a dorsolateral series of orange spots; upper surface head with pale green to yellow vermiculations; distinct pale vertical stripes on temporal region; indistinct pale spots on tail; venter bluish green; labials, chin-shields and throat pale greenish, with small black spots and mottling. Distribution. Found in the southwestern Cape Fold Mountains in Western Cape Province, South Africa (Branch 1998)., Published as part of Edwards, Shelley, Branch, William R., Vanhooydonck, Bieke, Herrel, Anthony, Measey, G. John & Tolley, Krystal A., 2013, Taxonomic adjustments in the systematics of the southern African lacertid lizards (Sauria: Lacertidae), pp. 101-114 in Zootaxa 3669 (2) on page 109, DOI: 10.11646/zootaxa.3669.2.1, http://zenodo.org/record/221968

Bates, Michael F., Tolley, Krystal A., Edwards, Shelley, Davids, Zoë, Da Silva, Jessica M., Branch, William R. (2013): A molecular phylogeny of the African plated lizards, genus Gerrhosaurus Wiegmann, 1828 (Squamata: Gerrhosauridae), with the description of two new genera. Zootaxa 3750 (5): 465-493, DOI: http://dx.doi.org/10.11646/zootaxa.3750.5.3

Monophyly of Australolacerta The two species of Australolacerta are endemic to South Africa (Branch 1998), and both were originally placed within Lacerta, creating a zoogeographic paradox as most congeners were restricted to Eurasia (Arnold 1989). Arnold (1989), when describing Australolacerta, gave only a minimal diagnosis, noting that “… the South African species share a number of features with other Ethiopian lacertids which are not found in the apparent closest Palaearctic relatives, namely Lacerta jayakari etc.”. The latter, now transferred to Omanosaura, was initially considered to form a basal lineage within the Eremiadini (Harris et al. 1998), although fuller taxon sampling of African lacertids (Arnold et al. 2007; Hipsley et al. 2009; Kapli et al. 2011) shows it to cluster with a suite of mainly north African genera (e.g. Acanthodactylus, Mesalina, Ophisops), with Atlantolacerta basal within the Eremiadini (Arnold et al. 2007). The sister relationship of Australolacerta and Tropidosaura proposed by Salvi et al. (2011) and Kapli et al. (2011) was based on the inclusion only of A. australis, and the inclusion of A. rupicola (Edwards et al. 2012; Engleder et al. 2013; this study) revealed the paraphyly of Australolacerta and the basal position of A. australis in a subclade including Ichnotropis, Tropidosaura and A. rupicola. Both species are rupicolous and Kirchhof and Richter (2009) and Kirchhof et al. (2010 a,b; 2012) give details of the species’ biology. They are morphologically similar, albeit that many of these similarities are plesiomorphic within lacertids (Arnold 1989). Due to their high-altitude and small ranges (Branch 1998), the two species have been difficult to collect and therefore little morphological data exists for either species. Recent morphological analyses (Edwards et al. 2012) confirm the similarity between the two species. Yet, important features of hemipenial ornamentation and everted hemipenis structure remain unknown. Whether these similar morphologies reflect adaptive convergence to rupicoly or the retention of plesiomorphic features remains unknown. The two species are allopatric and geographically separated from one another by a distance of approximately 1700km. Few other genera in southern Africa are known to show such large geographical disjunctions between congeners, and analysis of previous examples has often revealed deep genetic divergence best reflected in generic re-assignment. Examples include: the erection of the genera Kinyongia and Nadzikambia for non-South African dwarf chameleons previously included in Bradypodion (Tilbury et al. 2006), and Inyokia for the problematic Swazi rock snake that was shown to be sister to the tropical forest snake Homonotus modestus (Kelly et al. 2011). In one of the few exceptions of congeneric range disjunctions within the subcontinent, cordylid flat lizards of the Platysaurus capensis complex are separated geographically from other Platysaurus by approximately 850km (Branch & Whiting 1997). Other described lacertid species are also separated from congeners by large distances, for example Heliobolus lugubris is separated from its congeners (H. spekii, and H. nitidus) by> 2000km and Ichnotropis chapini is separated from other Ichnotropis by approximately 2000km (Branch, 1998; Spawls et al. 2002). However, the Central African region is undersampled and it is possible that with increased sample collection new species may be discovered or that ranges of described species may increase, lessening the geographic gap between congeners. Although there were no significant differences between the obtained trees and the constrained trees in the SH or AU test when the topology was constrained to monophyly for Australolacerta sequence divergence estimates and the long branch lengths in the phylogenetic analyses (Figure 1) all other evidence strongly suggests that the two Australolacerta species do not share a recent evolutionary history (Edwards et al. 2012).To provide consistency between taxonomic divisions in the Eremiadini, we propose that the two species of Australolacerta should be placed in separate genera. The type species of Australolacerta is Lacerta australis (Arnold 1989), and we therefore erect a new genus for the remaining species Lacerta rupicola, based on morphology and genetic divergence.

Ichnotropis Peters 1854. Type species. Ichnotropis macrolepidota (Peters 1854); = I. capensis (Smith 1838) Content. Uetz (2012) recognizes six species (excluding squamulosus): Ichnotropis bivittata Bocage 1866, Ichnotropis capensis (A. Smith 1838), Ichnotropis chapini Schmidt 1919, Ichnotropis grandiceps Broadley 1967, Ichnotropis microlepidota Marx 1956, Ichnotropis tanganicana Boulenger 1917. Characterization and diagnosis. The monophyly of the genus Ichnotropis remains to be established with complete taxon sampling of the referred species. The removal of M. squamulosus from Ichnotropis does not significantly alter the diagnosis for the genus given in FitzSimons (1943), as morphological variation within the remaining species still incorporates that of M. squamulosus. Remark. No modern revision of the genus has been undertaken, and the status of a number of taxa remains equivocal, e.g. Ichnotropis bivittata pallida Laurent 1964; Ichnotropis capensis nigrescens Laurent 1952; Ichnotropis microlepidota Marx 1956, and the generic assignment of many requires molecular confirmation., Published as part of Edwards, Shelley, Branch, William R., Vanhooydonck, Bieke, Herrel, Anthony, Measey, G. John & Tolley, Krystal A., 2013, Taxonomic adjustments in the systematics of the southern African lacertid lizards (Sauria: Lacertidae), pp. 101-114 in Zootaxa 3669 (2) on page 110, DOI: 10.11646/zootaxa.3669.2.1, http://zenodo.org/record/221968

Gerrhosaurus Wiegmann, 1828 Pleurotuchus Smith, 1837 Angolosaurus FitzSimons, 1953 Type species: Gerrhosaurus flavigularis Wiegmann, 1828 Content: Gerrhosaurus flavigularis Wiegmann, 1828; Gerrhosaurus typicus (Smith, 1837); Gerrhosaurus nigrolineatus Hallowell, 1857; Gerrhosaurus multilineatus Bocage, 1866 a; Gerrhosaurus auritus Boettger, 1887; Gerrhosaurus intermedius L��nnberg, 1907 comb. nov.; Gerrhosaurus skoogi Andersson, 1916; Gerrhosaurus bulsi Laurent, 1954. Diagnosis: The monophyly of Gerrhosaurus is established on the basis of a suite of nuclear and mitochondrial genetic characters (see above). These moderate-sized lizards are fairly well armoured and the head and body may be cylindrical, cyclotetragonal or slightly depressed; differentiated from the genera Broadleysaurus and Matobosaurus by its smaller size (maximum SVL 213 mm compared to 245 mm and 285 mm respectively for the latter two genera) and less robust appearance; most species of Gerrhosaurus have only eight ventral scale rows longitudinally (but 10 in G. typicus), whereas Broadleysaurus has 9���10 and Matobosaurus has 12���20; it also differs from Broadleysaurus by having 49���67 versus 31���38 transverse dorsal scale rows (Loveridge 1942; FitzSimons 1943, 1953; De Witte 1953; Laurent 1954, 1964; Broadley 1966; De Waal 1978; Jacobsen 1989). Description: Head large, moderate or small, its length included in SVL 3.3���4.8 times (young lizards) to 4.0��� 8.4 times (adults); head shields smooth or weakly striated; rostral in contact with, or separated from, the frontonasal; prefrontals well separated, slightly separated, in narrow contact, or in broad contact; supraoculars 4; supraciliaries 4���5 (rarely 3 or 6); tympanic shield narrow and band-like to broad and crescentic; body cyclotetragonal, slightly depressed in some G. t y pi cu s, or almost cylindrical (G. skoogi); dorsal scales weakly to strongly keeled, smooth or striated, in 20���28 (32���35 in G. skoogi) longitudinal and 49���67 transverse rows (usually counted from row posterior to nuchals to row above vent); lateral scales keeled, striated or smooth; ventral plates in 8 or 10 (G. typicus only) longitudinal and 30���42 transverse rows (counted ���from pectoral to anal shields��� according to Loveridge 1942; i.e. from axilla to row before enlarged ventral plate); femoral pores 9���27 per thigh; fourth toe with 14���22 subdigital lamellae; largest known specimens: unknown sex 613 mm (213 mm SVL + 400 mm tail length), male 485 mm (163 + 322), but another male had a SVL of 175 mm female: 475 (142 + 333), but another female had a SVL of 157 mm; tail 1.0 to 2.5 times SVL (Loveridge 1942; FitzSimons 1943, 1953; De Witte 1953; Laurent 1954, 1964; Broadley 1966; De Waal 1978; Jacobsen 1989). Distribution: Widespread in Africa south of the equator, extending northwestwards into Gabon and Cabinda, and north-eastwards through Uganda and Kenya to southern Sudan and Ethiopia (Loveridge 1942; FitzSimons 1943; De Witte 1953; Mertens 1955; Broadley 1966, 1971; De Waal 1978; Auerbach 1987; Jacobsen 1989; Lang 1991; Branch 1998; Spawls et al. 2002; Adolphs 2006, 2013; Bates et al. in press.). Note: Lizards in this genus are commonly known as ���plated lizards���. Status of ��� Gerrhosaurus major ��� The type locality of G. major (Fig. 4) is Zanzibar, an island off the coast of Tanzania, but G. m. major has an extensive range in the eastern half of Africa, from northern KwaZulu-Natal in South Africa to Ethiopia; G. m. bottegoi was described from Valley of Ghinda in Eritrea and has a fragmented distribution, extending from northeast Africa (where it occurs together with the nominate subspecies in Kenya) across the continent to West Africa (Dum��ril 1851; Del Prato 1895; Loveridge 1942; FitzSimons 1943; Broadley 1966; Jacobsen 1989; Branch 1998; Spawls et al. 2002; Adolphs 2006, 2013; Bates et al. in press.). The two subspecies are distinguishable only by their colour patterns (Broadley 1987). Our analysis included samples from southern and eastern Africa identifiable as G. m. major and one sample from Atakpame in Togo referable to G. m. bottegoi (Table 1). The Togo sample is embedded within samples of G. m. major. Based on our molecular data, plus the weak morphological differences (i.e. colour variation) used for recognition of the two subspecies, we relegate G. bottegoi Del Prato, 1895 to the synonomy of Broadleysaurus major (Dum��ril, 1851) comb. nov. Status of ��� Gerrhosaurus validus ��� The two currently recognized subspecies of G. va l i d us each form separate monophyletic clades. In addition, sequence divergences between these taxa are much larger than would be expected for subspecies and instead are at the level of species (i.e. 8.5 % ND 2, 4.1 % 16 S). The two taxa are morphologically well differentiated (e.g. subocular excluded from lip by a labial in validus, in contact with lip in maltzahni; longitudinal rows of dorsals 28��� 34 in validus, 25���30 in maltzahni; longitudinal rows of ventrals 14���20 in validus, 12���14 in maltzahni; Loveridge 1942, FitzSimons 1943) and occur allopatrically. Gerrhosaurus v. validus occurs from Limpopo Province in South Africa northwards to Mozambique, Zimbabwe, Zambia and Malawi, while G. v. maltzhani (type locality: Farm Roidina, north of Omaruru, Namibia; De Grys 1938) is restricted to northern Namibia and southern Angola (Loveridge 1942; FitzSimons 1943; Broadley 1966; Visser 1984 a; Jacobsen 1989; Branch 1998; Spawls et al. 2002; Adolphs 2006, 2013; Bates et al. in press.). The two taxa appear to be separated by the Kalahari Desert (Visser 1984 a). Our samples of G. v. validus were from Limpopo Province in South Africa, Mozambique and Zimbabwe; G. v. maltzahni was sampled in both Namibia and Angola (Table 1). The type locality for G. validus of ���towards the sources of the Garrep [Gariep], or Orange River��� (Smith 1849, Appendix, p. 9), i.e. in Lesotho, must be in error―as noted by FitzSimons (1943)―as the species is not known to occur anywhere south of 28 o latitude (Branch 1998; Bates et al. in press.). The combination of molecular, morphological and geographical evidence suggests that the two taxa represent separate evolutionary lineages, and we therefore revive G. maltzahni De Grys, 1938 as a full species, as Matobosaurus maltzahni (De Grys, 1938) comb. nov. The two species in the genus are illustrated in Figs 5 & 6. Status of taxa in the Gerrhosaurus nigrolineatus species complex The type locality of G. nigrolineatus is ��� Gaboon country, West Africa��� (= Gabon; Hallowell 1857). This species has now been collected at several localities in Gabon (Pauwels et al. 2006), confirming its occurrence there. As currently understood it has a large distribution range, from Gabon and the lower Congo eastwards through southern Democratic Republic of the Congo (D.R.C.) to Uganda and Kenya in the east, then southwards as far as northern Namibia, northern Botswana and north-eastern South Africa (Loveridge 1942; FitzSimons 1943; De Witte 1953; Broadley 1966, 1971; Auerbach 1987; Jacobsen 1989; Branch 1998; Spawls et al. 2002; Bates et al. in press.; Uetz 2013). Our samples were from Kouilou region, Republic of the Congo (west-Central Africa) adjacent to Gabon, and Tanzania, Mozambique and South Africa (East and Southern Africa) (Table 1). Our analysis showed that G. nigrolineatus as currently conceived is not monophyletic, although topology tests could not reject a monophyletic G. nigrolineatus as presently defined. However, given the observed topology, the well-supported west-Central African clade of G. nigrolineatus is more closely related to G. auritus, rather than to G. nigrolineatus from East and Southern Africa, and the nodes defining these groups are well-supported. Given the node support, as well as other lines of evidence (see below), we suggest that there is reasonably strong support that G. nigrolineatus as currently defined is not monophyletic. Although the phylogeny of Lamb et al. (2003) also recovered a sister relationship between G. nigrolineatus and G. auritus, only a single G. nigrolineatus sample from Mozambique was included. Because our analysis includes greater geographic coverage than previous studies, we were able to evaluate the status of G. nigrolineatus. In addition to the lack of monophyly for G. nigrolineatus, the west-Central African clade differs from the East and Southern African clade by large p -distances (13.0% ND 2, 6.9 % 16 S). One individual (HB057, Arusha, Tanzania; Fig. 1) was found less than 140 km to the south-east of the approximate type locality of Gerrhosaurus flavigularis intermedia L��nnberg, 1907 (i.e. ���steppe near the Natron lakes, Kibonoto���, northern Tanzania; p. 7). Taxonomic implications are that the East/Southern African clade represents a separate species, for which the name Gerrhosaurus intermedius L��nnberg, 1907 comb. nov. is available. Loveridge (1942) relegated G. f. intermedia to the synonymy of G. n. nigrolineatus without explanation. Because of the similarity of taxa associated with the names G. flavigularis and G. nigrolineatus, the applicability of the name G. intermedius for eastern populations previously referred to G. nigrolineatus requires explanation. Although not mentioned in the text of L��nnberg���s (1907) description of G. f. intermedia, it is evident from his fig. 1 b (left side of head) that there are four supraciliaries as in G. nigrolineatus (usually five in G. flavigularis; Loveridge 1942, FitzSimons 1943). The proportions and scutellation of the head (fig. 1 a) are also very similar to FitzSimons��� (1943) fig. 157 of G. nigrolineatus. In addition, L��nnberg���s description mentions that the flank scales of G. f. intermedia are strongly keeled, and minium red in colour with dark bars extending from the back. The prefrontals are shown to be in good contact, with a long median suture (indicated in L��nnberg���s fig. 1 a). All of these features are rare or absent in G. flavigularis and often associated with G. nigrolineatus, including eastern populations that we now refer to G. intermedius (Fig. 7). In the Congo and Gabon voucher specimens (G. nigrolineatus) examined (Appendix I) there were four supraciliaries on either side of the head (e.g. PEM R 20067, Fig. 8) in all but one specimen (PEM R 20066, Congo) which had five; flanks had weakly or moderately keeled scales in the two Congo specimens, weakly (5) or moderately (4) keeled in Gabon specimens; prefrontals in broad (PEM R 20067) or moderate (PEM R 20066) contact in Congo specimens, in broad (5) to moderate (4) contact in Gabon specimens. We refer all of the above specimens to G. nigrolineatus. The vouchered Mozambique sample of G. intermedius (TM 80959) from Moebase Village had four supraciliaries on either side of the head; flanks with strongly keeled scales; and prefrontals in broad contact. Although Loveridge (1942: 511) was tempted to ���separate an eastern race��� of G. nigrolineatus, the only character he found useful was the number of longitudinal rows of dorsal scales, which numbered 24���28 in ���West Africa��� and 20���26 (but usually 22���24) in ���East Africa���. Laurent (1954) later gave a count of 26 for a specimen from Dundo in north-eastern Angola that he assigned to G. nigrolineatus. For southern Africa these counts were given as 22���24 (usually 22) by FitzSimons (1943) and 20���24 (mostly 22���23) by Jacobsen (1989). The type description of G. nigrolineatus (Hallowell 1857) refers to 25 longitudinal rows of dorsals, while the holotype of G. flavigularis intermedia has 22 such rows (L��nnberg 1907). Laurent (1964) later referred a specimen from Mayombe (lower Congo) with 25 such rows to G. n. nigrolineatus, and four specimens from Pweto in Katanga, D.R.C., with 24���26 such rows to G. n. intermedius. The number of dorsal rows varied from 23 to 25 in both the Congo (N = 2) and Gabon (N = 9) specimens examined. The vouchered southern African sample of G. intermedius (TM 80959) had 24 longitudinal rows of ventrals. While there may be average differences in these counts between western and eastern populations, there is also some overlap, and the usefulness of this feature for separating G. nigrolineatus and G. intermedius requires further investigation. According to Broadley (2007), G. nigrolineatus from Gabon and the lower Congo region has ragged dorsolateral stripes and smooth plantar scales, features which he felt may distinguish it from populations of this species elsewhere in Africa. The plantar scales of eastern populations of G. nigrolineatus (= G. intermedius) are reportedly keeled (smooth and tubercular in G. flavigularis) (FitzSimons 1943; Broadley 1966). In the Congo specimens examined, the back and flanks were olive to light brown with distinct cream, black-bordered, dorsolateral stripes, with a similarly coloured vertebral stripe that was continuous in one specimen (PEM R 20066) and broken in the other (PEM R 20067). Gabon specimens examined were light brown with scattered black and white lateral scales, and similar stripes, but the vertebral stripe was continuous in one specimen, broken in three and absent in five. As shown in Fig. 9, MBUR 02993―a specimen sampled for the current analysis―also has typical dorsolateral stripes as described above, with a broken vertebral stripe. The original description of G. nigrolineatus refers to a yellow stripe on either side of the back, bordered internally (towards the centre of the back) by a black band; and also mentions that the centre of the back contains black spots in the form of longitudinal lines (Hallowell 1857). Colour photographs of the two syntypes of G. nigrolineatus indicated that both specimens have faded somewhat, but their colour patterns were not dissimilar to the Congo and Gabon material described above. ANSP 3729 had a pair of pale (cream) dorsolateral stripes with poorly defined black borders as well as a similar vertebral stripe anteriorly (not visible beyond the nape; Fig. 10), while ANSP 8825 (juvenile) was similar but lacked a discernible vertebral stripe (Fig. 11). Donald G. Broadley (in litt. 21 March 2013) noted that a specimen of G. nigrolineatus from Ponte Denis in Gabon in the collection of the Natural History Museum, Zimbabwe (Bulawayo) had smooth plantar scales, differing somewhat from the weakly keeled plantar scales of PEM R 20067 (a detailed photographic image was used for comparison) from Republic of the Congo (Appendix I). In the Congo specimens examined, plantar scales were almost smooth or weakly keeled, while in the Gabon sample they were weakly (7) or very weakly (2) keeled. Based on photographs of one foot of each of the syntypes of G. nigrolineatus, the scales on the soles were weakly keeled. The plantar scales of the sampled specimen (TM 80959) of G. intermedius were strongly keeled, while those of 10 additional specimens from Mozambique were moderately keeled; two out of three specimens from Limpopo Province in South Africa had moderately keeled palmar scales, while one had distinctly keeled scales (Appendix I). Although there was some variation in the extent and appearance of dorsal stripes and the keeling of plantar scales, the Congo and Gabon samples (including material referred to by Broadley) are all considered conspecific and referable to G. nigrolineatus. Nevertheless, the smooth to feebly keeled plantar scales in G. nigrolineatus from Gabon and Congo is in contrast to the moderately to strongly keeled scales in populations referable to G. intermedius (e.g. FitzSimons 1943), including those from Mozambique (e.g. TM 80959 and the other specimens listed in Appendix I) as discussed above. The minium red to vermillion flanks (with pale spots or bars) of adult eastern G. nigrolineatus (= G. intermedius) differ from the light and dark barred or mostly brown flanks of G. flavigularis (see descriptions and images in Jacobsen 1989; Branch 1998; Spawls et al. 2002; Alexander & Marais 2007). It should be noted however, that according to Broadley (1966), G. flavigularis from Mozambique and adjacent parts of Zimbabwe have vermillion flanks like G. nigrolineatus (= G. intermedius), although only in areas of allopatry. The same colour pattern has been recorded in G. flavigularis from eastern Limpopo Department and eastern North West Province, South Africa, where the underside of the head is blue-grey in males (Jacobsen 1989). The possibility that such populations represent unique evolutionary lineages was not investigated in the present study, although some genetic structuring is evident within G. flavigularis (Fig. 1). According to Loveridge (1942), the scales on the flanks of G. nigrolineatus (= G. intermedius) are striated, keeled, or more-or-less smooth, whereas those of adult G. f. flavigularis are smooth. For southern African material, FitzSimons (1943) noted that the laterals of G. nigrolineatus (= G. intermedius) are keeled and sometimes feebly striated, while those of G. flavigularis are smooth or feebly keeled and striated. However, Loveridge (1942: 515) also noted that in his ���ill-defined race��� G. flavigularis fitzsimonsi (a synonym of G. flavigularis) the laterals were striated and keeled, although occasionally almost smooth, whereas the prefrontals were in broad contact. The latter two features are consistent with G. nigrolineatus. However, Loveridge (1942: 515) noted that his new subspecies had a short head (head length into SVL 4.75 times in young to 6 times in adults) as in G. f. flavigularis, and ���should not be confused with G. f. intermedia ���which, from his [L��nnberg 1907] figure, is a synonym of the long-headed G. n. nigrolineatus ���. Head length into SVL was 4.7 ���5.0 times for the two Congo specimens examined, and 4.0���5.0 times (4.8 ���5.0 for three adults with SVL> 100 mm, 4.0��� 4.6 for seven juveniles with SVL G. intermedius (TM 80959) was similar with head length into SVL 4.4 times. Therefore, we conclude that G. f. intermedia L��nnberg, 1907 is conspecific with eastern populations currently referred to G. nigrolineatus Hallowell, 1857 and which we now refer to G. intermedius. In light of the phylogenetic and morphological differences mentioned above, we suggest that populations in Gabon and lower Congo (including Kouilou region) are all, Published as part of Bates, Michael F., Tolley, Krystal A., Edwards, Shelley, Davids, Zo��, Da Silva, Jessica M. & Branch, William R., 2013, A molecular phylogeny of the African plated lizards, genus Gerrhosaurus Wiegmann, 1828 (Squamata: Gerrhosauridae), with the description of two new genera, pp. 465-493 in Zootaxa 3750 (5) on pages 476-488, DOI: 10.11646/zootaxa.3750.5.3, http://zenodo.org/record/247067

Genus Chamaeleo Laurenti 1768 Type species: Chamaeleo chamaeleon (Linnaeus 1758) Generic synonyms: Phumanola Gray 1864. Type species Chamaeleo namaquensis Smith 1831 Calyptosaura Gray 1864. Type species Chamaeleo calyptratus Dumeril & Bibron 1851 Erizia Gray 1864. Type species Chamaeleo senegalensis Daudin 1802. Species content: africanus, anchietae, arabicus, calcaricarens, calyptratus, chamaeleon, dilepis, gracilis, laevigatus, monachus, namaquensis, necasi, senegalensis, zeylanicus. This genus has a wide ranging pan African distribution extending into Europe, the Middle East, Arabia and the Indian sub-continent (Fig. 2). One species is confined to the island of Socotra. The distribution areas of the mainland species tend to be large and continuous except for Ch. anchietae which appears to have population pockets restricted to highland plateaux. Although some species may penetrate into lowland forest, or high altitude grasslands, the species of the genus generally occupy moist and dry woodland savannahs, thorn scrub, semi desert and in one species, true desert. Apart from occipital lobes in some species and prominent parietal crests in others, they have little other head ornamentation. None of them possess horns or any form of rostro-nasal or pre-orbital projections. A gular-ventral crest of single cones is found in all species being more or less developed in the various forms from very prominent in Ch. calyptratus to almost indiscernible in Ch. namaquensis. None of the species of this genus demonstrate a temporal crest. The casque is edged in a lateral parietal crest originating as a posterior continuation of the supra-orbital ridge which delineates the posterior ramus of the squamosal bone. The temporal zone is undivided. The background scalation of the flanks is generally composed of relatively homogeneous to finely heterogeneous closely packed granular tubercles. The tail of all species within this group is smooth. The plantar surfaces are smooth and claws simple. This is the only genus where the presence of tarsal spurs is seen in several of the species (Ch. arabicus, Ch. monachus, Ch. chamaeleon, Ch. necasi, Ch zeylanicus, Ch. dilepis, Ch. gracilis, Ch. calyptratus, Ch. africanus). These tend to be best developed in males and usually absent or much reduced in females. Tarsal spurs may be a synapomorphy for the genus Chamaeleo. The basic internal lung morphology consists of two large septae arising from the region of the hilum of the lung which end freely within the lung, dividing it into three chambers viz a small dorsal, a large middle and a small ventral chamber. All species possess a gular pouch and in the lung - a membrano-fibrous diaphragm that partially separates off a small dorso-cranial compartment. Many species also have several small partial septae that arise from the dorsal wall of the lung near the cranial end. The lungs are invariably adorned with varying numbers of diverticulae that trail from the inferior and posterior margins of the lung. The diverticulae vary in length and number and may be branched (Klaver 1973, 1977, 1981). Hemipenes are calyculate, with a multi rotulae arrangement of between three to five pairs of denticulated rotulae except for Ch. arabicus and Ch. namaquensis which have retained the plesiomorphic four rotulae (two pairs) configuration (Klaver & B��hme 1986). The genus is oviparous with a cyclic reproductive strategy ��� usually a single brood but up to three clutches of eggs per year in some species in ideal conditions. These species tend to have relative longevity. Females are usually sexually mature within one year and over the next few years will produce at least one clutch of eggs annually. The parietal peritoneum is unpigmented. The documented karyotype of the several species so far examined (Matthey 1931, 1957, Matthey & van Brink 1956) is 2 n= 24 = 12 M+ 12m and appears to be restricted to this genus as a synapomorphic character (Klaver & B��hme 1986), Published as part of Tilbury, Colin R. & Tolley, Krystal A., 2009, A re-appraisal of the systematics of the African genus Chamaeleo (Reptilia: Chamaeleonidae), pp. 57-68 in Zootaxa 2079 on pages 63-65, DOI: 10.5281/zenodo.187324, {"references":["Klaver, C. J. J. (1973) Lung anatomy: aid in chameleon taxonomy. Beaufortia, 20 (269), 155 - 177.","Klaver, C. J. J. (1977) Comparative lung morphology in the genus Chamaeleo Laurenti, 1768 (Sauria: Chamaeleonidae) with a discussion of taxonomic and zoogeographic mplications. Beaufortia 25 (327), 167 - 199.","Klaver, C. J. J. (1981) Lung morphology in the Chamaeleonidae (Sauria) and its bearing on phylogeny, systematics and zoogeography. Zeitschriftfuer zoologishe Systematik Evolutionsforsch, 19, 36 - 58.","Klaver, C. J. J. & Bohme, W. (1986) Phylogeny and classification of the Chamaeleonidae (Sauria) with special reference to hemipenis morphology. Bonner Zoologische Monographien, 22, 1 - 64.","Matthey, R. (1931) Chromosomes de reptiles saurians, ophidians, chelonians. L'evolution de la formule chromosomiale chez les saurians. Revue Suisse de Zoologie 38 (9), 146 - 147.","Matthey, R. (1957) Cytologie comparee et taxonomie des Chamaeleontidae (Reptilia - Lacertilia). Revue Suisse de Zoologie 64, 709 - 732.","Matthey, R. & van Brink, J. M. (1956) Note preliminaire sur la cytologie chromosomique comparee des cameleons. Revue Suisse de Zoologie 63, 241 - 246."]}

Bradypodion ngomeense sp. nov. Holotype: PEM-R 16621, an adult male collected by Devi Stuart-Fox and Adnan Moussalli in the Ngome Forest, KZN 27 �� 49 ���S, 31 �� 25 ��� E, on 6 th January 2004. Paratypes: PEM-R 16617 adult female, PEM-R 16612 adult male, PEM-R 16613 adult male, PEM- R 16615 adult female, PEM-R 16618 adult male, PEM-R 16620 adult female, PEM-R 16622 adult female, all collected by Devi Stuart-Fox and Adnan Moussalli with same locality and date as holotype; PEM-R 16842 sub-adult male, PEM-R 16843 adult female collected by Colin Tilbury 05/01/ 2002 same locality as the holotype; PEM-R 5689 adult male & PEM-R 5690 adult female, collected by Colin Tilbury 08/01/ 1999, Ngome Forest; PEM-R 17518, PEM-R 17519, PEM-R 17520, PEM-R 17521 all collected by Krystal Tolley 14 / 11 / 2005, locality, Ngome Forest. Diagnosis: A moderately large-bodied, long-tailed species of the genus Bradypodion demonstrating the characteristics of the genus (Tilbury et al. 2006). This species differs from its various congeners in the following respects: the adults have a relatively tall casque angled at (usually) over 30 degrees or more to the supra-orbital line in keeping with 6 other South African species - viz: B. damaranum (Boulenger 1887), B. dracomontanum Raw 1976, some B. ventrale (Gray 1845), B. thamnobates Raw 1976, B. transvaalense (Fitzsimons 1930) and B. nemorale Raw, 1978 (Qudeni ecomorph). It differs from B. ventrale which has a tail that is less than 50 % of its snout /vent length; from B. thamnobates which has prominent enlarged plate-like flank tubercles; from B. damaranum which has areas of ���naked��� interstitium around the axilla and along the anterior para-vertebral zone; from B. nemorale which has a relatively much reduced gular crest, and a dorsal crest of cones larger than the diameter of the eye-opening; and from B. transvaalense (sensu stricta) where most adult specimens usually have dorsal cones larger than the eye opening and in which the superior temporal zone is pale coloured and the black mid-temporal stripe continues uninterrupted over the upper and mid flank (Figure 3). Description of Holotype: Adult male, Snout/vent length 65mm, Tail 81mm. Casque sharply elevated posteriorly. Pre-orbital, supra-orbital, and temporal crests rugose and composed of thick conical and subconical tubercles. A low median parietal crest extends forwards from the apex of the casque to terminate just before the mid-orbital region. The temporal crests originate from the mid post-orbital rim, are prominent and composed of a row of 6 more or less equally enlarged tubercles on the right side and 5 on the left. Posteriorly, the temporal crest then rises almost vertically as the squamosal crest to run parallel and adjacent to the lateral parietal crest to the apex of the casque. A prominent gular crest is present, composed of 14 granulated composite lobes, numbers 2 ��� 6 overlapping and numbers 2 ��� 4 are broader than long. The free distal edges of the lobes have a denticulated fringe. Five to six thin white gular interstitial grooves are present on each side of the throat, enclosing chains of enlarged rounded tubercles. A prominent dorsal crest is present, composed of 26 conical tubercles roughly triangular in outline, extending from the nuchal area to the sacrum and then extending along the proximal third of the tail. The tubercles of the dorsal crest are highest over the anterior to mid vertebral line and lowest over the sacral region, none of the cones are taller than the diameter of the eye opening. Background scalation of heterogeneous, rounded to polygonal tubercles with no obvious interstitial venation between the tubercles. A short row of slightly enlarged flattened tubercles skirts the upper mid-flank. Enlarged tubercles are scattered along the para-vertebral zone. Belly tubercles arranged in pallisades of more or less homogeneous rounded tubercles. Variation in the Paratypes: The largest specimen, a female (PEM-R 16617) has a total length of 166mm (snout/vent 77mm + tail 89mm). Considerable variation in the gular crest is present among the types. The gular crest varies from 11 to 19 individual lobes or tubercles. The length and width of the lobes is variable, in some specimens all the lobes are longer than wide, in others the length / width may be similar and in some the overlapping lobes may be broader than long. Although overlap of between 2 to 7 lobes is common, in several specimens there is no overlap of any of the lobes. In 9 / 12 specimens, the first gular projection is a simple conical tubercle, and in all specimens, the last gular projection is a simple cone. Up to the last 3 gular ornaments may be simple cones. In most specimens the flanks appear to have no interstitial web, but in 2 / 14 specimens, an indication of interstitial webbing was present. The dorsal crest in the type series consisted of between 19 ��� 26 tubercles extending between the nuchal fold to the mid-sacrum. It may additionally extend for a variable distance along the tail from being almost imperceptible to easily discernable up to half of the length of the tail. The tail is always longer than the snout/vent length in both sexes. Sexual dimorphism: Males tend to have a more rugose scalation, better developed body and head crests, and a prominent hemipenal bulge at the base of the tail. Tail length differs between the sexes, being 53���57 % of the total length in males compared to 53���55 % in females. Colour differences between the sexes are discussed below. Colour in life: Male���Head above the upper labials mostly black ��� labials and all cranial crests pale yellow. Gular region and gular crest white. Nuchal fold - white to pale yellow. Skin of eyeball, heavily flecked with red around the periphery and a single dark horizontal stripe transects the eye. Background colour of the body is blue-green as is the dorsal crest. A bright red to orange flash over the mid flank is fully enclosed within a black flank patch which terminates abruptly about 2 / 3 rds of the way along the flank. The colour of the flash is variable from white to various shades of red. Belly, front legs and tail blue-green. Female ��� Background pale green. The superior and inferior temporal zones have white patches. The superior half of the anterior 2 / 3 rds of the flank is dark green, with scattered lighter tubercles (Figures 4 & 5). Hemipenes: None of the specimens currently present in any museum collection have fully everted hemipenes so the structure of these organs remains un-described for the present. Habitat: Northern Afrotemporate ���Mist Belt��� forest between 500 and 1360 meters a.s.l. characterized by a tall, but multi-layered canopy up to 30 meters with both Afrotemperate species and sub-tropical elements (Mucina & Geldenhuys 2006). The forest occupies the deeply incised southeast facing slopes of the Ntendeka cliffs. The largest trees in the Ngome Forest are specimens of the forest waterwood (Syzygium gerrardii) reaching up to 30 meters. Other trees include Podocarpus latifolius, Octea kenyensis, Faurea macnaughtonii, Olinia radiata and Trichocladus grandiflorus. The forest is rich in species of epiphytic orchids and hydrophilous ferns. Most specimens collected in the type series were found along forest paths and the road verges in bushes and trees at perch heights between 1���10 meters from the ground. Conservation: The entire known range of this species is restricted to the Ngome Forest. It is the largest fragment of Mist Belt forest in KwaZulu-Natal with an estimated area of 3360 Ha (36.6 km 2 - Mucina & Rutherford 2006.). Between 1876 and 1899, the forest was heavily logged and most of the larger Podocarpus and other hard wood species were removed or depleted. The forest has been protected since 1905. Much of the forest (2,636 Ha) is managed by the South African Department of Water Affairs and Forestry (DWAF) and protected within the Ntendeka Wilderness Area. Although some of the adjacent scarp area has been given over to pine plantations, there is no pine within the Wilderness Area. Etymology: The species is named after the Ngome Forest - the only forest known to harbor this taxon., Published as part of Tilbury, Colin R. & Tolley, Krystal A., 2009, A new species of dwarf chameleon (Sauria; Chamaeleonidae, Bradypodion Fitzinger) from KwaZulu Natal South Africa with notes on recent climatic shifts and their influence on speciation in the genus, pp. 43-57 in Zootaxa 2226 on pages 47-50, DOI: 10.5281/zenodo.190207, {"references":["Raw, L. R. G. (1976) A survey of the dwarf chameleons of Natal, South Africa, with the description of three new species (Sauria: Chamaeleonidae). Durban Museum Novitates, 11, 139 - 161.","Raw, L. R. G. (1978) A further new dwarf chameleon from Natal, South Africa (Sauria: Chamaeleonidae). Durban Museum Novitates, 11, 265 - 269.","Mucina, L. & Rutherford, M. C. (eds) (2006) The vegetation of South Africa, Lesotho and Swaziland. Strelitzia 19. South African National Biodiversity Institute, Pretoria."]}

Bradypodion Fitzinger 1843 Type Species: Chamaeleo pumilus Daudin 1802 Composition: B. caffer (Boettger), B. damaranum, B. dracomontanum Raw, B. gutturale (Smith), B. kentanicum (Hewitt), B. melanocephalum, B. nemorale Raw, B. occidentale, B. pumilum, B. setaroi Raw, B. taeniabronchum, B. thamnobates Raw, B. transvaalense (Fitzsimons), B. ventrale, and several as yet undescribed species (Tolley et al. 2005; Branch et al. 2006). Characterization: The monophyly of the South African Bradypodion is established by a suite of nuclear and mitochondrial genes and can also be defined by three characteristics, namely; independently derived viviparity and the associated pigmentation of the parietal peritoneum, and a specific cranial structure with a broad roof-like parietal (interpreted as a retained symplesiomorphy) bearing supra-temporal processes (interpreted as a secondary character reversal Rieppel & Crumley 1997). External morphological features that are common to all Bradypodion include the presence of heterogeneous background scalation, a midline gular crest consisting in most species of composite lobes and cones, and the absence of a ventral crest. Rostronasal processes are absent in all species. The hemipenes are calyculate with a plesiomorphic 4 -rotulae apical ornamentation. All species are viviparous. The genus may also be characterized by lung morphology, comprising simple, adiverticulate, sac-like lungs with small ridge-like septae on the cephalic, dorsal and ventral walls and with an accessory gular pouch (Beddard 1997, Klaver 1973, 1881). However, not all Bradypodion species have been assessed. Distribution: Restricted to South Africa, ranging into adjacent Swaziland, and possibly Lesotho and southern Mozambique, with introduced populations in Namibia, and occupying a wide variety of habitats., Published as part of Tilbury, Colin R., Tolley, Krystal A. & Branch, William R., 2006, A review of the systematics of the genus Bradypodion (Sauria: Chamaeleonidae), with the description of two new genera, pp. 23-38 in Zootaxa 1363 on page 34, DOI: 10.5281/zenodo.174715, {"references":["Fitzinger, L. J. (1843) Systema Reptilium Auctore Leopoldo Fitzinger. Vindobonnae (Vienna).","Tolley, K. A., Tilbury, C. R., Branch, W. R. & Matthee, C. A. (2005) The dwarfs of Africa: taxonomy, distribution and diversity of dwarf chameleons (Bradypodion sensu lato). In Abstracts, Fifth World Conference of Herpetology, Stellenbosch, South Africa, 2005, pp. 156.","Branch, W. R., Tolley, K. A., & Tilbury, C. R. (2006) A new Dwarf Chameleon (Sauria: Bradypodion Fitzinger 1843) from the Cape Fold Mountains, South Africa. African Journal of Herpetology, in press.","Rieppel, O. & Crumley, C. (1997) Paedomorphosis and skull structure in Malagasy chamaeleons (Reptilia: Chamaeleoninae), Journal of Zoology (London), 243, 351 - 380.","Klaver, C. J. J. (1973) Lung anatomy: aid in chameleon taxonomy. Beaufortia, 20, 155 - 177."]}

Nadzikambia genus nova Type Species: Chamaeleo mlanjensis Broadley 1966 Composition: Nadzikambia mlanjense Characterisation: The monophyly of Nadzikambia is established on the basis of a suite of mitochondrial and nuclear genetic characters and a unique hemipenis. The latter is rather short and stout with a short pedicel and shallow calyces. There are no rotulae on the apex, which are replaced with a pair of large fleshy, papillate lobes with scalloped edges. At the sulcal base of each lobe are one or two pedunculated papillae (Klaver & B��hme 1986). The lungs show a structure similar to that of Kinyongia, with two pairs of long diverticulae trailing from the inferior and posterior surfaces of the lung. A series of four small septae alternate with five larger septae across the dorsal wall with several smaller septae arising from the ventral and cephalic walls. There is no gular pouch (Klaver 1977). The parietal peritoneum is unpigmented and reproduction oviparous. The external morphology is largely conservative, and gular and ventral crests are absent. There is a weak dorsal crest, finely heterogeneous scalation that forms rosettes of tubercles on the lower flanks, and a low casque. Although cranial morphology remains undescribed, the external appearance of the cranial crests suggest that the cranium is constructed in the same manner as Kinyongia. Distribution: Only known from sub-montane forest habitats in a few scattered localities on the Mulanje Massif in southern Malawi, Central Africa. Etymology: The name is derived from ���Nadzikambe���, the name for chameleon in ChiChewa, the language used by the tribe that lives in the area around Mulanje Mountain in southern Malawi. The name Nadzikambe is Latinised by terminating it with the suffix ia thus giving it a feminine gender., Published as part of Tilbury, Colin R., Tolley, Krystal A. & Branch, William R., 2006, A review of the systematics of the genus Bradypodion (Sauria: Chamaeleonidae), with the description of two new genera, pp. 23-38 in Zootaxa 1363 on pages 35-36, DOI: 10.5281/zenodo.174715, {"references":["Klaver, C. J. J. & Bohme, W. (1986) Phylogeny and classification of the Chamaeleonidae (Sauria) with special reference to hemipenis morphology. Bonner zoologische Monographisce 22, 1 - 64."]}

Kinyongia genus nova Type Species: Chamaeleo fischeri fischeri Reichenow 1887. Composition: K. adolfifriderici, K. carpenteri, K. excubitor, K. fischeri fischeri, K. fischeri multituberculatum (Nieden), K. fischeri uluguruense (Loveridge), K. tavetanum, K. tavetanum boehmei (Lutzman & Nečas), K. uthmoelleri, K. xenorhinum, K. oxyrhinum, and K. tenue Characterization: The monophyly of Kinyongia is established on the basis of a suite of nuclear and mitochondrial genetic characters. No morphological synapomorphy is known to define all members. Cranial structure has only been studied in K. fischeri (Rieppel & Crumley 1997). The parietal is reduced to a narrow posteriorly projecting sagittal process that meets the ascending squamosal processes at the apex of the casque to completely enclose the temporal fossa. This derived condition is similar to that found in the genera Chamaeleo, Furcifer and Calumma (Rieppel 1981, 1987, Rieppel & Crumley 1997). Scalation is generally of finely heterogeneous granules or flattened polygonal tubercles. In those species that are characterized by a head ornamentation of fused rostronasal projections (carpenteri, xenorhinum, tenue and oxyrhinum), the scalation is generally an unordered heterogeneous mix of tubercles. In species with paired rostronasal projections (fischeri, tavetanum, uthmoelleri) the flanks are adorned with tubercles clustered into “rosettes”, especially on the lower flanks. This rosetting is also seen in the hornless species excubitor. Plantar surfaces are smooth and claws are simple. None of the species have midline gular or ventral crests, occipital lobes, or annulated horns. Cranial ornamentation in some species, e.g. paired rostronasal blade-like horns in fischeri and tavetanum, and fusion of the canthal ridges into a single vertically flattened process in carpenteri, xenorhinum, tenue and oxyrhinum, are similar to features found in the Malagasy genera Calumma and Furcifer. Lung structure is relatively plesiomorphic. They are similar to those of Bradypodion and Malagasy Calumma and Furcifer, being generally simple with a number of small septae on the dorsal, cephalic and ventral walls. The lungs of Kinyongia appear to lack the accessory gular pouch and usually have trailing diverticulae from the posteroinferior surface of the lung (tavetanum, fischeri, tenue, and adolfifriderici) although these are lacking in K. xenorhinum (Klaver 1977, 1981). The lungs in the rest of the species of Kinyongia have not as yet been described. The hemipenes are calyculate with a plesiomorphic 4 rotulae apical ornamentation, and all the species are oviparous. Distribution: Distributed in East Africa with the most westerly species, K. adolfifriderici, extending into the eastern DRC, and K. excubitor reaching as far north as Mount Kenya. They are confined to tropical/sub-tropical forest biomes, often in relict montane or sub-montane forests (Fig. 1). Etymology: This genus is largely confined to the three countries that make up the central east African region namely Kenya, Tanzania and Uganda. The lingua franca for this region is Swahili. The name derives from the generic Swahili name for chameleon “Kinyonga” and identifies it as a genus that is largely confined to Swahili speaking countries. The name is Latinized by terminating the name spelling with the letters ia giving it a feminine gender. Thus the specific names remain unaltered.