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Micrelapidae new family. Type genus: Micrelaps Boettger, 1880. Type species: Micrelaps muelleri Boettger, 1880. Etymology: Boettger (Bottger ¨) did not give the etymology for the generic nomen but was almost certainly from the Latin adjective micro-, derived from the Greek mikros (small), and elaps, the Latinised form of the Greek noun´ellops or´elaps (literally sea-fish or serpent, but here in reference to the snake genus Elaps, now a synonym of Homoroselaps). Micrelapidae fam. nov. is derived from Micrelaps by the taking the stem elap- of the root word of the nomen. Content: Micrelaps muelleri Boettger, 1880, Micrelaps bicoloratus Sternfeld, 1908, Micrelaps vaillanti Mocquard, 1888, Brachyophis revoili Mocquard, 1888. Diagnosis and definition: In the crania of Micrelaps and Brachyophis we examined the ectopterygoid was laterally and medially expanded at the point of contact with the pterygoid, with this expansion not being contiguous with the ectopterygoid anterolateral and anteromedial lobes (Fig. 4, Supplementary material fig. 53, 54). The lateral expansion is a posterolaterally and somewhat ventrally directed, very prominent protuberance continuous with a ridge on the ventral surface of the pterygoid. This character state was not present in any other cranium we examined and is very likely a synapomorphy of the family. Other common cranial characters include a premaxilla adapted for a fossorial lifestyle, premaxillary transverse processes closely approaching the maxilla, a short maxilla with ascending processes abutting the prefrontal, well-developed, grooved fangs below the orbit, preceded by a diastema and 2 – 3 teeth, an ectopterygoid deeply forked into anterolateral and anteromedial lobes that articulate with maxillary ectopterygoid processes leaving a foramen in the middle, prefrontal and parietal supraorbital processes laterally bordering the frontal and almost meeting each other, a tendency towards fusion of cranial bones (especially because the supratemporal is absent, very likely fused to the quadrate in Brachyophis and to posterior chondrocranial elements in Micrelaps), and a short quadrate. Brachyophis, however, differs from the type genus in possessing a postorbital (versus postorbital absent in Micrelaps), dorsolateral adductor ridges on the parietal (versus a single sagittal ridge in Micrelaps), only a faint pseudocoronoid ridge on the dentary (versus a prominent process in Micrelaps). Scalation characters that are common in both genera include 1 nasal, 7 supralabials, 15 smooth dorsal scale rows, absence of a loreal, 2 anal shields. Ventrals range from 170 to 280 and subcaudals (paired) 16 – 32 in Micrelaps (Boulenger, 1896; De Witte and Laurent, 1947; Rasmussen, 2002; Werner et al., 2006; Spawls et al., 2018). In Brachyophis, ventrals range from 103 to 123 and subcaudals (single) 8 – 14 (De Witte and Laurent, 1947; Lanza, 1966). Brachyophis has a large, azygous occipital shield (Boulenger, 1896). Micrelaps and Brachyophis possess a rectal caecum and a short genital sinus in the female, two soft tissue traits used to cluster these two genera by Underwood and Kochva (1993). Distribution: Micrelaps spp. is distributed in eastern and northeastern Africa and western Asia. Brachyophis is limited to Somalia in north-eastern Africa. Distribution: Micrelaps spp. is distributed in eastern and northeastern Africa and western Asia. Brachyophis is limited to Somalia in north-eastern Africa. Remarks: Geniez (2018) commented that Micrelaps “could constitute a separate family within its own right, that of Micrelapsidae”. Bar et al. (2021) likewise wrote that “. The actual placement of the genus [Micrelaps] is often poorly supported within studies and inconsistent across them. We suspect it will soon be placed in its own family — as is the norm in taxonomy these days. We predict this family, containing a single genus (Micrelaps), will be called Micrelapidae.”. However, these authors did not explicitly express that they are erecting a new family for these snakes. Rather, it was a suggestion about what should/could be done. It therefore is not in accordance with Article 16.1 and Recommendation 16A of The Code (ICZN, 1999). They also did not also provide characters for the express purpose of differentiating or diagnosing “Micrelapsidae” or “ Micrelapidae ”, nor did they cite a work containing the same (again, very likely because a nomenclatural act presumably was not the intention of Geniez [2018] and it was not the intention of SM, who wrote this in Bar et al. [2021] either) and this contravenes Articles 13.1.1, 13.1.2 and Recommendation 13A of The Code. Hence, we regard the nomen “Micrelapsidae” as unavailable. The ZooBank LSID for this taxonomic action is urn:lsid:zoobank.org:pub:D8475246-AD8E-4886-AB55- 12F6F242E9C4., Published as part of Das, Sunandan, Greenbaum, Eli, Meiri, Shai, Bauer, Aaron M., Burbrink, Frank T., Raxworthy, Christopher J., Weinell, Jeffrey L., Brown, Rafe M., Brecko, Jonathan, Pauwels, Olivier S. G., Rabibisoa, Nirhy, Raselimanana, Achille P. & Merila, Juha, 2023, Ultraconserved elements-based phylogenomic systematics of the snake superfamily Elapoidea, with the description of a new Afro-Asian family, pp. 1-11 in Molecular Phylogenetics and Evolution 180 on pages 8-9, DOI: 10.1016/j.ympev.2022.107700, http://zenodo.org/record/7746501, {"references":["Sternfeld, R., 1908. Zur Schlangenfauna Ostafrikas. I. Schlangen aus Sud-Abessinien. Mitt. Zool. Mus. Berlin 4, 239 - 247.","Mocquard, F., 1888. Sur une collection de Reptiles et de Batraciens rapport´es des Pays Comalis et de Zanzibar par M. G. R´evoil. M´emoires Publies par la Soci´ete´ Philomathique a l' occasion du Centenaire de sa fondation 1788 - 1888, 109 - 134.","Boulenger, G. A., 1896. Catalogue of the Snakes in the British Museum (Natural History). Volume III, containing the Colubridae (Opisthoglyphae and Proteroglyphae), Amblycephalidae, and Viperidae. Trustees of the British Museum, London.","de Witte, G. - F., Laurent, R., 1947. R´evision d' un groupe de Colubridae africains: genres Calamelaps, Miodon, Aparallactus et formes affines. M´em. Mus. R. His. Nat. Belg. 29, 1 - 134.","Rasmussen, J. B., 2002. A review of the African members of the genus Micrelaps Boettger 1880 (Serpentes Atractaspididae). Tropical Zoology 15, 71 - 87.","Werner, Y. L., Babocsay, G., Carmely, H., Thuna, M., 2006. Micrelaps in the southern Levant: variation, sexual dimorphism, and a new species (Serpentes: Atractaspididae). Zool. Middle East 38, 29 - 48.","Spawls, S., Howell, K., Hinkel, H., Menegon, M., 2018. Field Guide to East African Reptiles, second edition. Bloomsbury Wildlife, London.","Lanza, B., 1966. Il genere Brachyophis e descrizione di una nuova forma (Reptilia, Serpentes, Colubridae). Monit. Zool. Ital. 74, 30 - 48.","Underwood, G., Kochva, E., 1993. On the affinities of the burrowing asps Atractaspis (Serpentes: Atractaspididae). Zool. J. Linn. Soc. 107, 3 - 64.","Geniez, P., 2018. Snakes of Europe, North Africa and the Middle East: A Photographic Guide. Princeton University Press, Princeton and Oxford.","Bar, A., Haimovitch, G., Meiri, S., 2021. Field guide to the amphibians and reptiles of, Israel. Edition. Chimaira, Frankfurt Am Main.","ICZN (International Commission on Zoological Nomenclature), 1999. International Code of Zoological Nomenclature, fourth edition. International Trust for Zoological Nomenclature, London."]}

Afrixalus lacustris Greenbaum, Dehling, Kusamba & Portik sp. nov. Great Lakes Spiny Reed Frog urn:lsid:zoobank.org:act: 41D3C41A-4501-4BEC-BFB6-B9C6C1518DC3 Afrixalus laevis (nec Megalixalus laevis Ahl 1930)— Laurent 1950: 24; Laurent 1972: 59 (partim); Laurent 1982: 33 (partim); Schiøtz 1999: 56 (partim); Channing & Howell 2006: 137 (partim); Spawls et al. 2006: 183 (partim); Channing & Rödel 2019: 158 (partim). Holotype. UTEP 20805 (field no. ELI 605), adult male, from the vicinity of Kalundu (03.15552° S, 28.42108° E, 1482 m), South Kivu Province, DRC, collected on 21 December 2010 by Chifundera Kusamba and Félix I. Alonda (Figs. 8D–F, 12A–B). Paratopotype. UTEP 20806 (field no. ELI 606), adult male, collected with the holotype. Referred specimens. Specimens with an asterisk were not included in morphological analyses due to poor quality of preservation, lack of a voucher, or unavailability of specimen(s): UTEP 20807–08 (field nos. ELI 644, 649), vicinity of Kalundu (03.15552° S, 28.42108° E, 1482 m), South Kivu Province, DRC; UTEP 20809, 22422 (field nos. ELI 669–70), Baraka, near shore of Lake Tanganyika (04.10832° S, 29.09705° E, 777 m), South Kivu Province, DRC; UTEP 20810–11 (field nos. EBG 1316, 1319), vicinity of Irangi (01.8873° S, 28.4495° E, 820 m), South Kivu Province, DRC; UTEP 22423 (field no. MUSE 10192), Kahuzi-Biega National Park, Nanwa (02.362138° S, 28.192589° E, 1566 m), South Kivu Province, DRC; UTEP 22424 (field no. MUSE 10137), Itombwe Plateau, Mbandakila (03.47937° S, 28.413049° E, 1135 m), South Kivu Province, DRC; RMCA 77-020 -B-133–136* (four specimens), Itombwe Plateau, Tubutubu (2300 m), South Kivu Province, DRC; RMCA 77-020 -B-89–92 (field nos. MF 14–15 [two specimens]), Kitutu (ca. 03.28° S, 28.11° E, 700 m), South Kivu Province, DRC; RMCA 77-020 - B-0069–71 (1 specimen), Itula (ca. 03.27° S, 28.12° E, 650 m), South Kivu Province, DRC; RMCA 77-020 -B-0081–83, 77-20-B-0086, Kamituga (ca. 03.54° S, 27.69° E, 1050 m), South Kivu Province, DRC; RMCA 77-020 - B-0072–73 (1 specimen), Mwana (ca. 03.15° S, 28.47° E, 1650 m), South Kivu Province, DRC; UTEP 22417 (field no. CRSN 2773), Toyokana (02.02734° N, 030.06653° E, 1294 m), Ituri Province, DRC; (field no. EPLU 395 *, photo and tissue only), ca. 0.5 km E of Epulu, near Mt. Mbiya (01.39594° N, 28.58093° E, 755 m), Ituri Province, DRC; UTEP 22416 (field no. DFH 247), Bwindi Impenetrable National Park, Buhoma (00.99045° S, 29.61884° E, 1523 m), Western Region, Uganda; (field nos. DFH 1102–03 * photos and tissues only), Kibale Forest National Park, Ngogo Research Center (00.49795° N, 30.42262° E, 1363 m), Western Region, Uganda; CAS 202032–36, Bwindi Impenetrable National Park, Buhoma rd., 2 km S (by rd.) of Bizenga River (01.00975° S, 29.620694° E), Western Region, Uganda; CAS 202133, Bwindi Impenetrable National Park, small tributary to Ishasha River (00.905° S, 29.704972° E, 1280 m), Western Region, Uganda; CAS 202109, Bwindi Impenetrable National Park, Bizenga River at Buhoma rd. (00.99275° S, 29.615722° E), Western Region, Uganda; CAS 256035 *, Bwindi Impenetrable National Park, rd. north of Ruhija, Western Region, Uganda; CAS 256128–31 *, Mabira Forest Reserve, marshy area adjacent to unpolluted stream (0.4508° N, 32.9493° E, 1121 m), Mukono District, Central Region, Uganda. Diagnosis. The species is referred to the genus Afrixalus for exhibiting the following characteristics: fingers and toes webbed; tips of fingers and toes enlarged to discs; eye large; pupil vertically elliptical; tympanum indistinct; vomer ridges and teeth absent; dorsal surfaces finely shagreened with minute pointed tubercles; outer metatarsal tubercle distinct; gular gland in males. It is readily distinguished from most other members of the genus by its small size (SVL in males 18.9–22.2 mm, in females 20.8–25.7 mm), being smaller than A. dorsalis (males 25–28 mm, females 26–30 mm), A. fornasini (males 30–38 mm, females 35–40 mm), A. fulvovittatus (males 23–27 mm, females 25–28 mm), A. lacteus (males 22–27 mm, females 25–29 mm), A. leucostictus (males 27–32 mm, females 25–36 mm), A. manengubensis (males to 32 mm), A. nigeriensis (males 28–34 mm, females 32–35 mm), A. osorioi (males 27–31, females 32–35 mm), A. paradorsalis (males 28–34 mm, females 32–35 mm), A. septentrionalis (males 19–21 mm), A. vittiger (males 22–25 mm, females 25–28 mm), and A. wittei (males 27–30 mm, females 29–33 mm); and larger size than A. delicatus (males 15–19 mm, females 17–22 mm) and A. stuhlmanni (males 15–21 mm, females 17–25 mm). Its differs in dorsal coloration and pattern from all species with longitudinal stripes or bands on a yellowish-brown background (A. brachycnemis, A. crotalus, A. delicatus, A. dorsalis, A. enseticola, A. fornasini, A. fulvovittatus, A. knysnae, A. morerei, A. orophilus, A. quadrivittatus, A. schneideri, A. spinifrons, A. stuhlmanni, A. upembae, A. septentrionalis, A. vittiger, and A. wittei); species showing a conspicuous large dark dorsal blotch on a uniform background on anterior dorsum in combination with large blotches on both sides of the hip (A. leucostictus, A. manengubensis, A. osorioi, A. paradorsalis, and A. schneideri) or with longitudinal lateral dark bands (A. equatorialis and A. nigeriensis); species with a uniformly colored dorsum without conspicuous pattern except a dark dorsolateral stripe (A. aureus, A. clarkei, A. delicatus, A. fornasini, A. lacteus, A. leucostictus [dorsum with small white tubercles], A. uluguruensis, and A. weidholzi [might have a black vertebral line and dark brown flanks]); and from all remaining species having either dark stripes running from tip of snout to back, crossing each other and continuing as dorsolateral stripes (A. vibekensis, also males without asperities); anterior part of dorsum yellowish with brown pattern, posterior part of dorsum translucent (A. laevis, also males without asperities); posterior part of dorsum and dorsal sides of limbs semi-translucent with dense small dark white-edged speckles (A. dorsimaculatus); and dark blotches and transverse marks with white speckles (A. sylvaticus). It is most similar to A. phantasma sp. nov. but differs from this species in being smaller (Table 1) and showing an overall darker coloration and more contrasting dorsal pattern that is obvious even in preserved specimens (Figs. 7–8, 12). It is readily distinguished from A. phantasma sp. nov. by the more extensive hand and toe webbing (Fig. 8E–F). Description of holotype. Measurements of the holotype are provided in Table 2. Body very slender, widest at temporal region, slightly tapering to groin (Fig. 8D–F); head small (HL/SVL 0.31, HW/SVL 0.31), about as long as wide (HW/HL 0.99); snout relatively long, rounded in dorsal view and in lateral profile, slightly wider than long; canthus rostralis hardly discernible, slightly concave between eye and nostril in dorsal view; loreal region oblique; nostrils rounded, directed anterolaterally and slightly dorsally; situated much closer to tip of snout and to eye, separated from each other by distance about equal to distance between eye and nostril (IN/EN 1.02); eye directed anterolaterally, strongly protruding, very large (ED/HL 0.39), its diameter shorter than snout (ED/SL 0.90); interorbital distance much wider than upper eyelid (IO/EW 1.61) and wider than internarial distance (IO/IN 1.31); tympanum covered by skin, not visible externally; upper jaw with dentition; teeth on premaxilla larger than teeth on maxilla; choanae small, located far anterolaterally at margins of roof of mouth, its anterior edge covered by maxillary bone, therefore appearing semicircular in ventral view; vomer ridges and teeth absent; tongue heartshaped, longer than broad, bilobed for about one-fourth of its length, free distally for about two-thirds its length; densely covered with minute papillae; median lingual process absent. Dorsal surfaces of head, trunk, and limbs and lateral surfaces of trunk finely shagreened; tiny pointed, widely scattered tubercles on dorsum, more numerous posteriorly than anteriorly; supratympanic fold absent; few tubercles at rear end of mandible without pointed tips; ventral side of head smooth; vocal sac present; gular gland of vocal sac smooth, large and wide, covering about 70 percent of throat width (Fig. 8E); chest smooth, abdomen weakly areolate; ventral side of limbs smooth; short transverse fold above vent. Forelimbs slender; hand large (HaL/SVL 0.28); tips of fingers enlarged into large disks, each with circummarginal groove; relative length of fingers: I I 2-/2 II 2-/3- III 2.25/2 IV; thenar tubercle oval, small and very low, about one-fourth length of metacarpal of Finger I; inner palmar tubercle and outer palmar tubercle very low, small, rounded, almost indiscernible. Hind limbs slender, moderately long; heel reaching to level of eye when legs adpressed forwardly to body; crus moderately long (TibL/SVL 0.44), slightly shorter than thigh; heels not touching each other when knees flexed and thighs held perpendicularly to median plane; foot subequal in length to crus (FoL/TibL 0.94); relative length of toes: I I 1.5/2+ II 1+/2+ III 1+/2+ IV 2/1+ V (Fig. 8E); inner metatarsal tubercle moderately prominent, elongated, about half length of metatarsus of Toe I; outer metatarsal tubercle small, rounded. Coloration in life. During night, skin on dorsal side of head, trunk, forelimbs, crus and tarsus light brown with more or less regular, locally more densely clustered dark brown speckling; distinct dark brown transverse stripe between upper eyelids, another less distinct one in scapula region, and another narrow one at level of anterior end of pelvis; dorsal skin of thigh largely unpigmented with only a broad stripe of widely scattered brown pigmentation; distinct dark brown band along canthus, continuing behind eye on both sides of trunk to level of pelvis; tubercles below eye and in tympanic region at rear end of jaw white; dark brown stripe running diagonally on middle of tibia, forming interrupted, inverted U-shaped band together with pelvic spots when legs folded against body; skin of ventral side of head, trunk and limbs largely unpigmented and translucent; gular gland bright yellow; fingers and toes yellow (Fig. 12). During day, basic dorsal coloration light brown; dark brown dorsal speckles, spots and stripes in more pronounced contrast to basic coloration, more distinctly visible. Coloration in preservative. Dorsal basic coloration largely faded to light brown; darker dorsal pattern elements dark brown, clearly visible; color of gular gland, fingers and toes faded to white (Fig. 8D–F). Variation. The paratypes match the holotype in general appearance and proportions. Coloration and color pattern varies from light-cream to yellowish-orange with a few relatively large and darker blotches and vermiculation, light speckling and indistinct dorsolateral stripe to an almost reticulated pattern of contrasting bright and dark elements with a broad and conspicuous brown dorsolateral band (Fig. 12). Pedal webbing variation is I 1.5[100]/2+[100] II 1+[100]/2+[100] III 1 +[50],1.25[50]/2+[100] IV 2 [100]/1+[100] V. Bioacoustics. The advertisement call and other vocalizations are unknown. Ecology and natural history. We collected males and females in swamps in forest openings and near forest edges (Fig. 11B), from vegetation at the edge of streams (or on vegetation overhanging them) in forest, and from non-forest vegetation near the shore of Lake Tanganyika. Egg deposition has not been observed and tadpoles are unknown. An adult male (CAS 256035) was found in Bwindi Impenetrable National Park (Uganda) ca. 2 m above ground in a tree fern “either calling or moving to a calling site (similar to ‘running’ of Kassina)” (D. Blackburn, in litt., 22 July 2021). Laurent (1955) noted the species from transitional forest. In his paper on amphibians of Virunga National Park, Laurent (1972) noted the species is most commonly found in swamps with herbaceous vegetation in secondary forest, and more rarely, in primary forest. Laurent (1982) noted that, in general, the species is common in the foliage of shrubs in dense forests. He also recorded it from “une mare sombre” (i.e., a dark pond) in syntopy with Chiromantis at Kitutu (DRC), marshes where it reproduces by sticking its eggs under the leaves of fountain grass (Pennisetum, Shabunda, DRC), a shaded bank of a pond in syntopy with Hyperolius “ tuberculatus ” (likely H. hutsebauti sensu Bell et al. 2017) and H. frontalis; the center of this pond had reeds harboring H. kivuensis and Afrixalus orophilus (Shabunda, DRC). Laurent (1983:352) listed the species (along with Phrynobatrachus petropedetoides and Hyperolius frontalis) from “shadowy puddles of the thick transition forest.” Etymology. The species epithet is the Latin adjective “ lacustris,” meaning belonging to or dwelling in lakes; in allusion to the distribution of the new species in the region of the African Great Lakes. Distribution and conservation. The species is distributed from lowland rainforest of the Congo Basin at 460 m (Omaniundu, DRC, Laurent 1982) to transitional forests of the Albertine Rift at 1650 m (Mwana, DRC) and east as far as Mabira Forest, Uganda (1121 m). As shown in Figure 5, the species has a relatively large distribution, including several national parks and protected areas (e.g., Virunga National Park, Bwindi Impenetrable National Park), and thus is not likely to be threatened based on a limited distribution. Using GeoCAT, its extent of occurrence is estimated as 409,238 km ² and its area of occupancy as 160 km ² (Bachman et al. 2011). We therefore classify this species as Least Concern according to the IUCN Red List criteria (IUCN 2021). Four specimens (RMCA 77-020-B-133–136) from Tubutubu, Itombwe Plateau (2300 m) are problematic because they are morphologically consistent with A. lacustris, but they were collected from a substantially higher elevation than all other known specimens. Coordinates for Tubutubu provided by RMCA (4° S, 28.933° E) suggest the locality is east of the highest point of the plateau at an elevation of about 1400 m on Google Earth. Although it is possible that the specimens were collected at the latter elevation, Laurent (1982) mentions bamboo forest as the habitat, which usually occurs at the highest elevations of the Itombwe Plateau (Doumenge 1998). No clarity is offered by Laurent (1964), because he does not mention any Afrixalus in his study of the ecology and distribution of amphibians of the Itombwe Plateau. Given this conflicting information, we do not include the Tubutubu specimens in our current understanding of the elevational distribution of A. lacustris (see gray triangle in Fig. 5)., Published as part of Greenbaum, Eli, Portik, Daniel M., Allen, Kaitlin E., Vaughan, Eugene R., Badjedjea, Gabriel, Barej, Michael F., Behangana, Mathias, Conkey, Nancy, Dumbo, Bonny, Gonwouo, Legrand N., Hirschfeld, Mareike, Hughes, Daniel F., Igunzi, Félix, Kusamba, Chifundera, Lukwago, Wilber, Masudi, Franck M., Penner, Johannes, Reyes, Jesús M., Rödel, Mark-Oliver, Roelke, Corey E., Romero, Soraya & Dehling, J. Maximilian, 2022, Systematics of the Central African Spiny Reed Frog Afrixalus laevis (Anura Hyperoliidae), with the description of two new species from the Albertine Rift, pp. 201-232 in Zootaxa 5174 (3) on pages 218-223, DOI: 10.11646/zootaxa.5174.3.1, http://zenodo.org/record/6986207, {"references":["Ahl, E. (1930) Ueber die afrikanischen Arten der Baumfroschgattung Megalixalus. Sitzungsberichte der Gesellschaft Naturforschender Freunde zu Berlin, 1930, 89 - 102.","Laurent, R. (1950) Genres Afrixalus et Hyperolius (Amphibia Salientia). Exploration du Parc National Albert, Mission G. F. De Witte (1933 - 1935), 64, 5 - 120, 7 pls.","Laurent, R. F. (1972) Amphibiens. Exploration du Parc National des Virunga, Deuxieme Serie, 22, 1 - 125, 11 pls.","Laurent, R. F. (1982) Le genre Afrixalus Laurent (Hyperoliidae) en Afrique Centrale. Annales du Musee Royal de l'Afrique Centrale, Serie in Octavo, Science Zoologique, 235, 1 - 93.","Schiotz, A. (1999) Treefrogs of Africa. Edition Chimaira, Frankfurt am Main, 350 pp.","Channing, A. & Howell, K. M. (2006) Amphibians of East Africa. Cornell University Press, Ithaca, New York, New York, xi + 418 pp.","Spawls, S., Howell, K. M. & Drewes, R. C. (2006) Pocket Guide to the Reptiles and Amphibians of East Africa. A & C Black Publishers, London, 240 pp.","Channing, A. & Rodel, M. - O. (2019) Field Guide to the Frogs & Other Amphibians of Africa. Struik Nature, Cape Town, 408 pp.","Laurent, R. F. (1955) Apercu de la biogeographie des batraciens et des reptiles de la region des grand lacs. Bulletin de la Societe Zoologique de France, 79, 290 - 310.","Bell, R. C., Parra, J. L., Badjedjea, G., Barej, M. F., Blackburn, D. C., Burger, M., Channing, A., Dehling, J. M., Greenbaum, E., Gvozdik, V., Kielgast, J., Kusamba, C., Lotters, S., McLaughlin, P. J., Nagy, Z. T., Rodel, M. - O., Portik, D. M., Stuart, B. L., VanDerWal, J. & Zamudio, K. R. (2017) Idiosyncratic responses to climate-driven forest fragmentation and marine incursions in reed frogs from Central Africa and the Gulf of Guinea Islands. Molecular Ecology, 26 (19), 5223 - 5244. https: // doi. org / 10.1111 / mec. 14260","Laurent, R. F. (1983) About the herpetofauna of Central African montane forest. In: Rhodin, A. G. J. & Miyata, K. (Eds.), Advances in Herpetology and Evolutionary Biology: Essays in Honor of Ernest E. Williams. Museum of Comparative Zoology, Cambridge, Massachusetts, pp. 350 - 358.","Bachman, S., Moat, J., Hill, A., de la Torre, J. & Scott, B. (2011) Supporting Red List threat assessments with GeoCAT: Geospatial Conservation Assessment Tool. ZooKeys, 150, 117 - 126. https: // doi. org / 10.3897 / zookeys. 150.2109","IUCN (2021) IUCN Red List Categories and Criteria: Version 2021 - 3. Available from: https: // www. iucnredlist. org / (accessed 25 May 2022)","Doumenge, C. (1998) Forest diversity, distribution, and dynamique in the Itombwe Mountains, South-Kivu, Congo Democratic Republic. Mountain Research and Development, 18 (3), 249 - 264. https: // doi. org / 10.2307 / 3674036","Laurent, R. F. (1964) Adaptive modifications in frogs of an isolated highland fauna in Central Africa. Evolution, 18 (3), 458 - 467. https: // doi. org / 10.1111 / j. 1558 - 5646.1964. tb 01622. x"]}

Afrixalus phantasma Dehling, Greenbaum, Kusamba & Portik sp. nov. Ghost Spiny Reed Frog urn:lsid:zoobank.org:act: 1B2DBF26-5B0A-4DA8-B65E-0BC01704C1B4 Afrixalus laevis (nec Megalixalus laevis Ahl 1930)— Laurent 1972: 59 (partim); Laurent 1982: 33 (partim); Schiøtz 1999: 56 (partim); Channing & Howell 2006: 137 (partim); Spawls et al. 2006: 183 (partim); Channing & Rödel 2019: 158 (partim). Holotype. ZFMK 103454 (field no. JMD 723), adult male, from Gishwati Forest (01.823745° S, 29.360373° E, 2084 m), now part of Gishwati-Mukura National Park, Western Province, Rwanda, collected on 5 April 2011 by J. Maximilian Dehling and Bonny Dumbo (Figs. 7A, 8A–C). Paratypes. ZFMK 103455 (field no. JMD 722), adult female, collected with the holotype; ZFMK 103456 (field no. JMD 2015-30), adult male, ZFMK 103457 (field no. JMD 2015-31), adult female, from the type locality in Gishwati Forest, both collected on 27 September 2015 by J. Maximilian Dehling and Bonny Dumbo; ZFMK 103458–60 (field nos. JMD 677–679), three adult males, collected on 19 March 2011 by J. Maximilian Dehling, ZFMK 103461–62 (field nos. JMD 954–955), two adult males, collected on 16 February 2013 by J. Maximilian Dehling, all from Kamiranzovu Swamp (02.477165° S, 29.158243° E, 1962 m), Nyungwe National Park, Western Province, Rwanda; UTEP 20791–20792 (field nos. EBG 2838, 2843), two adult males, collected on 21 December 2009 by Chifundera Kusamba, Wandege M. Muninga, Mwenebatu M. Aristote, and Maurice Luhumyo from Nyakasanza Swamp near Tshibati (02.22886° S, 28.78017° E, 1979 m), South Kivu Province, DRC. Referred specimens. UTEP 20802 (field no. EBG 1198), forest ca. 4 km NW of Lwiro (02.2226° S, 28.7754° E, 2077 m), South Kivu Province, DRC; UTEP 20803, 22418–20 (field nos. EBG 1232, 1238–40), Kahuzi-Biega National Park, Mugaba (02.2750° S, 28.6631° E, 2298 m), South Kivu Province, DRC; UTEP 20804 (field no. EBG 1283), Kahuzi-Biega National Park, Chinya (02.2671° S, 28.6455° E, 2267 m), South Kivu Province, DRC; UTEP 20793–94 (field nos. EBG 2844–45), Nyakasanza Swamp near Tshibati (02.22886° S, 28.78017° E, 1979 m), South Kivu Province, DRC; UTEP 20795–20800, 22421 (field nos. ELI 414–20), Nyakasanza Swamp near Tshibati (02.22829° S, 28.77972° E, 1979 m), South Kivu Province, DRC; UTEP 20801 (field no. ELI 425), Chanjoka (02.21261° S, 28.77644° E, 2115 m), South Kivu Province, DRC. Diagnosis. The species is referred to the genus Afrixalus by exhibiting the following characteristics: fingers and toes webbed; tips of fingers and toes enlarged to discs; eye large; pupil vertically elliptical; tympanum indistinct; vomer ridges and teeth absent; dorsal surfaces finely shagreened with minute pointed tubercles; outer metatarsal tubercle distinct; gular gland in males. It is readily distinguished from most other members of the genus by its small size (SVL in males 20.1–24.6 mm, in females 22.6–26.4 mm), being smaller than A. dorsalis (males 25–28 mm, females 26–30 mm), A. fornasini (males 30–38 mm, females 35–40 mm), A. lacteus (males 22–27 mm, females 25– 29 mm), A. leucostictus (males 27–32 mm, females 25–36 mm), A. manengubensis (males to 32 mm), A. nigeriensis (males 28–34 mm, females 32–35 mm), A. osorioi (males 27–31 mm, females 32–35 mm), A. paradorsalis (males 28–34 mm, females 32–35 mm), A. septentrionalis (males 19–21 mm), and A. wittei (males 27–30 mm, females 29–33 mm); and larger than A. brachycnemis (males 18–21 mm, females 20–22 mm), A. delicatus (males 15–19 mm, females 17–22 mm), A. spinifrons (males to 20 mm, females to 25 mm), and A. stuhlmanni (males 15–21 mm, females 17–25 mm). Its differs in dorsal coloration and pattern from all species with longitudinal stripes or bands on a yellowish-brown background (A. brachycnemis, A. crotalus, A. delicatus, A. dorsalis, A. enseticola, A. fornasini, A. fulvovittatus, A. knysnae, A. morerei, A. orophilus, A. quadrivittatus, A. schneideri, A. spinifrons, A. stuhlmanni, A. upembae, A. septentrionalis, A. vittiger, and A. wittei); species with a conspicuous large dark dorsal blotch on a uniform background on anterior dorsum in combination with large blotches on both sides of the hip (A. leucostictus, A. manengubensis, A. osorioi, A. paradorsalis, and A. schneideri) or with longitudinal lateral dark bands (A. equatorialis and A. nigeriensis); species with a uniformly colored dorsum without conspicuous pattern except a dark dorsolateral stripe (A. aureus, A. clarkei, A. delicatus, A. fornasini, A. lacteus, A. leucostictus [dorsum with small white tubercles], A. uluguruensis, and A. weidholzi [might have a black vertebral line and dark brown flanks]); and from all remaining species having either dark stripes running from tip of snout to back, crossing each other and continuing as dorsolateral stripes (A. vibekensis, also males without asperities); anterior part of dorsum yellowish with brown pattern, posterior part of dorsum translucent (A. laevis, also males without asperities); posterior part of dorsum and dorsal sides of limbs semi-translucent with dense small dark white-edged speckles (A. dorsimaculatus); and dark blotches and transverse marks with white speckles (A. sylvaticus). The advertisement calls of the following species of Afrixalus consist of a long series of clicks, often initiated by a long buzzing note, and thus differ from the call of A. phantasma: A. aureus, A. brachycnemis, A. clarkei, A. crotalus, A. delicatus, A. dorsalis, A. fornasini, A. fulvovittatus, A. knysnae, A. nigeriensis, A. morerei, A. osorioi, A. quadrivittatus, A. septentrionalis, A. spinifrons, A. vibekensis, A. vittiger, A. weidholzi, and A. wittei. The advertisement calls of the remaining species for which information is available differ from the call of A. phantasma in the following characteristics (in parentheses): A. dorsimaculatus (soft, short buzzing rattle, usually repeated three times), A. equatorialis (series of five notes, repeated at 15–20/s and initiated by a long buzz, energy maximum at 2000–2500 Hz), A. lacteus (8–9 notes, 14–15 pulses,>300 pulses/s), A. paradorsalis (2–3 notes, energy maximum at 2700 Hz), A. stuhlmanni (2–9 notes, energy maximum at 4200–5100 Hz), and A. sylvaticus (2–5 notes, energy maximum at 4000–4500 Hz). The new species is most similar to A. lacustris sp. nov. from which it differs by a number of characteristics (see below). Description of holotype. Measurements of the holotype are provided in Table 2. Body very slender, widest at temporal region, slightly tapering to groin (Fig. 8A–C); head small (HL/SVL 0.31, HW/SVL 0.30), about as long as wide (HW/HL 0.98); snout relatively long, rounded in dorsal view and in lateral profile, slightly longer than wide; canthus rostralis hardly distinct, straight between eye and nostril; loreal region oblique; nostrils rounded, directed anterodorsally and slightly laterally; situated much closer to tip of snout and to eye, separated from each other by distance larger than distance between eye and nostril (IN/EN 1.15); eye directed anterolaterally, strongly protruding, very large (ED/HL 0.41), its diameter shorter than snout (ED/SL 0.84); interorbital distance much wider than upper eyelid and wider than internarial distance (IO/IN 1.21); tympanum covered by skin, not visible externally; upper jaw with dentition; teeth on premaxilla larger than teeth on maxilla; choanae small, located far anterolaterally at margins of roof of mouth, its anterior edge covered by maxillary bone, therefore appearing semicircular in ventral view; vomer ridges and teeth absent; tongue short and moderately broad, bilobed for about one-sixth of its length, free distally for about half its length; densely covered with minute papillae; median lingual process absent. Dorsal surfaces of head, trunk, and limbs and lateral surfaces of trunk finely shagreened; minute pointed, widely scattered tubercles on dorsum, more numerous posteriorly than anteriorly; supratympanic fold absent; small, subcircular, elevated glandular area in tympanic region posterior to eye and rear end of mandible, bearing large tubercles with pointed keratinous tips; ventral side of head smooth; vocal sac present; gular gland of vocal sac smooth, large and wide, covering about 70 percent of throat width (Fig. 8B); chest smooth, abdomen weakly areolate; ventral side of limbs smooth; short transverse fold above vent. Forelimbs slender;hand large (HND/SVL 0.32); tips of fingers enlarged into large disks, each with circummarginal groove; relative length of fingers: I I 2+/2.25 II 2/3+ III 2.75/2.25 IV; thenar tubercle oval, small and low, about one-fourth length of metacarpal of Finger I; palmar tubercles indiscernible. Hind limbs slender, moderately long; heel reaching to level of eye when legs adpressed forwardly to body; crus moderately long (TibL/SVL 0.48), about as long as thigh; heels meeting each other when knees flexed and thighs held perpendicularly to median plane; foot subequal in length to crus (FoL/TibL 0.96); relative length of toes: I I 2/2.25 II 1.5/3- III 1.75/3 IV 3-/1.75 V (Fig. 8C); inner metatarsal tubercle moderately prominent, elongated, about twothirds length of metatarsus of Toe I; outer metatarsal tubercle small, rounded. Coloration in life. During night, skin on dorsal side of head, trunk, forelimbs, crus and tarsus light yellowishbrown with more or less regular dark brown speckling; indistinct darker brown transverse stripe between upper eyelids; large dark brown, irregularly shaped spot in scapula region, two less distinct, somewhat lighter brown, irregularly shaped spots at level of anterior end of pelvis, separated from each other by half their width; cloacal region with dark brown mottling; dorsal skin of thigh largely unpigmented with only a broad stripe of widely scattered brown pigmentation; weak brown stripe along canthus, continuing behind eye on anterior half of trunk on both sides; tubercles below eye, and in tympanic region at rear end of jaw white; indistinct dark brown stripe running diagonally on middle of tibia, forming interrupted, inverted U-shaped band together with pelvic spots when legs folded against body; ventral side of head, trunk and limbs largely unpigmented and translucent; gular gland bright yellow; fingers and toes yellow (Fig. 7A). During day, basic dorsal coloration brighter, very light brown to bright cream-colored; dark brown dorsal speckles, spots and stripes in more pronounced contrast to basic coloration, more distinctly visible (Fig. 7B). Coloration in preservative. Dorsal basic coloration largely faded to yellowish white; darker dorsal pattern elements light to dark brown, clearly visible; yellow color of gular gland, fingers and toes faded to white (Fig. 8A–C). Variation. The paratypes match the holotype in general appearance, proportions, coloration and color pattern. Mensural variation within the species is shown in Table 1. Pedal webbing variation is I 2+[100]/2.5[90],2.25[10] II 2[50],2+[40],2.25[10]/3[10],3+[40],3.5[50] III 1.75 [80],2-[20]/3[100] IV 3 -[40],3[60]/1.75[100] V. Bioacoustics. Series of advertisement calls of six different males were recorded at the following ambient temperatures: 10.9°C (N = 1), 13.6°C (N = 3), and 16.2°C (N = 2). The advertisement call consisted of five to six, rarely four pulse groups (notes) (Fig. 9). Depending on the ambient temperature, notes were repeated at a rate of 7.1–10.0/s (10.9°C), 8.2–11.7/s (13.6°C), and 10.2–11.4/s (16.2°C). The highest repetition rate was always between the first three notes of a series, the lowest at the end of the series. Each note consisted of 10–11 pulses (Fig. 9). Probably due to echo effects, pulsation was often veiled in the waveforms, especially towards the end of the note. Pulse repetition rate varied from 190–210/s at 10.9°C, 222–250/s at 13.6°C, and 277–292/s at 16.2°C, resulting in a note length of 72–94 ms, 55–70 ms, and 36–50 ms, respectively. Amplitude modulation was prominent within individual notes (Fig. 9). Total call length showed linear temperature dependence and varied from 388–397 ms at 16.2°C, 472–499 ms at 13.6°C, and 572–620 ms at 10.9°C for five-note calls (Fig. 10). Energy maximum showed linear temperature dependence, with 3020–3150 Hz at 10.9°C, 3370–3550 Hz at 13.6°C, and 3660–3810 Hz at 16.2°C (Figs. 9–10). There was no marked frequency modulation. Prominent harmonics were at about 6000–7000 Hz and 9000–11000 Hz (Fig. 9). Ecology and natural history. We collected males and females in swamps in forest openings, or at forest edges (Fig. 11A). Males were found calling above standing bodies of water with thick lower vegetation cover. In Gishwati Forest and Kamiranzovu Swamp in Nyungwe Forest, the species was found calling in syntopy with Hyperolius castaneus and H. discodactylus. One adult male (UTEP 20802) from Kahuzi-Biega National Park (DRC) was found 3.5 meters above ground in montane forest. Tadpoles are unknown. Laurent (1955, 1983) listed the species from bushes in montane forest. Laurent (1982) noted that, in general, the species is common in the foliage of shrubs in dense forests. He also recorded it in banana trees in a deep valley (Nyungwe, Rwanda) and “des trous de prospection” (i.e., prospecting holes likely for gold mining, Upper Lubitshako, Kabobo Plateau, DRC). Etymology. The species epithet derives from the Greek noun φάντασμα (phántasma), meaning ghost or phantom, in allusion to the coloration and general appearance of the new species. The epithet is used as an invariable noun in apposition. Distribution and conservation. The occurrence of the species has been confirmed for several locations in western Rwanda (Nyungwe and Gishwati Forest) and eastern DRC (in and near Kahuzi-Biega National Park, Itombwe Plateau and Kabobo Plateau [Laurent 1982], Fig. 5). We expect the species to be eventually found in the AR of southwestern Uganda and northwestern Burundi. So far, the species has been recorded from only a narrow elevation range between 1962 m (Kamiranzovu Swamp, Rwanda) and ca. 2400 m (May ya Moto [and possibly Luemba], Itombwe Plateau, Laurent 1982). Given the relatively limited overall geographic distribution of the species with an estimated extent of occurrence of 19,088 km 2 (Fig. 5), an estimated area of occupancy of 52 km 2 (both calculated with GeoCAT; Bachman et al. 2011), the detection of the amphibian chytrid fungus in one adult male (UTEP 20791, Greenbaum et al. 2015), and the conservation challenges facing natural areas of the AR (Greenbaum 2017; Ayebare et al. 2018), we categorize this species as Vulnerable under the IUCN Red List criteria (IUCN 2021).

Toxicodryas blandingii (Hallowell, ���1844��� 1845) (Table 1, Figs. 5���8) Dipsas Blandingii: Hallowell (���1844��� 1845:170); type locality: ��� Liberia, West Africa. ��� Triglyphodon fuscum: Dum��ril, Bibron & Dum��ril (1854:1101); type locality: ��� Grand-Bassam, sur la C��te d���Ivoire (Guin��e)��� [Ivory Coast]. Dipsas fasciata: Fischer (1856:84); type locality: ��� Peki (West-Afrika)��� [Ghana]. Dipsas valida: Fischer (1856:87); type locality: ��� Edina (Grand Bassa County, West-Afrika)��� [Liberia]. Dipsas globiceps: Fischer (1856:89); type locality: ��� Edina (Grand Bassa County, Liberia, West-Afrika).��� Toxicodryas Blandingii: Hallowell (1857:60); comb. nov. Dipsas Fischeri: Jan in Dum��ril (1859:212); no type locality provided. Triglyphodon fuscum var. obscurum: Dum��ril (1861:211); type locality: ��� C��te d���Or ��� [Ghana]. Dipsas regalis: Jan (1871:3, Livraison 38, pl. vi, fig. 2) in Jan & Sordelli (1870 ���1881); type locality: ��� C��te d���Or ��� [Ghana]. Dipsas globiceps var. tumboensis: M��ller (1885:688); type locality: ��� Tumbo-Insel ��� [Guinea]. Boiga blandingi occidentalis: Stucki-Stirn (1979:377); inferred type locality: ���Besongabang��� Cameroon. Boiga blandingi subfulva: Stucki-Stirn (1979:381); type locality: none provided, but limited to Cameroon according to book title. Toxicodryas blandingii was originally described by Hallowell (1845:170) based on a single specimen collected by his friend, ���Dr. Blanding,��� in Liberia. The dorsum and venter of the specimen was noted to have a ���light yellow��� color with a series of blotches of ���leaden colour.��� This specimen reportedly possessed 2 preoculars, 2 postoculars, 272 ventrals, 131 subcaudals, body length (i.e., SVL) of 1.22 meters, and tail length of 0.39 meters. Hallowell (1854) provided additional details of the specimen���s teeth, noted it had 17 ���rows of scales,��� and corrected the tail length to 0.37 meters. Hughes & Barry (1969), Wallach et al. (2014) and Uetz et al. (2019) stated that the type was lost, which is consistent with Malnate (1971), who did not list a type specimen from the ANSP collection. Wallach et al. (2014) noted the type was a 1.67 m specimen, slightly longer than the total length of 1.61 m reported by Hallowell (1845) in the original description, but the longer measurement is likely a typographical error in reference to the latter citation (V. Wallach, pers. comm.). A query by EG to the Philadelphia Academy of Sciences in spring 2020 resulted in location of the type specimen (ANSP 10083, Fig. 6), and a redescription of this specimen is provided below. According to Loveridge (1957:269), the name Dipsas Fischeri was proposed by Jan (in Dum��ril 1859) to combine the minor color pattern variants Dipsas fasciata, D. valida, and D. globiceps named by Fischer (1856). Pel (1852:171) coined the name Dipsas regalis, and as translated by Savage & McDiarmid (2017:73), Pel stated, ���the third species of venomous snake, Naja atropos, belongs to cobras (spectacled snakes) and reaches a length of 6 to 7 feet [1.8���2.1 m]. Its color is entirely black... As this snake in general shows much similarity to a tree snake, Dipsas regalis, which equals it in color and size, but is not venomous.��� Perhaps because of this poor description, Boulenger (1896:78) attributed the latter name to Jan & Sordelli (1870 ���1881), who provided an illustration that served as an appropriate description. Jan listed the name in the Index des Planches for Livraison 38 as Dipsas cynodon Cuv. vari��t��? (D. regalis Schlegel), but according to Savage & McDiarmid (2017:73), the attribution to Hermann Schlegel is in error because he never used the name in any publication. M��ller (1885:687) seemed to suggest that Jan illustrated his specimen (collected by Dr. M��hly from ���Goldk��ste��� [i.e., Gold Coast or modern-day Ghana]) from Basel, but Hughes & Barry (1969:1020) listed a personal communication from M.S. Hoogmoed, who noted the type of D. regalis (specimen ���Leiden 958���) was collected by Pel in February 1844 from Accra, Ghana. Hallowell (1857:60) coined the genus Toxicodryas because he noticed that his specimen of T. blandingii had a ���single channelled posterior tooth on each side... and therefore...[it] cannot belong to the genus Triglophodon [sic] of Dum. and Bibron, which has three.��� Subsequent herpetological publications in the 19 th and early 20 th centuries seemingly ignored Hallowell���s new genus and continued to recognize the taxon in either the genus Dipsas (e.g., Mocquard 1896) or more commonly, Dipsadomorphus (e.g., Boulenger 1896, 1919; de Witte 1933). Schmidt (1923) transferred the taxon to the genus Boiga in his opus on Congolese snakes, recognizing B. (Toxicodryas) blandingii and B. (Toxicodryas) pulverulenta, an action that was followed by most subsequent authors for decades. Based on the placement of African Boiga in the ��� Dipsadidae: Lycodontinae ��� by Underwood (1967), Welch (1982) seems to have been the first to recognize the genus Toxicodryas for both species of the genus, an action followed by Meirte (1992) and observed by most herpetologists in the 21 st century (e.g., Uetz et al. 2020). Boulenger (1896) included all of the above West African, 19 th- century names in the synonymy of Dipsas (Toxicodryas) blandingii. Because the dubious subspecies described by Stucki-Stirn (1979) both occur in Cameroon, where one molecular sample (CAS 253611) from Allen et al. (in press) is recovered in a well-supported clade with West African samples (Fig. 2), we confirm the taxonomic nomenclature of Wallach et al. (2014) in treating these taxa as synonyms of T. blandingii. Marques et al. (2018) noted that northwestern Angolan records from ���Chinchoxo,��� ���Piri-Dembos,��� and ���Quirimbo��� by Peters (1877), Bocage (1895), Parker (1936), and Hellmich (1957a, b) are attributable to T. pulverulenta. However, the morphometric data provided by Hellmich (1957b) for Angolan snakes are inconsistent with the size and scale rows of the latter species, and herein, we consider his records to be attributable to T. vexator sp. nov. Diagnosis. Toxicodryas blandingii, as recognized herein, is restricted to West Africa and west-central Africa (west of the confluence of the Congo and Ubangi rivers), defined by the following combination of characters: maximum SVL> 1 meter (vs. maximum SVL T. pulverulenta and T. adamanteus sp. nov.); DSRN 23���25 (vs. 19���21 in T. pulverulenta and 18���23 in T. adamanteus sp. nov.); DSRM 21���25 (vs. 19���21 in T. pulverulenta and 18���21 in T. adamanteus sp. nov.); cloacal plate usually divided (vs. divided or undivided in T. vexator sp. nov., and always undivided in T. pulverulenta and T. adamanteus sp. nov.); adult males glossy or velvety black with a yellow venter, and adult females light brown, gray, or yellowish-brown with light-brown or cream cross-bars on the flanks, with yellowish-brown venters (vs. both sexes brown to pink with darker cross-bars that often enclose a whitish spot, and the dorsum and venter sprinkled with fine dark brown or black spots in T. pulverulenta and T. adamanteus sp. nov.); hemipenis relatively short and massive (i.e., broad), proximal third covered with spines, distal two-thirds dimpled with a flattened apex (vs. relatively long with long spines mid-way along the shaft that decrease in size towards the apex and base, and with a domed apex in T. pulverulenta and T. adamanteus sp. nov.); venom toxicity LD 50 = 2.85���3.55 mg /kg in mice (vs. venom toxicity LD 50 = 4.88 mg /kg in mice for T. vexator sp. nov.). Redescription of the holotype. ANSP 10083 (Fig. 6) adult female in poor condition, 1330 mm SVL; head triangular and distinct from neck, 1.92% of SVL (25.5 mm); right loreal missing, left loreal partially obscured by supralabials due to cranial damage, upper side tapering superiorly; body triangular; tail moderately long (400 mm; 30.1% of SVL). Supralabials ���/9, ���/4 th, 5 th, and 6 th contacting orbit; infralabials 14/13, 1 st on each side in contact behind mental, 1 st ���4 th /1 st ���4 th contacting anterior chin shields and 4 th ���7 th /4 th ���7 th contacting posterior chin shields; 2 preoculars; 3 postoculars (on left, missing on right); temporals ���/2 + 2; 2 internasals; nasal divided (on left, missing on right); frontal slightly longer than wide, only left side undamaged; dorsal scale rows 23 one head length posterior to jaw rictus, 23 at midbody, 17 one head length anterior to vent, smooth and oblique, vertebral scales broad and apically flattened; ventrals 273 (standard), 271 (Dowling); cloacal plate undivided; paired subcaudals 132. Coloration (in preservative). After approximately 176 years in preservation, specimen is faded, with creamy tan background color in dorsal and ventral views. Brown markings on posterior edge of supralabials and dorsum of head. Irregular brown and dark brown blotches and saddles on dorsum from neck to tip of tail (Fig. 6). Variation. Morphometric variation of Toxicodryas blandingii is shown in Table 1. M��ller (1885:688) provided data for a snake from Ghana with 15 infralabials and noted that most of its scales have two ���Endgruben��� [terminal pits], which likely refer to apical pits. In his description of Dipsas globiceps var. tumboensis M��ller (1885) noted his specimen from Guinea (Fig. 7) had 147 subcaudals. In snakes from West Africa (without separating by sex), Angel (1933) noted a temporal formula of 2 + 2 or 2 + 3, 21���25 scales at midbody, 240���289 ventrals, 120���147 subcaudals, either an undivided or divided cloacal plate, and a maximum total size of 2290 mm, and nearly verbatim variation was listed by Villiers (1950a), Doucet (1963), Stucki-Stirn (1979), Chippaux (2006), and Trape & Man�� (2006). However, in snakes from Ghana, Swiecicki (1965:302) noted a maximum total length of 2450 mm for a ���black form��� individual, Gauduin (1970) listed the maximum total length as 2700 mm (600 mm tail) for Cameroon, Chirio & LeBreton (2007) provided a slightly larger total length of 2740 mm for Cameroon, and Luiselli et al. (1998a) noted a maximum size of 2800 mm, presumably for Nigeria. Villiers (1951) noted an unsexed individual from Benin with 115 subcaudals. Cansdale (1965) documented 21���25 scales at midbody, and Segniagbeto et al. (2011) documented snakes from Togo with 19���24 scales at midbody and 102���159 subcaudals. In our examined specimens, temporal formula includes the variation noted by Angel (1933), but is more extensive (2 +5, 3 + 4, 3 + 3, 1 + 5, 3 + 2, or 2 + 4), and either supralabials 3���5 or 4���6 contact the eye, which is consistent with the observations of Angel (1933), Villiers (1950b), and Chippaux (2006); the latter author also noted that sometimes only 2 supralabials contact the eye. Rasmussen (1997a) noted specimens with the 4th���5th, 5th���7 th, or 4th���7 th supralabials in contact with the eye, and in general, this species has sloping and smooth scales with apical pits, and the vertebral row is greatly enlarged. The holotype, one male and one female from Liberia (Loveridge 1941; Johnsen 1962), one male from Gabon (Pauwels et al. 2002b), one male from DRC (RBINS 10888), and a juvenile from Cameroon (Werner 1897) are unusual in having an undivided cloacal plate, because all other examined specimens have a divided cloacal plate, including the type specimens of Dipsas fasciata, D. valida, and D. globiceps (Fischer 1856). Rasmussen (1997a) remarked that his specimens have either a divided or undivided cloacal plate. In his book on West and Central African snakes, including countries west of the Congo River, Chippaux (2006:154) noted the anal [cloacal plate] is sometimes entire, but more often divided. Trape & Man�� (2006) stated that the cloacal plate is almost always divided. Segniagbeto et al. (2011) noted individuals with divided or undivided cloacal plates in Togo. Combined descriptions by Fischer (1856) of the teeth of Dipsas fasciata, D. valida, and D. globiceps (all now synonyms of T. blandingii) suggest the species has 9 maxillary teeth that increase in size posteriorly, followed by two fangs (three on the right side in one specimen), and 12 mandibular teeth, which decrease in size posteriorly. In snakes from Ghana, Leeson (1950) noted 10���11 maxillary teeth, becoming larger posteriorly, and two fangs followed by a shorter fang; fourteen large palatine and pterygoid teeth, and 15 mandibular teeth (anterior ones largest). Taylor & Weyer (1958:1217) described a Liberian specimen with 9���10 maxillary teeth (on different sides) that increase in size from the 1 st to 4 th tooth, and then become subequal; two fangs occur after this series of teeth, and after a short diastema, there is a third fang with only traces of a groove. A second Liberian specimen had 10 maxillary teeth followed by three fangs, the last of which had only ���a suggestion of a groove.��� Based on specimens ranging from Guinea to Congo, Rasmussen (1997a:98) noted 10 maxillary teeth followed by three enlarged, furrowed venomous teeth, with the 3 rd fang slightly smaller than the previous two. Fischer (1856) provided detailed descriptions of the color patterns of West African Dipsas fasciata, D. valida, and D. globiceps (all now synonyms of T. blandingii), which seem to suggest he examined a subadult male that had been kept in alcohol for a long (unspecified amount) time (D. fasciata), an adult female (D. valida), and a subadult that retained juvenile coloration (D. globiceps). In his description of Dipsas globiceps var. tumboensis M��ller (1885:689) noted his specimen from Guinea (Fig. 7) had a gray-reddish dorsum with 30 black transverse bands, usually containing milky white spots. The frontal, supraoculars, and occipital scales had large black spots, the labial scales and postoculars were edged with black, and the head shields had multiple milk-white speckles. The tail was bright red with dark, irregular transverse bands. Aspects of this coloration description are highly unusual for this species (e.g., bright red tail), and more typical of T. pulverulenta, but the black edging of the labial scales, number of preoculars (3), supralabials in contact with the eye (5 and 6), ventrals (269), and subcaudals (147) clearly indicate this taxon is a synonym of T. blandingii (Fig. 7, Tables 1���2). Mocquard (1887:80) described a recently collected, unsexed subadult (���la longueur du tronc��� [trunk length] 1.1 m) from Gabon as having a dorsal color of a general tint of Burgundy with slightly darker spots on the flanks that have a dirty white spot a little above their lower edge. Mertens (1938) described an adult male from Cameroon as solid black dorsally and ventrally, with the exception of the anterior third of the venter, which was white, but the ventral scales had dark gray edges. The labial scales were gray with vertical black borders. Villiers (1950b) described the color pattern of an unsexed individual from Ivory Coast as sooty black or brownish in places on the dorsum; underside iridescent dark gray posteriorly, becoming whitish anteriorly, with the posterior edge of the ventrals edged with gray; underside of head white, and labials whitish and edged in black. Another unsexed individual from Liberia was described as bluish black above, yellow below; supralabials yellow with black edges, and posterior part of venter and underside of tail black. A third unsexed individual from Liberia had identical coloration to the latter one, except for the presence of whitish bars on the neck. Leeson (1950) noted that snakes from Ghana have a dull green or gray dorsum. Monard (1951:162) described an unsexed individual from Cameroon as a beautiful light redbrown, ���barr����� [barred] with dark brown. Taylor & Weyer (1958:1217) described a Liberian brown-phase female with pale grayish green on the ventral side of the head and neck, merging into gray with ���a greenish cast��� 12.7 cm posteriorly, and at 40.6 cm behind the head, it transitioned into plain tan to the tip of the tail. Isemonger (1962:12) remarked that this species has a ���delicate bloom on the skin.��� Cansdale (1965:43) described a highly unusual color pattern for juveniles by noting that ���the young brown form is pink with irregular chocolate markings that break up its outline very effectively and make it difficult to pick out in a tree or shrub.��� Leston & Hughes (1968:753) described an unusual specimen from Ghana as ���pale grey with darker greyish-green transverse bands, the bands irregular but more or less diamond shaped on each side. The ventrals are also grey but more glossy.��� Groves (1973:107) described the coloration of hatchlings from a captive Liberian female as ���light grey background colour with pinkish undertones; black, roughly oval, lateral blotches narrowing as they approach the midline, where many of them fail to conjoin; top of head light grey; belly dark grey.��� Rasmussen (1997a:98) noted the scales of his specimens were dull and almost dusty, a sentiment also shared by Cansdale (1965). Adult males were solid black on the dorsum and yellow on the venter (becoming black posteriorly), whereas adult females were noted to be gray, brown or yellow-brown on the dorsum and yellow-brown on the venter, sometimes without transverse bands. Hughes (2000:8) noted juvenile and subadult (approximately 1 meter in total length or less) snakes had a dorsal coloration that was ���a distinctively bright and contrasting pattern of chocolate brown blotches.��� He noted that most male specimens lose this coloration as they age, becoming increasingly melanistic, and although exceptions are possible, this melanistic progression does not seem to occur in females. Chippaux (2006:154) noted there are two dorsal color morphs: (1) uniform black or dark blue with ���reflets velout��s��� [velvety reflections] or (2) gray with darker, poorly defined transverse spots. The venter was noted as dull yellow to charcoal gray, and juvenile coloration as light brown with darker transverse ring-shaped spots. Stucki-Stirn (1979), perhaps confused by the tw, Published as part of Greenbaum, Eli, Allen, Kaitlin E., Vaughan, Eugene R., Pauwels, Olivier S. G., Wallach, Van, Kusamba, Chifundera, Muninga, Wandege M., Aris- Tote, Mwenebatu M., Mali, Franck M. M., Badjedjea, Gabriel, Penner, Johannes, R��del, Mark-Oliver, Rivera, Jacqueline, Sterkhova, Viktoria, Johnson, Grant, Tapondjou, Walter P. & Brown, Rafe M., 2021, Night stalkers from above: A monograph of Toxicodryas tree snakes (Squamata Colubridae) with descriptions of two new cryptic species from Central Africa, pp. 1-44 in Zootaxa 4965 (1) on pages 7-17, DOI: 10.11646/zootaxa.4965.1.1, http://zenodo.org/record/4723024, {"references": ["Fischer, J. G. (1856) Neue Schlangen des Hamburgischen Naturhistorischen Museums. Abhandlungen aus dem Gebiete der Naturwissenschaften herausgegeben von Naturwissenschaftlichen Verein in Hamburg, 3, 79 - 116 + 3 pl.", "Hallowell, E. (1857) Notice of a collection of reptiles from the Gaboon country, West Africa, recently presented to the Academy of Natural Sciences of Philadelphia, by Dr. Henry A. Ford. Proceedings of the Academy of Natural Sciences of Philadelphia, 9 (1857 - 1858), 48 - 72.", "Dumeril, A. H. A. (1859) Reptiles et poissons de l'Afrique occidentale. Etude precedee de considerations generales sur leur distribution geographique. Archives du Museum National d'Histoire Naturelle, 10, 137 - 240.", "Jan, G. & Sordelli, F. (1870 - 1881) Iconographie Generale des Ophidiens. Tome Troisieme (Livrais. 35 a 50), Contenant Quatre Vingt Seize Planches. Georges Jan and Ferdinand Sordelli, Milan, 96 pls.", "Muller, F. (1885) Vierter nachtrag zum Katalog der herpetologischen Sammlung der Basler Museums. Verhandlungen der Naturforschenden Gesellschaft in Basel (1887 - 1890), 7, 668 - 717. https: // doi. org / 10.5962 / bhl. part. 2388", "Stucki-Stirn, M. C. (1979) A Comparative Study of the Herpetological Fauna of the former West Comeroon [sic] / Africa: With a Classification and Synopsis of 95 Different Snakes and Description of Some New Sub-species. Snake Report 721. Herpeto- Verlag, Teuffenthal, Switzerland, 650 pp.", "Hallowell, E. (1854) Remarks on the geographical distribution of reptiles, with descriptions of several species supposed to be new, and corrections of former papers. Proceedings of the Academy of Natural Sciences of Philadelphia, 7 (1854 - 1855), 98 - 105.", "Hughes, B. & Barry, D. H. (1969) The snakes of Ghana: a checklist and key. Bulletin de l'Institut fondamental d'Afrique noire. Serie A, Sciences naturelles, 31 (3), 1004 - 1041.", "Wallach, V., Williams, K. L. & Boundy, J. (2014) Snakes of the World: A Catalogue of Living and Extinct Species. CRC Press, Boca Raton, Florida, xxvii + 1209 pp.", "Uetz, P., Cherikh, S., Shea, G., Ineich, I., Campbell, P. D., Doronin, I. V., Rosado, J., Wynn, A., Tighe, K. A., McDiarmid, R., Lee, J. L., Kohler, G., Ellis, R., Doughty, P., Raxworthy, C. J., Scheinberg, L., Resetar, A., Sabaj, M., Schneider, G., Franzen, M., Glaw, F., Bohme, W., Schweiger, S., Gemel, R., Couper, P., Amey, A., Dondorp, E., Ofer, G., Meiri, S. & Wallach, V. (2019) A global catalog of primary reptile type specimens. Zootaxa, 4695 (5), 438 - 450. https: // doi. org / 10.11646 / zootaxa. 4695.5.2", "Malnate, E. V. (1971) A catalog of primary types in the herpetological collections of the Academy of Natural Sciences, Philadelphia (ANSP). Proceedings of the Academy of Natural Sciences of Philadelphia, 123, 345 - 375.", "Loveridge, A. (1957) Check list of the reptiles and amphibians of East Africa (Uganda; Kenya, Tanganyika; Zanzibar). Bulletin of the Museum of Comparative Zoology, Harvard, 117 (2), 151 - 362 + i - xxxvi (index).", "Pel, H. S. (1852) Over de jagt aan de Goudkust, volgens eene tienjarige eigene ondervinding. Nederlandsch Tijdschrift voor Liefhebbers der Jagt und Visscherei, 1, 149 - 173.", "Savage, J. M. & McDiarmid, R. W. (2017) The Herpetological Contributions of Giorgio Jan (1791 - 1866): With an Introduction, Annotated Bibliography, Synopsis of Herpetological Taxa, and a Comprehensive Guide to the Iconographie generale des Ophidiens. Society for the Study of Amphibians and Reptiles, Villanova, Pennsylvania, viii + 926 pp.", "Boulenger, G. A. (1896) Catalogue of the Snakes in the British Museum (Natural History). Vol. III. Containing the Colubridae (Opisthoglyphae and Proteroglyphae), Amblycephalidae, and Viperidae. Trustees of the British Museum (Natural History), London, xiv + 727 pp.", "Mocquard, F. (1896) Notes sur quelques ophidiens de Conakry. Bulletin du Museum Paris, 20, 59 - 60.", "Boulenger, G. A. (1919) Batraciens et reptiles recueillis par le Dr C. Christy au Congo Belge dans le districts de Stanleyville, Haut-Uele et Ituri en 1912 - 1914. Revue Zoologique Africaine, 7, 1 - 29. https: // doi. org / 10.5962 / bhl. part. 15108", "de Witte, G. - F. (1933) Reptiles recoltes au Congo Belge par le Dr H. Schouteden et par M. G. - F. de Witte. Annales du Musee du Congo Belge, C. - Zoologie, 3 (1), 55 - 98.", "Schmidt, K. P. (1923) Contributions to the herpetology of the Belgian Congo based on the collection of the American Museum Congo Expedition, 1909 - 1915. Part II. - Snakes. Bulletin of the American Museum of Natural History, 49, 1 - 146, pls. I - XXII.", "Underwood, G. (1967) A Contribution to the Classification of Snakes. British Museum (Natural History), London, x + 179 pp.", "Welch, K. R. G. (1982) Herpetology of Africa: A Checklist and Bibliography of the Orders Amphisbaenia, Sauria and Serpentes. Robert E. Krieger Publishing Company, Malabar, Florida, 293 pp.", "Meirte, D. (1992) Cles de determination des serpents d'Afrique. Museum Royal d'Afrique Centrale, Tervuren Belgique Annual Series Octavo Science Zoologique, 267, 1 - 152.", "Uetz, P., Freed, P. & Hosek, J. (Eds.) (2020) The Reptile Database. Available from: http: // www. reptile-database. org (accessed 12 December 2020)", "Marques, M. P., Ceriaco, L. M. P., Blackburn, D. C. & Bauer, A. M. (2018) Diversity and distribution of the amphibians and terrestrial reptiles of Angola: Atlas of historical and bibliographic records (1840 - 2017). Proceedings of the California Academy of Sciences, Series 4, 65 (Supplement II), 1 - 501.", "Peters, W. C. H. (1877) Ubersicht der Amphibien aus Chinchoxo (Westafrika), welche von der Afrikanischen Gesellschaft dem Berliner zoologischen Museum ubergeben sind. Monatsberichte der Koniglichen Preussischen Akademie der Wissenschaften zu Berlin, 1877, 611 - 621.", "Bocage, J. V. B. (1895) Herpetologie d'Angola et du Congo. Ministerio da Marinha e das Colonias, Lisbonne, XX + 203 pp., pls. I - XIX.", "Parker, H. W. (1936) Dr. Karl Jordan's expedition to South-West Africa and Angola: Herpetological collection. Novitates Zoologicae, 40, 115 - 146.", "Hellmich, W. (1957 a) Herpetologische Ergebnisse einer Forschungreise in Angola. Veroffentlichungen der Zoologischen Staatssammlung Munchen, 5, 1 - 92.", "Hellmich, W. (1957 b) Die Reptilienausbeute der hamburgischen Angola-Expedition. Mitteilungen aus dem Hamburgischen Zoologischen Museum und Institut, 55, 39 - 80.", "Angel, F. (1933) Les Serpents d'Afrique Occidentale Francaise. Larose Editeur, Paris, 246 pp.", "Villiers, A. (1950 a) Initiations Africaines. II. Les Serpents de l'Ouest Africain. Institut Francais d'Afrique Noire, Dakar, Senegal, 148 pp.", "Doucet, J. (1963) Les serpents de la Republique de Cote d'Ivoire. Serpents venimeux. Acta Tropica, 20 (3 - 4), 297 - 340.", "Chippaux, J-P. (2006) Les Serpents d'Afrique Occidentale et Centrale. 3 rd Edition. IRD Editions, Paris, 311 pp.", "Trape, J. - F. & Mane, Y. (2006) Guide des Serpents d'Afrique Occidentale: Savane et Desert. IRD editions, Paris, 226 pp. https: // doi. org / 10.4000 / books. irdeditions. 37382", "Swiecicki, A. W. (1965) Snakes and snake bite in the western region, Ghana. Journal of Tropical Medicine and Hygiene, 68 (12), 300 - 304.", "Gauduin, P. (1970) Contributions a la faune de la region de Yaounde. Annales de la Faculte des Sciences du Cameroun, 4, 73 - 99.", "Chirio, L. & LeBreton, M. (2007) Atlas des Reptiles du Cameroun. Publications Scientifiques du MNHN, IRD Editions, Paris, 686 pp.", "Luiselli, L., Akani, G. C. & Barieenee, I. F. (1998 a) Observations on habitat, reproduction and feeding of Boiga blandingi (Colubridae) in south-eastern Nigeria. Amphibia-Reptilia, 19 (4), 430 - 436. https: // doi. org / 10.1163 / 156853898 X 00098", "Villiers, A. (1951) Mission A. Villiers au Togo et au Dahomey (1950). II. - Ophidiens. Etudes Dahomeennes, 5, 17 - 43.", "Cansdale, G. S. (1965) West African Snakes. Longmans, Green and Co. Ltd., London, vi + 74 pp.", "Segniagbeto, G. H., Trape, J. F., David, P., Ohler, A., Dubois, A. & Glitho, I. A. (2011) The snake fauna of Togo: Systematics, distribution and biogeography, with remarks on selected taxonomic problems. Zoosystema, 33 (3), 325 - 360. https: // doi. org / 10.5252 / z 2011 n 3 a 4", "Villiers, A. (1950 b) Catalogues. IV. La Collection de Serpents de l'I. F. A. N. Institut Francais d'Afrique Noire, Dakar, Senegal, 155 pp.", "Rasmussen, J. B. (1997 a) Afrikanske slanger (10). Boiga blandingii. Nordisk Herpetologisk Forening, 40 (3), 97 - 103.", "Loveridge, A. (1941) Report on the Smithsonian-Firestone expedition's collection of reptiles and amphibians from Liberia. Proceedings of the United States National Museum, 91 (3128), 113 - 140. https: // doi. org / 10.5479 / si. 00963801.91 - 3128.113", "Johnsen, P. (1962) Notes on African snakes, mainly from northern Rhodesia and Liberia. Videnskabelige Meddelelser fra Dansk Naturhistorisk Forening i Kjobenhavn, 124, 115 - 130.", "Pauwels, O. S. G., Kamdem Toham, A. & Chimsunchart, C. (2002 b) Recherches sur l'herpetofaune des Monts de Cristal, Gabon. Bulletin de l'Institut Royal des Sciences Naturelles de Belgique, Biologie, 72, 59 - 66.", "Werner, F. (1897) Ueber Reptilien und Batrachier aus Togoland, Kamerun und Tunis aus dem kgl. Museum fur Naturkunde in Berlin. Verhandlungen der Kaiserlich-Koniglichen Zoologisch-Botanischen Gesellschaft in Wien, 47, 395 - 408. https: // doi. org / 10.5962 / bhl. part. 11464", "Leeson, F. (1950) Identification of Snakes of the Gold Coast. The Crown Agents for the Colonies, London, x + 142 pp.", "Taylor, E. H. & Weyer, D. (1958) Report on a collection of amphibians and reptiles from Harbel, Republic of Liberia. The University of Kansas Science Bulletin, 38 (14), 1191 - 1229. https: // doi. org / 10.5962 / bhl. part. 10973", "Mocquard, F. (1887) Sur les ophidiens repportes du Congo par la Mission de Brazza. Bulletin de la Societe Philomathique de Paris, Serie 7, 11, 62 - 92.", "Mertens, R. (1938) Herpetologische Ergebnisse einer Reise nach Kamerun. Abhandlungen der Senckenbergischen Naturforschenden Gesellschaft, 442, 1 - 52.", "Monard, A. (1951) Resultats de la mission zoologique suisse au Cameroun. Reptiles. Batraciens. Memoires de l'Institut Francais d'Afrique Noire, Sciences Naturelles, (1), 123 - 170.", "Isemonger, R. M. (1962) Snakes of Africa: Southern, Central and East. Thomas Nelson and Sons (Africa) (Party) Ltd., Johannesburg, viii + 236 pp.", "Leston, D. & Hughes, B. (1968) The snakes of Tafo, a forest cocoa-farm locality in Ghana. Bulletin de l'Institut Francaise d'Afrique Noire, 30 A, 737 - 770.", "Groves, F. (1973) Reproduction and venom in Blanding's tree snake. International Zoo Yearbook, 13, 106 - 108. https: // doi. org / 10.1111 / j. 1748 - 1090.1973. tb 02114. x", "Hughes, B. (2000) Polymorphism, sex, size and other aspects of the African snake, Boiga blandingi (Hallowell, 1844). Herpetological Bulletin, 74, 8 - 12.", "Lawson, D. (1993) The reptiles and amphibians of the Korup National Park Project, Cameroon. Herpetological Natural History, 1 (2), 27 - 90.", "Greenbaum, E. & Carr, J. L. (2005) The herpetofauna of Upper Niger National Park, Guinea, West Africa. Scientific Papers, Natural History Museum, The University of Kansas, 37, 1 - 21. https: // doi. org / 10.5962 / bhl. title. 8470", "Pauwels, O. S. G., Pauly, A., Araldi, A., Ndonda Makemba, R., Fonteyn, D., Oslisly, R. & Whittaker, A. M. (2020) Miscellanea Herpetologica Gabonica XVI. Bulletin of the Chicago Herpetological Society, 55 (5), 93 - 99.", "Sternfeld, R. (1909) Die Fauna der deutschen Kolonien, Herausgegeben mit Unterstutzung des Reichs-Kolonialamtes vom Zoologischen Museum in Berlin. Reihe I. Kamerun. Heft I. Die Schlangen Kameruns. R. Friedlander & Sohn, Berlin, iv + 28 pp.", "Leache, A. D., Grummer, J. A., Miller, M., Krishnan, S., Fujita, M. K., Bohme, W., Schmitz, A., Lebreton, M., Ineich, I., Chirio, L., Ofori-Boateng, C., Eniang, E. A., Greenbaum, E., Rodel, M. - O. & Wagner, P. (2017) Bayesian inference of species diffusion in the West African Agama agama species group (Reptilia, Agamidae). Systematics and Biodiversity, 15 (3), 192 - 203. https: // doi. org / 10.1080 / 14772000.2016.1238018", "Cansdale, G. (1955) Reptiles of West Africa. Penguin Books, London, United Kingdom, 104 pp.", "Dekeyser, P. L. (1955) Les Mammiferes de l'Afrique Noire Francaise. 2 nd Edition. Institut Francais d'Afrique Noire, Initiations Africaines, Dakar, Senegal, 426 pp.", "Villiers, A. (1955) Note sur quelques Ixodidae et Gamasidae parasites des Vertebres rencontres en Afrique Occidentale francaise. Bulletin de l'Institut fondamental d'Afrique noire, Serie A, 17 (2), 444 - 454.", "Wickler, V. W. & Uhrig, D. (1969) Verhalten und okologische Nische der Gelbflugelfledermaus, Lavia frons (Geoffroy) (Chiroptera, Megadermatidae). Zeitschrift fur Tierpsychologie, 26 (6), 726 - 736. https: // doi. org / 10.1111 / j. 1439 - 0310.1969. tb 01972. x", "Happold, M. & Happold, D. (Eds.) (2013) Mammals of Africa. Vol. IV. Hedgehogs, Shrews and Bats. Bloomsbury, London, New Delhi, New York, and Sydney, 800 pp.", "Villiers, A. (1956) La collection de serpents de l'I. F. A. N. (Acquisitions 1954 - 1955). Bulletin de l'Institut fondamental d'Afrique noire, Serie A, 18 (3), 878 - 883.", "Wuster, W., Chirio, L., Trape, J-F., Ineich, I., Jackson, K., Greenbaum, E., Kusamba, C., Barron, C., Nagy, Z., Storey, R., Hall, C., Wuster, C., Barlow, A. & Broadley, D. G. (2018) Integration of nuclear and mitochondrial gene sequences and morphology reveal unexpected diversity in the forest cobra (Naja melanoleuca) species complex in Central and West Africa (Serpentes: Elapidae). Zootaxa, 4455 (1), 68 - 98. https: // doi. org / 10.11646 / zootaxa. 4455.1.3", "Pitman, C. R. S. (1958) Snake and lizard predators of birds. Bulletin of the British Ornithologists' Club, 78, 82 - 86.", "Woodward, S. F. (1960) A bat-eating snake. The Nigerian Field, 25, 172 - 174.", "Menzies, J. I. (1961) Correspondence: bat-eating snakes. The Nigerian Field, 26, 44.", "Cozens, B. (1961) Correspondence: bat-eating snakes. The Nigerian Field, 26, 44.", "Pitman, C. R. S. (1962) More snake and lizard predators of birds. Bulletin of the British Ornithologists' Club, 82, 45 - 55.", "Barry, D. H. (1961) Correspondence: bat-eating snakes. The Nigerian Field, 26, 42 - 43.", "Dunger, G. T. (1961) Correspondence: bat-eating snakes. The Nigerian Field, 26, 43.", "Greene, H. W. (1989) Ecological, evolutionary, and conservation implications of feeding biology in Old World cat snakes, Genus Boiga (Colubridae). Proceedings of the California Academy of Sciences, 46 (8), 193 - 207.", "Jones, T. (1961) Notes on bat-eating snakes. The Nigerian Field, 26, 69 - 70.", "Menzies, J. I. (1966) The snakes of Sierra Leone. Copeia, 1966 (2), 169 - 179. https: // doi. org / 10.2307 / 1441123", "Leston, D. (1970) Some snakes from the forest zone of Ghana. British Journal of Herpetology, 4 (6), 141 - 144.", "Greene, H. W. (1997) Snakes: The Evolution of Mystery in Nature. University of California Press, Berkeley, Los Angeles and London, xiii + 351 pp.", "Luiselli, L., Akani, G. C. & Capizzi, D. (1998 b) Food resource partitioning of a community of snakes in a swamp rainforest of south-eastern Nigeria. Journal of Zoology, London, 246 (2), 125 - 133. https: // doi. org / 10.1111 / j. 1469 - 7998.1998. tb 00141. x", "Bohme, W. (2000) Diversity of a snake community in a Guinean rain forest (Reptilia, Serpentes). In: Rheinwald, G. (Ed.), Isolated Vertebrate Communities in the Tropics. Proceedings of the 4 th International Symposium in Bonn. Bonner zoologische Monographien, 46, 69 - 78.", "Akani, G. C., Ebere, N., Luiselli, L. & Eniang, E. A. (2007) Community structure and ecology of snakes in fields of oil palm trees (Elaeis guineensis) in the Niger Delta, southern Nigeria. African Journal of Ecology, 46 (4), 500 - 506. https: // doi. org / 10.1111 / j. 1365 - 2028.2007.00885. x", "Pauwels, O. S. G., Carlino, P., Chirio, L., Daversa, D. R., Lips, J., Oslisly, R. & Testa, O. (2019 a) Amphibians and reptiles found in caves in Gabon, western Equatorial Africa. Cave and Karst Science, 46 (1), 3 - 12.", "Butler, J. A. & Reid, J. C. (1990) Records of snakes from Nigeria. The Nigerian Field, 55 (1), 19 - 40.", "Pauwels, O. S. G., Carlino, P., Chirio, L., Meunier, Q., Vivien Okouyi Okouyi, J., Orbell, C., Rousseaux, D. & Testa, O. (2017 c) Miscellanea Herpetologica Gabonica IX. Bulletin of the Chicago Herpetological Society, 52 (6), 97 - 102.", "Young, R. A. (1983) Feeding behavior of captive Boiga. Bulletin of the Chicago Herpetological Society, 18 (3 - 4), 89 - 91.", "Rodel, M. - O. & Glos, J. (2019) Herpetological surveys in two proposed protected areas in Liberia, West Africa. Zoosystematics and Evolution, 95 (1), 15 - 35. https: // doi. org / 10.3897 / zse. 95.31726", "Pauwels, O. S. G. & Vogel, G. (2011). Ein Fall von Thanatose (Totstellreflex) bei Boiga blandingii (Squamata, Serpentes, Colubridae) in Gabun. Elaphe, 4, 24 - 25.", "Pasqual, J. R. H. (1962) Correspondence: West African snakes. The Nigerian Field, 27, 189 - 190.", "Knoepffler, L. - P. (1966) Faune du Gabon (amphibiens et reptiles). I. Ophidiens de l'Ogooue-Ivindo et du Woleu N'tem. Biologia Gabonica, 2 (1), 3 - 23.", "Roux-Esteve, R. (1969) Les serpents de la region de Lamto (Cote-d'Ivoire). Annales de l'Universite d'Abidjan, Ecologie, 2 (1), 81 - 140.", "Perret, J. - L. (1961) Etudes herpetologiques africaines III. Bulletin de la Societe de Sciences Naturelles de Neuchatel, 84, 133 - 138.", "Hughes, B. (1983) African snake faunas. Bonner zoologische Beitrage, 34 (1 - 3), 311 - 356.", "Mertens, R. (1941) Zur Kenntnis der Herpetofauna von Fernando-Poo. 3. Beitrag zu den wissenschaftlichen Ergebnissen der Forschungreise H. Eidmann nach Spanisch-Guinea 1939 / 40. Zoologischer Anzeiger, 135, 275 - 281.", "Blackwell, K. (1967) A preliminary survey of the reptiles of the Ibadan area. British Journal of Herpetology, 3, 307 - 311.", "Butler, J. A. & Reid, J. C. (1986) Habitat preferences of snakes in the southern Cross River State, Nigeria. In: Rocek, Z. (Ed.), Studies in Herpetology. Charles University, Prague, pp. 483 - 488.", "Hughes, B. (1988) Herpetology in Ghana (West Africa). British Herpetological Society Bulletin, (25), 29 - 37.", "Rodel, M. - O., Kouadio, K. & Mahsberg, D. (1999) Die Schlangenfauna des Comoe-Nationalparks, Elfenbeinkuste: Erganzungen und Ausblick. Salamandra, 35 (3), 165 - 180.", "Leache, A. D. (2005) Results of a herpetological survey in Ghana and a new country record. Herpetological Review, 36 (1), 16 - 19.", "Auliya, M., Wagner, P. & Bohme, W. (2012) The herpetofauna of the Bijagos archipelago, Guinea-Bissau (West Africa) and a first country-wide checklist. Bonn zoological Bulletin, 61 (2), 255 - 281.", "Akaffou, M. H., Chippaux, J. - P., Kouadio Allali, B., Coulibaly, Z. & Dosso, M. (2017) Peuplement ophidien des plantations d' Hevea brasiliensis d'Anguededou (sud-est de la Cote d'Ivoire). Bulletin de la Societe Herpetologique de France, 162, 31 - 38.", "Pauwels, O. S. G., Albert, J. - L., Arrowood, H., Mvele, C., Casanova, M., Dodane, J. - B., Morgan, J., Primault, L., Thepenier, L. & Fenner, J. N. (2017 a) Miscellanea Herpetologica Gabonica X. Bulletin of the Chicago Herpetological Society, 52 (8), 133 - 138.", "Leache, A. D., Oaks, J. R., Ofori-Boateng, C. & Fujita, M. K. (2020) Comparative phylogeography of West African amphibians and reptiles. Evolution, 74 (4), 716 - 724. https: // doi. org / 10.1111 / evo. 13941", "Taub, A. M. (1967) Comparative histological studies on Duvernoy's gland of colubrid snakes. Bulletin of the American Museum of Natural History, 138 (1), 1 - 50.", "Levinson, S. R., Evans, M. H. & Groves, F. (1976) A neurotoxic component of the venom from Blanding's tree snake (Boiga blandingi). Toxicon, 14 (4), 307 - 312. https: // doi. org / 10.1016 / 0041 - 0101 (76) 90027 - 1", "Weinstein, S. A. & Kardong, K. V. (1994) Properties

Toxicodryas pulverulenta (Fischer, 1856) (Table 2, Figs. 8, 11) Dipsas pulverulenta: Fischer (1856:11); type locality: ��� Edina, Grand Bassa County in Liberia (West-Afrika).��� Dipsadomorphus Boueti: Chabanaud (1917a:373); type locality: none provided, but subsequently clarified to ��� Porto-Novo ��� ��� Dahomey ��� [Porto-Novo, Benin] by Chabanaud (1917b:12). As noted by Cer��aco et al. (2018), the description of this species was based on two specimens (only one syntype remains today according to Hallermann 1998 and Uetz et al. 2019), one of which was depicted in plate 4 of livraison 38 by Jan & Sordelli (1871). In a footnote to the nomenclatural history of Boiga (Toxicodryas) pulverulenta, Hughes & Barry (1969:1020) wrote ���Dr. Ladiges [then curator at ZMH] tells us that the type is still in Hamburg and is from St. Thom��, not Edina, Grand Bassa County, Liberia as stated by Loveridge (1958:269).��� The latter citation is in error (Loveridge 1957). Cer��aco et al. (2018) deduced that the information from Ladiges was based on an old label associated with the presumed remaining type specimen. Because extensive fieldwork on the island of S��o Tom�� over the last century failed to document this species, Cer��aco et al. (2018) concluded that it does not occur there, and the original type locality from Fischer (1856) ���Edina, Grand Bassa County in Liberia (West-Afrika)��� is correct. Moreover, the former authors determined that the pholidosis from this presumed type (Cer��aco et al. 2018:fig. 1) is not consistent with the extensively detailed original description, and thus, it is not a type specimen (contra Uetz et al. 2019). Although we agree with this conclusion, an examination of this specimen (ZMH R04376) by J. Hallermann (pers. comm.) found some differences in the morphometric data provided by Cer��aco et al. (2018) (e.g., SVL = 635 mm, not 880 mm), and we used these updated data in our analyses of morphological data (Table 2). Chabanaud (1917a:375), convinced that the odd configuration of subcaudals (combination of single and double subcaudals near the vent) seen in one male and one female was unique to Colubridae, named the taxon Dipsadomorphus boueti from these specimens that were donated by ���Dr Bouet,��� who collected for the Paris Museum in Ivory Coast (1909) and Dahomey (1910���1913). Unfortunately no specific type locality was provided. Chabanaud (1917b) then clarified and corrected himself by stating that both types were males (the mistaken female was a juvenile), and they were collected from ���Porto-Novo��� in Dahomey, which is modern-day Porto-Novo, the capital city of Benin. Chabanaud (1917c) subsequently realized that his specimens of D. boueti represented T. pulverulenta individuals with abnormal subcaudals, and he synonymized the former taxon with the latter (as D. pulverulentus). We agree with this taxonomic placement, because the types of D. boueti have subcaudal counts (106 and 116) and TL/SVL ratios (0.25 and 0.28) that are most consistent with T. pulverulenta (Table 2). The Porto-Novo locality is on the eastern edge of the Dahomey Gap (sensu Demenou et al. 2016:fig. 1), a renowned biogeographic barrier to at least some forest-specialist species in West Africa. Because T. pulverulenta does not seem to be a strict forest specialist (see Habitat below), and the Porto-Novo locality occurs on the eastern edge of the Dahomey Gap, it is likely that the Gap is not a barrier to dispersal for this species. We therefore hypothesize that the Niger Delta is the biogeographic barrier separating T. pulverulenta from its cryptic sister taxon T. adamanteus sp. nov. Diagnosis. A species of Toxicodryas restricted to West Africa, west of the Niger Delta, defined by the following combination of characters: maximum SVL 1 meter in T. blandingii and T. vexator sp. nov.), DSRN 19���21 (vs. 23���25 in T. blandingii and 23���29 in T. vexator sp. nov.), DSRM 19���21 (vs. 21���25 in T. blandingii and T. vexator sp. nov.); cloacal plate undivided (vs. usually divided in T. blandingii, and divided or undivided in T. vexator sp. nov.); both sexes brown to pink with darker cross-bars that often enclose a whitish spot, and the dorsum and venter sprinkled with fine dark brown or black spots (vs. adult males glossy or velvety black with a yellow venter, and adult females light brown, gray, or yellowish-brown with light-brown or cream cross-bars on the flanks, with yellowish-brown venters in T. blandingii and T. vexator sp. nov.); hemipenis relatively long with long spines mid-way along the shaft that decrease in size towards the apex and base, with a domed apex (vs. hemipenis relatively short and massive [i.e., broad], proximal third covered with spines, distal two-thirds dimpled with a flattened apex in T. blandingii and T. vexator sp. nov.). Variation. Morphometric variation of Toxicodryas pulverulenta is shown in Table 2. Chabanaud (1917a) first noted that some individuals (including the type of Dipsadomorphus boueti) can have a few single subcaudals posterior to the cloacal plate. Bogert (1940) provided morphometric data, but he did not distinguish between populations from Liberia (T. pulverulenta) and former French Cameroon (T. adamanteus sp. nov.), except that ���the Liberia specimens are distinctly more reddish brown than the Cameroon specimens.��� However, he listed the maximum total length of a male (1112 mm) and female (1050 mm) T. pulverulenta from Liberia. In snakes from West Africa (without noting sex), including Cameroon where T. adamanteus sp. nov. occurs, Angel (1933:146) noted temporal formula variation of 2 + 2 (rarely 2 + 3 or 3 + 2 or 1 +2), 236���276 ventrals, 96���132 subcaudals, and a maximum size of 1225 mm; nearly identical data were reported by Villiers (1950a), Doucet (1963), Stucki-Stirn (1979), and Chippaux (2006). Although all of our examined specimens and most literature records noted the 3 rd to 5 th supralabial in contact with the eye (e.g., Villiers 1950b), Chippaux (2006) documented individuals with the 4 th to 6 th supralabial in contact with the eye, and sometimes, only two scales in contact with the eye, but some of these specimens might be attributable to T. adamanteus sp. nov. Segniagbeto et al. (2011) reported snakes from Togo with 165���175 ventrals, which is undoubtedly erroneous. Fischer (1856:83) noted his type specimen had seven maxillary teeth that were oriented nearly backwards, becoming larger posteriorly. These were followed by two larger teeth (i.e., fangs) that were furrowed in a ���besonderen Hauttasche��� [special skin pocket]. There were 10���12 mandibular teeth, which were slightly curved towards the back of the mouth, and increased in size posteriorly. Bogert (1940:61) remarked ��� five specimens examined show variation from eleven to thirteen anterior subequal teeth followed after a very short diastema by two larger grooved fangs and a smaller fang, the total number of teeth being fourteen to eighteen.��� Because only two of his examined specimens originated from former French Cameroon (attributable to T. adamanteus sp. nov.), at least three of these specimens were from Liberia, which are attributable to T. pulverulenta. In snakes from Ghana, Leeson (1950) noted 11���13 maxillary teeth, becoming slightly larger posteriorly, followed by 2 fangs, and sometimes a 3 rd, smaller fang; 14���16 palatine and pterygoid teeth, and 15 mandibular teeth with the anteriormost ones largest. Johnsen (1962) reported that his specimen from Liberia had 11 maxillary teeth on one side, and six on the other. Based on a specimen from Liberia, Malnate (1972) noted the species lacks posterior hypapophyses. In his original description of the species, Fischer (1856:83) described the coloration in great detail as chocolate brown above, yellow below, finely dotted everywhere. On each side, near the back, there was a large number (60��� 70) of pale red spots that lacked dark edging and extended over 4���6 scales. Usually the spots on one side alternated with those on the other; sometimes they also were opposite to each other and in this case were connected to weak ���Querbinden��� [cross-ties] by bright red connecting strips that extended over the back. Usually there was a small black spot under each of these spots on the outermost tip of the corresponding ventral shields. The innumerable fine black points, with which the whole body was sown, were grouped on the abdominal shields on each side at the point where they bent over to rise sideways, to form a black spot, which in their succession looked like a black longitudinal band, whereby the narrow belly appeared delimited from the flanks. Head was brown and without black lines. Upper lip, lower lip, throat yellow, dotted with black. G��nther (1858:173) described the dorsum of two Nigerian specimens as ���brown with a strong cast of purple��� with elliptical transverse streaks, and at mid-body, these streaks had a small yellow spot in their center. In his description of Dipsadomorphus boueti, Chabanaud (1917a) noted the types were light brownish gray and dotted with brown, whereas the supralabials and venter were yellowish gray. Leeson (1950) described animals from Ghana as brownish red on the dorsum, with ���dull brown��� heads, and ���lightly coloured��� patches on the flanks that had a black edge on the lower margins. There were more narrow light gray bands on the anterior third of the body that gradually diminished to form light gray patches on the posterior two-thirds of the body. The ventral surface of the head was white or cream, whereas the venter of the body was pinkish with numerous brown spots. Brown lines ���commence���[ed] just before the middle of the body on each side of the ventrals, and continued along the edge of the ventrals to the tip of the tail. In his description of two specimens from Ivory Coast, Villiers (1950b) noted their color pattern was pale brownish gray, the dorsal scales dotted with black, the back with alternate dark brown diamond spots (some ocellated with white). Yellowish ventral surface with two black lateral lines. Doucet (1963) noted the dorsal coloration ranged from uniform reddish or yellowish or powdery brown, sometimes with dark bars, with ventral coloration ranging from yellowish to pinkish. Chippaux (2006) noted dorsal coloration as dark beige or reddish with occasional dark gray designs or crossbars; venter pinkish with two dark lateral lines. Based on photographs of an adult male from Guinea (Fig. 11B), the base of the tongue is orangish red, and the forked tip is silvery white with black edging. Our observations of live specimens suggest the dorsal scales have an almost satiny sheen, similar to the appearance of a spider web (MOR, pers. obs.). Hemipenis. Bogert (1940) did not note whether his hemipenis description was based on specimens from Liberia (T. pulverulenta) or former French Cameroon (T. adamanteus sp. nov.). Doucet (1963:301) described the hemipenis of a snake from Ivory Coast as very different from that of B. [Toxicodryas] blandingi. Quite elongated, thorny at the base and dimpled at the apex, it is not bifid. The spines on either side of the sperm groove are very long in the middle part and decrease in size towards the apex and the base. The apex is domed, not flattened. Diet. Villiers (1950a) noted both species of Toxicodryas in West Africa feed mainly on birds. Cansdale (1954) reported an individual from Ghana with two young mice in its stomach. Johnsen (1962:121) reported an adult male from Liberia with ���a small insect-eating bird��� in its stomach. In their paper on snakes of Ghana, Leston & Hughes (1968:754) noted the presence of undigested hair and a 10-cm long rodent tail in the stomachs of two specimens, and that ���this suggests the attraction of Cocoa is for its Rodent fauna.��� Another specimen contained the foot and tail of an Agama sp. lizard (sensu Leach�� et al. 2017). B��hme (2000) noted the diet of snakes from Guinea included small mammals (shrews and rodents), adult and nestling birds, and lizards with well-developed limbs. Greene (1989) suggested that there is an ontogenetic shift in diet from mostly lizards in young individuals to mostly birds and/or mammals as adults, and some individuals can ingest a third of their body weight if they eat more than one prey item in a nest or roost. Behavior. In a paper on Guinean snakes, B��hme (2000) classified the species as relatively common and nocturnal. Reproduction. An adult female found in Ghana in early September contained eggs ���about 8 mm maximum breadth and between 30 and 35 mm long��� attached to the left and right ovaries (Leston & Hughes 1968). Habitat. Menzies (1966) documented several specimens from forest in Sierra Leone, but one animal found at Rokupr seemed to occur in a habitat labeled as mangrove and coastal savanna. Leston & Hughes (1968:754) noted that this species was very common in cocoa tree pods in Ghana. Hughes & Barry (1969) listed the habitat as forest. Leston (1970:143) encountered several snakes in Ghana in trees (including one in a rotten cocoa pod), ���amidst old files on an office shelf,��� on a 1.5-meter high cocoa tree in a greenhouse, and in savanna. Hughes (1988) noted the species only from rainforest in Ghana. R��del & Mahsberg (2000) found snakes in Ivory Coast from thick undergrowth in a swamp bordered by fields and secondary forest, and swampy primary forest. Branch & R��del (2003) captured a juvenile in a funnel trap on the ground in closed-canopy forest in Ivory Coast. In Togo, Segniagbeto et al. (2011) noted the species is most common in forest, but some specimens were found outside of it, presumably in savanna. Geographic distribution and habitat. Based on molecular data from Allen et al. (in press) and patterns of our morphometric data (Table 2), we hypothesize that this species occurs west of the Niger Delta from Guinea to Nigeria. Venom. Venom composition and effects are unknown (Weinstein et al. 2011)., Published as part of Greenbaum, Eli, Allen, Kaitlin E., Vaughan, Eugene R., Pauwels, Olivier S. G., Wallach, Van, Kusamba, Chifundera, Muninga, Wandege M., Aris- Tote, Mwenebatu M., Mali, Franck M. M., Badjedjea, Gabriel, Penner, Johannes, R��del, Mark-Oliver, Rivera, Jacqueline, Sterkhova, Viktoria, Johnson, Grant, Tapondjou, Walter P. & Brown, Rafe M., 2021, Night stalkers from above: A monograph of Toxicodryas tree snakes (Squamata Colubridae) with descriptions of two new cryptic species from Central Africa, pp. 1-44 in Zootaxa 4965 (1) on pages 24-27, DOI: 10.11646/zootaxa.4965.1.1, http://zenodo.org/record/4723024, {"references":["Fischer, J. G. (1856) Neue Schlangen des Hamburgischen Naturhistorischen Museums. Abhandlungen aus dem Gebiete der Naturwissenschaften herausgegeben von Naturwissenschaftlichen Verein in Hamburg, 3, 79 - 116 + 3 pl.","Chabanaud, P. (1917 a) Enumeration des ophidiens non encore etudies de l'Afrique occidentale, appartenant aux collections du Museum avec la description des especes et des varietes nouvelles. Bulletin du Museum d'Histoire Naturelle, Paris, 22, 362 - 382. https: // doi. org / 10.5962 / bhl. part. 17132","Chabanaud, P. (1917 b) Note complementaire sur les ophidiens de l'Afrique occidentale, avec la description d'une espece nouvelle. Bulletin du Museum d'Histoire Naturelle, Paris, 23, 7 - 14.","Ceriaco, L. M. P., Marques, M. P. & Bauer, A. M. (2018) Miscellanea Herpetologica Sanctithomae, with a provisional checklist of the terrestrial herpetofauna of Sao Tome, Principe and Annobon islands. Zootaxa, 4387 (1), 091 - 108. https: // doi. org / 10.11646 / zootaxa. 4387.1.4","Hallermann, J. (1998) Annotated catalog of the type specimens of the herpetological collection in the Zoological Museum of the University of Hamburg. Mitteilungen aus dem Hamburgischen Zoologischen Museum und Institut, 95, 197 - 223.","Uetz, P., Cherikh, S., Shea, G., Ineich, I., Campbell, P. D., Doronin, I. V., Rosado, J., Wynn, A., Tighe, K. A., McDiarmid, R., Lee, J. L., Kohler, G., Ellis, R., Doughty, P., Raxworthy, C. J., Scheinberg, L., Resetar, A., Sabaj, M., Schneider, G., Franzen, M., Glaw, F., Bohme, W., Schweiger, S., Gemel, R., Couper, P., Amey, A., Dondorp, E., Ofer, G., Meiri, S. & Wallach, V. (2019) A global catalog of primary reptile type specimens. Zootaxa, 4695 (5), 438 - 450. https: // doi. org / 10.11646 / zootaxa. 4695.5.2","Hughes, B. & Barry, D. H. (1969) The snakes of Ghana: a checklist and key. Bulletin de l'Institut fondamental d'Afrique noire. Serie A, Sciences naturelles, 31 (3), 1004 - 1041.","Loveridge, A. (1957) Check list of the reptiles and amphibians of East Africa (Uganda; Kenya, Tanganyika; Zanzibar). Bulletin of the Museum of Comparative Zoology, Harvard, 117 (2), 151 - 362 + i - xxxvi (index).","Chabanaud, P. (1917 c) Revision de quelques reptiles d'Afrique et description de trois especes nouvelles. Bulletin du Museum d'Histoire Naturelle, Paris, 23, 442 - 454.","Demenou, B. B., Pineiro, R. & Hardy, O. J. (2016) Origin and history of the Dahomey Gap separating West and Central African rain forests: Insights from the phylogeography of the legume tree Distemonanthus benthamianus. Journal of Biogeography, 43 (5), 1020 - 1031. https: // doi. org / 10.1111 / jbi. 12688","Bogert, C. M. (1940) Herpetological results of the Vernay Angola expedition. Bulletin of the American Museum of Natural History, 77, 1 - 107, pl. I.","Angel, F. (1933) Les Serpents d'Afrique Occidentale Francaise. Larose Editeur, Paris, 246 pp.","Villiers, A. (1950 a) Initiations Africaines. II. Les Serpents de l'Ouest Africain. Institut Francais d'Afrique Noire, Dakar, Senegal, 148 pp.","Doucet, J. (1963) Les serpents de la Republique de Cote d'Ivoire. Serpents venimeux. Acta Tropica, 20 (3 - 4), 297 - 340.","Stucki-Stirn, M. C. (1979) A Comparative Study of the Herpetological Fauna of the former West Comeroon [sic] / Africa: With a Classification and Synopsis of 95 Different Snakes and Description of Some New Sub-species. Snake Report 721. Herpeto- Verlag, Teuffenthal, Switzerland, 650 pp.","Chippaux, J-P. (2006) Les Serpents d'Afrique Occidentale et Centrale. 3 rd Edition. IRD Editions, Paris, 311 pp.","Villiers, A. (1950 b) Catalogues. IV. La Collection de Serpents de l'I. F. A. N. Institut Francais d'Afrique Noire, Dakar, Senegal, 155 pp.","Segniagbeto, G. H., Trape, J. F., David, P., Ohler, A., Dubois, A. & Glitho, I. A. (2011) The snake fauna of Togo: Systematics, distribution and biogeography, with remarks on selected taxonomic problems. Zoosystema, 33 (3), 325 - 360. https: // doi. org / 10.5252 / z 2011 n 3 a 4","Leeson, F. (1950) Identification of Snakes of the Gold Coast. The Crown Agents for the Colonies, London, x + 142 pp.","Johnsen, P. (1962) Notes on African snakes, mainly from northern Rhodesia and Liberia. Videnskabelige Meddelelser fra Dansk Naturhistorisk Forening i Kjobenhavn, 124, 115 - 130.","Malnate, E. V. (1972) Observations on the vertebral hypapophyses and associated musculature in some snakes, with special reference to the Colubridae. Zoologische Mededelingen, 47 (18), 225 - 239.","Gunther, A. (1858) Catalogue of Colubrine Snakes in the Collection of the British Museum. Trustees of the British Museum, London, xvi + 281 pp. https: // doi. org / 10.5962 / bhl. title. 13272","Cansdale, G. (1954) Gold Coast snakes - a complete list. The Nigerian Field, 19, 118 - 132.","Leston, D. & Hughes, B. (1968) The snakes of Tafo, a forest cocoa-farm locality in Ghana. Bulletin de l'Institut Francaise d'Afrique Noire, 30 A, 737 - 770.","Leache, A. D., Grummer, J. A., Miller, M., Krishnan, S., Fujita, M. K., Bohme, W., Schmitz, A., Lebreton, M., Ineich, I., Chirio, L., Ofori-Boateng, C., Eniang, E. A., Greenbaum, E., Rodel, M. - O. & Wagner, P. (2017) Bayesian inference of species diffusion in the West African Agama agama species group (Reptilia, Agamidae). Systematics and Biodiversity, 15 (3), 192 - 203. https: // doi. org / 10.1080 / 14772000.2016.1238018","Bohme, W. (2000) Diversity of a snake community in a Guinean rain forest (Reptilia, Serpentes). In: Rheinwald, G. (Ed.), Isolated Vertebrate Communities in the Tropics. Proceedings of the 4 th International Symposium in Bonn. Bonner zoologische Monographien, 46, 69 - 78.","Greene, H. W. (1989) Ecological, evolutionary, and conservation implications of feeding biology in Old World cat snakes, Genus Boiga (Colubridae). Proceedings of the California Academy of Sciences, 46 (8), 193 - 207.","Menzies, J. I. (1966) The snakes of Sierra Leone. Copeia, 1966 (2), 169 - 179. https: // doi. org / 10.2307 / 1441123","Leston, D. (1970) Some snakes from the forest zone of Ghana. British Journal of Herpetology, 4 (6), 141 - 144.","Hughes, B. (1988) Herpetology in Ghana (West Africa). British Herpetological Society Bulletin, (25), 29 - 37.","Rodel, M. - O. & Mahsberg, D. (2000). Vorlaufige Liste der Schlangen des Tai-Nationalparks / Elfenbeinkuste und angrenzender Gebiete. Salamandra, 36 (1), 25 - 38.","Branch, W. R. & Rodel, M. - O. (2003) Herpetological survey of the Haute Dodo and Cavally forests, western Ivory Coast, Part II: Trapping results and reptiles. Salamandra, 39 (1), 21 - 38.","Weinstein, S. A., Warrell, D. A., White, J. & Keyler, D. E. (2011) \" Venomous \" Bites from Non-Venomous Snakes: A Critical Analysis of Risk and Management of \" Colubrid \" Snake Bites. Elsevier, Amsterdam, Boston, Heidelburg, London, New York, Oxford, Paris, San Diego, San Francisco, Singapore, Sydney and Tokyo, xxvii + 336 pp."]}

Toxicodryas adamanteus sp. nov. (Table 2, Figs. 8, 12–13) We hypothesize that this new, cryptic species occurs in western, central and eastern Africa, east of the Niger Delta, and it has been considered to be conspecific with Toxicodryas pulverulenta since it was first documented to occur in Angola by Peters (1877). The recognition of this new species is supported by evidence from molecular data (Fig. 2) and significant differences in subcaudal scale counts (see Results). Diagnosis. A species of Toxicodryas restricted to West, Central and East Africa, east of the Niger Delta, defined by the following combination of characters: maximum SVL 1 meter in T. blandingii and T. vexator sp. nov.), DSRN 18–23 (vs. 23–25 in T. blandingii and 23–29 in T. vexator sp. nov.), DSRM 18–21 (vs. 21–25 in T. blandingii and T. vexator sp. nov.); cloacal plate undivided (vs. usually divided in T. blandingii, and divided or undivided in T. vexator sp. nov.); both sexes brown to pink with darker cross-bars that often enclose a whitish spot, and the dorsum and venter sprinkled with fine dark brown or black spots (vs. adult males glossy or velvety black with a yellow venter, and adult females light brown, gray, or yellowish-brown with light-brown or cream cross-bars on the flanks, with yellowish-brown venters in T. blandingii and T. vexator sp. nov.); hemipenis relatively long with long spines mid-way along the shaft that decrease in size towards the apex and base, with a domed apex (vs. hemipenis relatively short and massive [i.e., broad], proximal third covered with spines, distal twothirds dimpled with a flattened apex in T. blandingii and T. vexator sp. nov.). Holotype. UTEP 22204 (field no. ELI 2213; Figs. 8, 12–13), adult male from Npenda village, NE of Lake Tumba (00.7465° S, 18.2243° E, 311 m), Equateur Province, DRC, collected by local Twa people and brought to Eli Greenbaum, Chifundera Kusamba, Wandege M. Muninga, and Mwenebatu M. Aristote on 8 July 2013. Paratopotype. UTEP 22203 (field no. ELI 2212) adult female with same collection details as the holotype. Paratype. RBINS 2699 (formerly RBINS 9127) (field no. Paul Leloup #26), adult female from Région Tshabondo (2.690861° S, 27.341972° E), South Kivu Province, DRC, collected by Paul Leloup on 11 October 1958. Description of the holotype. Adult male, 812 mm SVL; head weakly triangular and moderately distinct from neck, 2.4% of SVL (19.3 mm); interocular distance 13.4 mm, pupil elliptical, eye diameter 5.2 mm; loreal nearrectangular, shorter (1.7 mm) than high (2.0 mm), slightly tapering superiorly and vertically; body semi-triangular; tail moderately long (27.7% of SVL). Supralabials 8/8, 3 rd –5 th /3 rd –5 th contacting orbit; infralabials 13/12, 1 st on each side in contact behind mental, 1 st –5 th /1 st –5 th contacting anterior chin shields and 5 th –7 th /5 th –7 th contacting posterior chin shields; 1/1 preocular; 2/2 postoculars; temporals 2 + 3 + 3/2 + 3 + 3; 2 internasals; nasal divided; frontal width and length equal (6.6 mm); dorsal scale rows 21 one head length posterior to jaw rictus, 19 at midbody, 15 one head length anterior to vent, smooth and oblique with apical pits visible on the neck, vertebral scales broad and apically flattened; ventrals 264 (standard), 262 (Dowling); cloacal plate undivided; paired subcaudals 117; both hemipenes everted. Coloration (in life) of the holotype. Silvery gray-brown over the entire length of the dorsum including the head, with similarly colored but darker diamonds occurring laterally over the full length of the neck, body, and tail. These diamonds are elongated vertically, have pale gray centers, and often have black spots at the top and bottom corners. Between each diamond on the body and neck is a more vaguely defined, pale gray diamond or stripe. The lateral and antero-dorsal sides of the head are speckled, as is the ventral side of the head, neck, body, and tail. The venter is otherwise white, with the speckles forming two pale stripes running down either side of the ventrals and subcaudals. The base of the tongue is orangish red, and the forked tip is silvery white with black edging. Coloration (in preservative) of the holotype. This specimen’s coloration has become slightly darker and browner than it was in life, with less pronounced patterning on the flanks, but otherwise the appearance is similar to the coloration in life. Variation. Morphometric variation of Toxicodryas adamanteus sp. nov. is shown in Table 2. We observed extensive temporal scale variation, including 1 + 1, 1 + 2, 2 + 2 (most commonly), 2 + 3, 2 + 5, 3 + 2, 3 + 3, and 4 + 3. Chabanaud (1917c) described a male from Gabon with supralabials 3–6 contacting the eye, and this specimen seems to be the size record at 1,225 mm total length (995 mm SVL, 230 mm tail length). Schmidt (1923) listed ventral counts of 251–269 in snakes from DRC, and noticed one individual that had a preocular fused to the supraocular on one side, and in two individuals, the loreal was fused to the lower portion of the preocular and thus contacted the eye. Laurent (1956) noticed that snakes from DRC (i.e., T. adamanteus sp. nov.) had more subcaudals than snakes from West Africa (i.e., T. pulverulenta). Skinner (1973) noted ventral scale counts range from 240–269, subcaudal scale counts range from 105–126 (identical to Pitman 1974), and the maximum size is about 2 meters, substantially larger than all other published records, and thus, highly doubtful. De Witte (1975) provided data for ventral scale counts of DRC snakes ranging from 235–242 (males) and 239–249 (females), and subcaudal counts of 112–120 (both sexes). Spawls & Branch (2020) listed the maximum size as “about 1.25 m ” but no specific locality or record was provided. Although most of our examined specimens and literature records noted the 3 rd to 5 th supralabial in contact with the eye, some individuals have the 4 th to 6 th supralabial in contact with the eye (e.g., Loveridge 1937) and one specimen from Gabon (CAS 258155) had only the 4 th and 5 th supralabial in contact with the eye. In snakes from Uganda, Pitman (1974) reported ventral scale counts of 240–269 and subcaudal counts of 105–126 (not distinguished by sex). In de Witte’s (1975) study of snakes from Virunga National Park in eastern DRC, he noted ventrals range from 235–242 in males and 239–249 in females; subcaudals ranged from 112–120 in both sexes. Rasmussen (1997b:106) described individuals with 3 or 4 postoculars, 7–9 supralabials, 9–13 infralabials, 236–278 ventrals, and 96–132 subcaudals (sometimes undivided). In general, he noted this species has sloping and smooth scales with apical pits, and the vertebral row is more or less enlarged. Bogert (1940:fig. 8) illustrated the maxillary teeth of a specimen (AMNH 50590) from former French Cameroon, noting “ five specimens examined show variation from eleven to thirteen anterior subequal teeth followed after a very short diastema by two larger grooved fangs and a smaller fang, the total number of teeth being fourteen to eighteen.” Because only two of his examined specimens originated from former French Cameroon, at least three of these specimens are from Liberia, which are attributable to T. pulverulenta. Schmidt (1923:103) quoted field notes of Herbert Lang for DRC snakes as “coloration, in life, reddish brown above, head darker brown. Irregular dark gray lateral bars, wider in the middle, extend from the vertebral line to the venter, tipped above and below with black. A cream-colored central spot in the broad portion of each lateral bar. Faint narrow grayish crossbars between the wider ones, disappearing posteriorly. The wider crossbars are usually alternate, sometimes confluent on the back. Venter pinkish gray, heavily dotted with brown which forms two lines at the inner edges of the ventral edges of the ventral angle. These lines are more distinct beneath the tail.” Hellmich (1957b) described several individuals from Angola with a reddish brown to reddish blue-gray dorsum, with varying degrees of contrasting rhombic patterns and spots. Pitman (1974) provided a similar description for specimens from Uganda, noting they range from pinkish-brown to reddish-brown. Hedges (1983:21) described Kenyan specimens as, “a rather pretty pattern of coffee coloured diamond shaped markings on a mushroom pink body.” Rasmussen (1997b) described coloration in this species (presumably based on examined specimens from Nigeria and Cameroon) as reddish brown on the dorsum of the body; dorsum of the head darker brown; irregular, dark gray, diamondshaped spots on the flanks that are widest at mid-body; spots edged in black on superior and inferior edges with a cream spot in the center that sometimes fuse together in the midline; faint grayish transverse bands between spots that fade and disappear posteriorly; venter pinkish-gray, with dense brown spots that form a line just inside and parallel to the keeled ventrals, most salient on the tail. Pauwels et al. (2019b) noted a specimen from Gabon with an orange tongue in life. Spawls & Branch (2020:239) noted the dorsal color can range from pinkish to brown, redbrown or pinky gray with “darker” cross-bars that enclose a “pale” spot. The dorsum and venter are “finely dusted” with brown or black specks, whereas the venter is pale pink with “dashed dark lines” on each side of the ventrals. Contrary to Pauwels et al. (2019b), they noted the tongue is pink with a white tip. Based on our photographs of DRC snakes (e.g., Fig. 12G), the tongue is orangish red, and the forked tip is silvery white with black edging, which is consistent with the coloration of the holotype in life (EG pers. obs.). An unsexed individual from Banalia, DRC (Fig. 12F) is unique in having a golden yellow mid-dorsal stripe. Diet. Pitman (1938) mentioned a juvenile from Uganda that had a small mouse in its stomach. In Kenya, Hedges (1983) noted the species eats small chameleons, geckos, and frogs. Butler & Reid (1990) described an adult female from Nigeria that contained the remains of an Agama sp. lizard (sensu Leaché et al. 2017). Spawls & Branch (2020) listed the diet as arboreal lizards and rodents. Parasites. Pitman (1938) noted a juvenile from Uganda with numerous ticks. Behavior. Pitman (1974:128) described the species as “arboreal, nocturnal... amiable disposition,” and suggested that it might rely on camouflage to avoid predation, a sentiment also suggested by Gans (1961). Chirio & LeBreton (2007) suggested the species is shy and rarely bites. Spawls & Branch (2020) noted that when threatened, it elevates the anterior part of its body, “flickers its red tongue,” hisses, and strikes. Reproduction. Schmidt (1923) described a DRC female collected in June 1914 with an unspecified number of eggs that measured 11 x 29 mm. Butler & Reid (1990) described an adult female from Nigeria (captured 21 January 1988) that contained two developing eggs (34–35 mm long, 11 mm wide). Branch (2005) noted the species lays 2– 5 eggs. Habitat. Parker (1936) noted this species is restricted to rain forest and “its outliers.” Perret (1961) listed the species from forest in Cameroon. De Witte (1962) listed the habitats for DRC snakes as equatorial forest and gallery forest, and this classification was repeated by Thys van den Audenaerde (1965). Blackwell (1967) listed the species from forest, thickets, and gardens in Nigeria. Pitman (1974:128) described this snake as a “forest species” in Uganda, but also noted one individual from a “fowl house,” and another from an epiphytic fern (Platycerium angolense) on a forest tree. Butler & Reid (1986) listed the species only from forest in Nigeria. Lawson (1993) documented the species from forest and “farmbush” in Cameroon. Rasmussen (1997b) described the species as nocturnal and restricted to forest. Chirio & LeBreton (2007) recorded the species from forests and savanna-forest mosaic in Cameroon. Spawls & Branch (2020) included forest, woodland, and forest-savanna mosaic from sea level to about 2,000 m elevation. Among the 19 snake species recorded within the garden of a villa in Yenzi ( 2.77261° S, 10.03403° E), Gamba, Nyanga Province, southwestern Gabon (forest-savanna mosaic), inhabited by one of the authors from 2004 to 2011 (OSGP, unpubl. data), T. adamanteus sp. nov. was among the most rarely encountered species. Geographic distribution and habitat. Based on molecular data from Allen et al. (in press) and patterns of our morphometric data (Table 2), we hypothesize that this species occurs east of the Niger Delta in Nigeria, Cameroon, Equatorial Guinea, Gabon, CAR, Republic of Congo, DRC, Angola, Uganda, South Sudan, and Kenya. Ullenbruch & Böhme (2017) recently listed the species from South Sudan. Venom. Based on a specimen from Cameroon, Taub (1967) described the histological morphology of the Duvernoy’s gland. Venom composition and effects are unknown (Weinstein et al. 2011). Etymology. The specific epithet adamanteus is a Latin adjective referring to the diamond-shaped marks on the flanks and dorsum of this species.

Toxicodryas vexator sp. nov. (Table 1, Figs. 1, 8���10) We hypothesize that this new, cryptic species occurs east of the confluence of the Congo and Ubangi rivers, and it has been considered to be conspecific with Toxicodryas blandingii since it was first documented to occur east of these rivers in DRC (Boulenger 1919). The recognition of this new species is supported by molecular data (Fig. 2), several significant differences in scale counts (see Results), and less toxic venom in comparison to its sister taxon T. blandingii (see Venom). Diagnosis. A species of Toxicodryas restricted to east-central and East Africa (east of the confluence of the Congo and Ubangi rivers), defined by the following combination of characters: maximum SVL> 1 meter (vs. maximum SVL T. pulverulenta and T. adamanteus sp. nov.); DSRN 23���29 (vs. 19���21 in T. pulverulenta and 18���23 in T. adamanteus sp. nov.); DSRM 21���25 (vs. 19���21 in T. pulverulenta and 18���21 in T. adamanteus sp. nov.); cloacal plate divided or undivided (vs. usually divided in T. blandingii, and always undivided in T. pulverulenta and T. adamanteus sp. nov.); adult males glossy or velvety black with a yellow venter, and adult females light brown, gray, or yellowish-brown with light-brown or cream cross-bars on the flanks, with yellowish-brown venters (vs. both sexes brown to pink with darker cross-bars that often enclose a whitish spot, and the dorsum and venter sprinkled with fine dark brown or black spots in T. pulverulenta and T. adamanteus sp. nov.); hemipenis relatively short and massive (i.e., broad), proximal third covered with spines, distal two-thirds dimpled with a flattened apex (vs. relatively long with long spines mid-way along the shaft that decrease in size towards the apex and base, and with a domed apex in T. pulverulenta and T. adamanteus sp. nov.); venom toxicity LD 50 = 4.88 mg /kg in mice (vs. venom toxicity LD 50 = 2.85���3.55 mg /kg in mice for T. blandingii). Holotype. UTEP 22196 (field number MUSE 10341; Fig. 10), adult male collected in Mulisi, Nzovu Est, Kahuzi-Biega National Park (02.447291�� S, 28.2825378�� E, 1101 m), South Kivu Province, DRC, collected by Guillain M. Mitamba, Deo Kujirakwinja, Emmanuel Muhindo, Radar Nushili, Wandege M. Muninga, and Andrew J. Plumptre on 13 November 2015. Paratypes. UTEP 22195 (field number EBG 1362; Figs. 1B, 9C), adult female collected in the vicinity of Irangi (01.8780�� S, 28.4524�� E, 811 m), South Kivu Province, DRC, collected from a tree by a resident and brought to Maurice Luhumyo, Chifundera Kusamba, Mwenebatu M. Aristote, Wandege M. Muninga, and Eli Greenbaum on 30 August 2007; RBINS 2698 (formerly RBINS 8621) (field no. Leloup #27), adult male from Bunyakiri (2.075630�� S, 28.573194�� E, 1000 m), South Kivu Province, DRC, collected by Paul Leloup on 6 March 1958. Description of the holotype. Adult male, 1438 mm SVL; head strongly triangular and distinct from the neck, 2.1% of SVL (30.9 mm); interocular distance 21.7 mm, pupil elliptical, maximum horizontal eye diameter 7.5 mm; loreal near-rectangular, longer (3.0 mm) than high (2.4 mm), slightly tapering superiorly; body triangular; tail moderately long (32.1% of SVL). Supralabials 9/9, 4 th, 5 th, and 6 th /4 th, 5 th, and 6 th contacting orbit; infralabials 14/14, 1 st on each side in contact behind mental, 1 st ���4 th /1 st ���4 th contacting anterior chin shields, 5 th ���8 th /5 th ���8 th contacting posterior chin shields, and 1 cuneate between the 5 th and 6 th left infralabials; 2/2 preoculars; 2/2 postoculars; temporals 2 + 2/2 + 2; 2 internasals; nasal divided on both sides; frontal slightly wider (10.1 mm) than long (9.2 mm); dorsal scale rows 23 one head length posterior to jaw rictus, 21 at midbody, 15 one head length anterior to vent, smooth and oblique with apical pits visible on the neck, vertebral scales broad and apically flattened; ventrals 261 (standard), 260 (Dowling); cloacal plate undivided; paired subcaudals 136 (terminal spine missing); both hemipenes everted. Coloration (in life) of the holotype. Solid black over the entire length of the dorsum, and black ventrally except for the anterior half of the body, where the ventral scales are pale yellow with a black posterior edge that increases in thickness in correlation with increased distance from the head. The head is black dorsally and laterally except for yellow on the lower, anterior region of the 5 th ���9 th supralabials, and yellow ventrally except for thin black rims on the posterior edges of the infralabials. Coloration (in preservative) of the holotype. The specimen���s coloration in preservative is nearly the same as in life, although the ventral yellow coloration has dulled to a yellowish off-white; some scales also seem cloudy under close lighting, indicative of some stage of ecdysis in life. Variation. For years, the size record for this species was attributed to Goodman (1985), who recorded a total length of 2515 mm for an adult female collected in Uganda. However, Hedges (1983:20) measured a specimen from Kenya that was 2707 mm total length, and said it ���is reputed to exceed 3300 mm.��� Weinstein et al. (2011) noted the species may exceed 3.5 meters. Spawls & Branch (1995, 2020) listed the maximum size as ���about 2.8 m ��� (no specific location provided), but Weinstein (pers. comm.) estimated a captive specimen from Kakamega was approximately 3.0 meters total length. Schmidt (1923:104) remarked that two specimens from DRC had fused prefrontals, creating a single transverse scale. Five of his 20 DRC specimens had a divided cloacal plate, ���with indications of a groove in two others,��� and as a result, he stated that the condition of the cloacal plate is ���obviously useless��� as a diagnostic character. Laurent (1956) noted that, with the exception of DRC specimens from Mayombe (north of the Congo River in Kongo Central Province and thus attributable to T. blandingii as recognized herein) with consistently divided cloacal plates, 13 of 31 snakes from other areas of DRC had an undivided cloacal plate. Our examined specimens confirm this sentiment, because 17/ 92 specimens have a divided cloacal plate, and an additional five specimens have a partially divided cloacal plate. Hellmich (1957b) noted maximum sizes of Angolan males (1740 mm SVL; 523 mm TL) and females (1730 mm SVL; 505 mm TL), and aberrant individuals that had 11 supralabials and 9���14 infralabials. De Witte (1966) noted that the species has 17 scale rows at midbody, but this datum is so aberrant that it is likely an error (Pitman 1974; Table 1). Skinner (1973) provided ranges of scale counts (161���274 ventrals, 86���147 subcaudals) that are so aberrant that they are clearly erroneous (at least for lower counts), and noted a maximum length of about 3 meters. For Uganda snakes, Pitman (1974) listed ventral scale counts from 240���260 (males) and 240���259 (females). De Witte (1975) noted a single individual with 25 scale rows at midbody, and Broadley et al. (2003) also noted 25 scale rows as the maximum amount of variation. Based on two specimens from Lukolela, Belgian Congo (AMNH 45907) and Akenge, Belgian Congo (AMNH 12243), Bogert (1940:61) described their maxillary teeth as ���ten in number, followed after a short diastema by three enlarged, grooved fangs, the posterior one of which is smaller than the other two.��� Schmidt (1923:105) described two color phases of DRC specimens, including (1) a black phase with the anterior portion of the venter yellow, each ventral scale bordered with black on its posterior edge, the border increasing in width until the yellow color disappears on the posterior two-thirds of the venter; and (2) a brownish phase, ���with more or less distinct wide dark cross-bars, confluent anteriorly, alternate posteriorly on the vertebral line.��� Pitman (1938:211) described an unsexed individual from Uganda as ���darkish gunmetal with a purplish mottling towards the tail, belly pale yellow from head to tail but with slight brown markings on the posterior half increasing in occurrence towards and on the tail.��� A female from Uganda was described as rich chestnut dorsally and ventrally, with handsome chocolate blotches on the flanks, each one containing a small white spot. On the posterior part of the body, including most of the tail, these blotches were confluent with pale edging. The ventral aspect of the head and the first 50 ventrals were ���paly yellowish tinged green.��� The dorsum of the head was dark brown, and the posterior supralabials were dull greenish grey. The supralabials had black edging posteriorly, except for the last one. There were two elongated dark brown blotches behind the eye, which was iridescent hazel. Bogert (1940:61) provided data for a specimen (AMNH 45907; determined to be a subadult male based on photographs examined by EG) that had been described by James P. Chapin as ���olive-brown, with dark brown patches; below grayish brown.��� Hellmich (1957b) described the coloration of several individuals from Angola, including an adult male that had a blue-reddish-black dorsum with a venter that was pale yellow in the front half of the body, with an ever-widening blue-black border that eventually enclosed the entire ventral scales towards the posterior side. Another adult male had a dark black-brown dorsum, and light yellow venter on the first third of the body, but starting on the 5 th ventral, there was a dark gray-blue spot that increased in size posteriorly until it covered the entire venter towards the posterior end of the body. An adult female���s dorsum was described as clay yellow with mostly alternating dark transverse bars and a pale yellow venter. A second adult female had similar coloration to the latter specimen, but the transverse bands were only faintly visible on the posterior sixth of the body and tail. A juvenile dorsum was described as gray-brown with transverse black-brown spots that converge in the anterior part of the body, but then alternate posteriorly, with a light yellowish-white spot in the lower center of the spots. Laurent (1956:195) listed a section about possible sexual dimorphism for this species, but stated that nothing glaring appears in his data. Pitman (1974:125) described several individuals from Uganda that were similar to the descriptions above, and added that both color phases ���exhibit a handsome suede effect.��� He noted that although subadult males might retain the brown phase, black females are unknown, and the black and brown color phases have ���sexual significance.��� The dark blotches of juveniles were noted to vary from ���blackish, chocolate, reddishbrown or dark or bright chestnut,��� with interspaces ranging from light gray, pale brown or chestnut, sometimes with a pinkish tinge. Goodman (1985:56) described two adult females from Uganda as ���dingy olive-brown.��� Hughes (2000) noted that Laurent (1964) seemed to imply that his largest male specimen from Dundo, Angola was not black, but this is not clear, because the latter author only mentioned in passing that a smaller male from Andrada, Angola represented the black phase. Spawls & Branch (1995, 2020) remarked that the yellow ventral pigmentation might form a stripe in the middle of the venter. In the brown color morph, the skin between the scales is bluish gray, and especially visible when the snake inflates its body during a threat display. The eye can be yellowish or brown. The black phase snakes are usually male, whereas the brown phase ones are usually female. Based on photos of an adult male from Banalia, DRC (individual shown in Fig. 9A), the base of the tongue is bluish black, and the forked tip is silvery gray. Hemipenis. Bogert (1940:61) described the hemipenis of an individual, presumably from DRC, as ���not bifurcate, extending to the tenth caudal, the undivided sulcus deeply buried between two fleshy fold [sic]. The basal half is heavily armed with stout spines but distally the organ is calyculate, the edges of the calyces crenulated.��� Hemipenes of our examined specimens had a simple, subcylindrical shape, simple sulcus spermaticus, and spinose ornamentation with a rough apical structure, as also noted for T. blandingii. Because only two specimens with everted hemipenes were available for both T. blandingii and T. vexator sp. nov., any finer-scale differences cannot be attributed to species-level differentiation (Dowling & Savage 1960; Doucet 1963). Diet. Laurent (1964) noted a juvenile snake contained the remains of a bird. In snakes from Uganda, Pitman (1974) noted the diet included weaver finches, robber birds��� eggs, small rodents, bats, frogs, large agamids, and chameleons. Hedges (1983) described a Kenyan snake that regurgitated a Nectarinia famosa sunbird. Spawls & Branch (1995, 2020) described the diet as birds, bird eggs, arboreal lizards, frogs, arboreal rodents, and bats. Nagy et al. (2011) documented an adult female from DRC that predated a Short-palated Fruit Bat (Casinycteris argynnis) from a bat net in secondary forest. Another male specimen from DRC was captured with a nestling bushshrike (Laniarius sp.) in its stomach. Parasites. Loveridge (1937) noted a tick on the scales of a specimen (ANSP 20504) from Belgian Congo. Parasites of Ugandan snakes included internal nematodes (Kalicephalus sp.) and cestodes, ectoparasites (ticks, including Aponomma latum), and captive specimens had bacterial infections, including mouth rot (Pitman, 1974). Goodman (1985:56) documented dozens of ticks, ���probably Aponomma sp.,��� and linguatulid worms from the posterior saccular lung cavity of an adult female from Uganda. Behavior. Spawls & Branch (1995, 2020) described the snake as arboreal, sometimes climbing to heights of 30 meters in trees, but also descending to the ground to cross open spaces and roads. It is mostly nocturnal, sheltering in leaf clumps and tree hollows during the day. When threatened, it will open its mouth widely and expose the pink lining inside, inflate its body, flatten the head, lift the anterior part of its body off the ground and into C-shaped coils, and strike at perceived threats. It is adept at smelling sleeping birds in nests at night, and will make a slow, deliberate approach to attack them. Reproduction. A personal communication from ���Leakey��� (most likely Richard Leakey) to Pitman (1974) noted that a female from Kakamega (Kenya) laid 9 eggs that were approximately 20 x 40 mm on 9 January. Spawls & Branch (1995) reported an average of 7��� 14 eggs in this species (3���14 in Spawls & Branch 2020), presumably based mostly on East African data, which is substantially more than clutch sizes reported for T. blandingii by Luiselli et al. (1998a). Habitat. Laurent (1954) remarked that an adult female from Dundo, Angola was captured after it fell out of a mango tree. Laurent (1960) stated that his colleague M. Leloup often encountered the species in eastern DRC. De Witte (1962) listed the habitats for DRC snakes as equatorial forest and gallery forest, a classification that was repeated by Thys van den Audenaerde (1965). Pitman (1974) also limited the species to forest in Uganda, but he remarked that the species frequently enters houses (where it hunts for bats), and one individual was killed in a tree over 20 meters above the ground. Broadley & Cotterill (2004) stated the species inhabits forests and wooded savannas. Spawls & Branch (1995, 2020) noted the species from forest, woodland, forest-savanna mosaic, riverine woodland, and human habitations from sea level to about 2,200 m elevation. The holotype was found basking on a shrub (ca. 4 meters above ground) in a clearing created by a fallen tree in primary forest. The Irangi paratype was found in secondary forest near a road. Geographic distribution. Based on molecular data from Allen et al. (in press) and patterns of our morphometric data (Table 1), we hypothesize that this species occurs east of the Congo and Ubangi Rivers in CAR, DRC, Angola, Zambia, South Sudan, Uganda, Kenya, and Tanzania. Although Boulenger (1896) listed a specimen from ��� Zanzibar,��� the locality likely referred to mainland records (Parker et al. 1940; Loveridge 1957; Pitman 1974). Venom. Wakeman (1966) documented a 2.59 m (presumably total length) captive adult from Uganda that struck an adult rat, which died after 8.5 minutes. Pitman (1974:127) described the case of a young European who was bitten on the thumb three times in rapid succession by a small (305 mm) individual (erroneously reported as 350 mm by Goodman 1985); only ���initial smarting��� at the puncture sites occurred. Spawls (1979) described the bite of a captive 2.1 meter snake on a man, including three fang punctures on his thumb, which resulted in mild swelling of his hand. Anecdotal observations of a bite by a captive snake on a mouse resulted in death in under a minute. Hedges (1983:20) was bitten, ���and although it drew a considerable amount of blood, I felt no effects whatsoever.��� He also remarked that this species ���chews��� when biting, ostensibly to ensure the rear fangs penetrate the flesh to inject venom. Based on venom collected from a snake originating from Kenya, Weinstein & Smith (1993) estimated the LD 50 value as 4.88 mg /kg in mice. Weinstein & Smith (1993:88) also noted ���the differences in lethal potencies between the two samples of B. blandingi [compare with LD 50 = 2.85���3.55 mg /kg in T. blandingii from West Africa] demonstrate variation of secretion toxicity in this species,��� suggesting possible species-level differences in venom composition of these sister taxa (Weinstein & Kardong 1994). Immunological cross reactivity with elapid venom antisera was demonstrated, but Weinstein & Smith (1993) did not distinguish between samples originating from West Africa (i.e., T. blandingii) or East Africa (i.e., T. vexator sp. nov.). Branch (2005) suggested bites cause nausea and headaches, but fatalities are unknown. Dashevsky et al. (2018) included a Toxicodryas from Tanzania in their proteomics study, and they noted its venom is dominated by three-finger toxins. Scott Weinstein (pers. comm., October 2020) clarified that a snakebite mentioned by Weinstein & Smith (1993) and a second one that occurred one year after this publication were caused by snakes originating from Kakamega forest, Kenya. ���The bite mentioned in the paper was delivered by [an approximately] 3.0 m jet black male that was a favourite member of my collection for nearly 7 years. He typically performed the whole gamut of open mouthed posturing and mock strikes, but was also a very aggressive feeder, and although I tried to condition him to receiving food (freeze-thawed chicks and rodents) out of his enclosure, he still occasionally would ���pour��� ou, Published as part of Greenbaum, Eli, Allen, Kaitlin E., Vaughan, Eugene R., Pauwels, Olivier S. G., Wallach, Van, Kusamba, Chifundera, Muninga, Wandege M., Aris- Tote, Mwenebatu M., Mali, Franck M. M., Badjedjea, Gabriel, Penner, Johannes, R��del, Mark-Oliver, Rivera, Jacqueline, Sterkhova, Viktoria, Johnson, Grant, Tapondjou, Walter P. & Brown, Rafe M., 2021, Night stalkers from above: A monograph of Toxicodryas tree snakes (Squamata Colubridae) with descriptions of two new cryptic species from Central Africa, pp. 1-44 in Zootaxa 4965 (1) on pages 17-21, DOI: 10.11646/zootaxa.4965.1.1, http://zenodo.org/record/4723024, {"references":["Boulenger, G. A. (1919) Batraciens et reptiles recueillis par le Dr C. Christy au Congo Belge dans le districts de Stanleyville, Haut-Uele et Ituri en 1912 - 1914. Revue Zoologique Africaine, 7, 1 - 29. https: // doi. org / 10.5962 / bhl. part. 15108","Goodman, J. D. (1985) Two record size Blanding's tree snakes from Uganda. East African Natural History Society Bulletin, 1985, 56 - 57.","Hedges, N. G. (1983) Reptiles and Amphibians of East Africa. Kenya Literature Bureau, Nairobi, Kenya, xii + 139 pp.","Weinstein, S. A., Warrell, D. A., White, J. & Keyler, D. E. (2011) \" Venomous \" Bites from Non-Venomous Snakes: A Critical Analysis of Risk and Management of \" Colubrid \" Snake Bites. Elsevier, Amsterdam, Boston, Heidelburg, London, New York, Oxford, Paris, San Diego, San Francisco, Singapore, Sydney and Tokyo, xxvii + 336 pp.","Spawls, S. & Branch, B. (1995) The Dangerous Snakes of Africa: Natural History, Species Directory, Venoms and Snakebite. Ralph Curtis - Books, Sanibel Island, Florida, 192 pp.","Spawls, S. & Branch, B. (2020) The Dangerous Snakes of Africa. Bloomsbury, London, Oxford, New York, New Delhi and Sydney, 336 pp.","Schmidt, K. P. (1923) Contributions to the herpetology of the Belgian Congo based on the collection of the American Museum Congo Expedition, 1909 - 1915. Part II. - Snakes. Bulletin of the American Museum of Natural History, 49, 1 - 146, pls. I - XXII.","Laurent, R. F. (1956) Contribution a l'herpetologie de la region des Grands Lacs de l'Afrique centrale. I. Generalites - II. Cheloniens - III. Ophidiens. Annales du Musee Royal du Congo Belge, Serie in- 8 °, 48, 1 - 390, pls. I - XXXI.","Hellmich, W. (1957 b) Die Reptilienausbeute der hamburgischen Angola-Expedition. Mitteilungen aus dem Hamburgischen Zoologischen Museum und Institut, 55, 39 - 80.","de Witte, G. - F. (1966) Exploration du Parc National de la Garamba. Reptiles. Institut du Parcs Nationaux du Congo, 48, 1 - 108, pls. I - V.","Pitman, C. R. S. (1974) A Guide to the Snakes of Uganda, Revised Edition. Wheldon & Wesley, Ltd., Codicote, xxii + 290 pp.","Skinner, H. A. (1973) Snakes and Us: An Introduction to East African Herpetology. East African Literature Bureau, Nairobi, Kampala and Dar es Salaam, xvi + 146 pp.","de Witte, G. - F. (1975) Serpents, Boidae, Colubridae, Elapidae et Viperidae. Fondacion pour Favoriser les Recherches Scientifiques en Afrique, Serie 2, 24, 5 - 121.","Broadley, D. G., Doria, C. T. & Wigge, J. (2003) Snakes of Zambia: An Atlas and Field Guide. Edition Chimaira, Frankfurt am Main, 280 pp.","Bogert, C. M. (1940) Herpetological results of the Vernay Angola expedition. Bulletin of the American Museum of Natural History, 77, 1 - 107, pl. I.","Pitman, C. R. S. (1938) A guide to the snakes of Uganda. Part XI. (Conclusion). The Uganda Journal, 5, 160 - 244, pls. XV - XVIII.","Hughes, B. (2000) Polymorphism, sex, size and other aspects of the African snake, Boiga blandingi (Hallowell, 1844). Herpetological Bulletin, 74, 8 - 12.","Laurent, R. F. (1964) Subsidos para o estudo da biologia na Lunda, Museo do Dundo. Reptiles et amphibiens de l'Angola (Troisieme contribution). Publicacoes Culturais da Companhia de Diamantes de Angola, 67, 17 - 165.","Dowling, H. G. & Savage, J. M. (1960) A guide to the snake hemipenis: A survey of basic structure and systematic characteristics. Zoologica, 45 (Part 1), 17 - 28, pls. I - III.","Doucet, J. (1963) Les serpents de la Republique de Cote d'Ivoire. Serpents venimeux. Acta Tropica, 20 (3 - 4), 297 - 340.","Nagy, Z. T., Chifundera Kusamba, Z., Gembu Tungaluna, G. - C., Lotana Lokasola, A., Kolby, J. & Kielgast, J. (2011) Foraging acrobatics of Toxicodryas blandingii in the Democratic Republic of the Congo. Herpetology Notes, 4, 91 - 92.","Loveridge, A. (1937) Zoological results of the George Vanderbilt African expedition of 1934. Part VII - Reptiles and amphibians. Proceedings of the Academy of Natural Sciences of Philadelphia, 89, 265 - 296.","Luiselli, L., Akani, G. C. & Barieenee, I. F. (1998 a) Observations on habitat, reproduction and feeding of Boiga blandingi (Colubridae) in south-eastern Nigeria. Amphibia-Reptilia, 19 (4), 430 - 436. https: // doi. org / 10.1163 / 156853898 X 00098","Laurent, R. F. (1954) Reptiles et batraciens de la region de Dundo (Angola). Deuxieme note. Publicacoes Culturais da Compan- hia de Diamantes de Angola, 23, 35 - 84.","Laurent, R. F. (1960) Notes complementaires sur les cheloniens et les ophidiens du Congo Oriental. Annales du Musee Royal du Congo Belge, Serie in- 8 °, 84, 1 - 86.","de Witte, G. - F. (1962) Genera des serpents du Congo et du Ruanda-Urundi. Annales du Musee Royal de l'Afrique Centrale, Sciences Zoologique, Serie in Octavo, 104, i - viii + 1 - 203.","Thys van den Audenaerde, D. F. E. (1965) Les serpents des environs de Leopoldville. Revue de zoologie et de botanique africaines, 72 (3 - 4), 366 - 388.","Broadley, D. G. & Cotterill, F. P. D. (2004) The reptiles of southeast Katanga, an overlooked ' hot spot. ' African Journal of Herpetology, 53 (1), 35 - 61. https: // doi. org / 10.1080 / 21564574.2004.9635497","Boulenger, G. A. (1896) Catalogue of the Snakes in the British Museum (Natural History). Vol. III. Containing the Colubridae (Opisthoglyphae and Proteroglyphae), Amblycephalidae, and Viperidae. Trustees of the British Museum (Natural History), London, xiv + 727 pp.","Parker, H. W., Moreau, R. E. & Pakenham, R. H. W. (1940) XXV. - Notes on the herpetology of the East African Islands. Journal of Natural History, 5 (27), 309 - 314. https: // doi. org / 10.1080 / 00222934008527048","Loveridge, A. (1957) Check list of the reptiles and amphibians of East Africa (Uganda; Kenya, Tanganyika; Zanzibar). Bulletin of the Museum of Comparative Zoology, Harvard, 117 (2), 151 - 362 + i - xxxvi (index).","Wakeman, B. N. (1966) Uganda's poisonous snakes: Further observations on feeding habits. The Uganda Journal, 30, 101 - 103","Spawls, S. (1979) Sun, Sand & Snakes. William Morrow and Company, Inc., New York, New York, 254 pp.","Weinstein, S. A. & Smith, L. A. (1993) Chromatographic profiles and properties of Duvernoy's secretions from some boigine and dispholidine colubrids. Herpetologica, 49 (1), 78 - 94.","Weinstein, S. A. & Kardong, K. V. (1994) Properties of Duvernoy's secretions from opisthoglyphous and aglyphous colubrid snakes. Toxicon, 32 (10), 1161 - 1185. https: // doi. org / 10.1016 / 0041 - 0101 (94) 90347 - 6","Branch, W. R. (2005) A Photographic Guide: Snakes Other Reptiles and Amphibians of East Africa. Ralph Curtis Publishing, Inc., Sanibel Island, Florida, 132 pp.","Dashevsky, D., Debono, J., Rokyta, D., Nouwens, A., Josh, P. & Fry, B. G. (2018) Three-finger toxin diversification in the venoms of cat-eye snakes (Colubridae: Boiga). Journal of Molecular Evolution, 86, 531 - 545. https: // doi. org / 10.1007 / s 00239 - 018 - 9864 - 6"]}

Greenwood, Laura, Loader, Simon P., Lawson, Lucinda, Greenbaum, Eli, Zimkus, Breda M. (2020): A new species of Phrynobatrachus (Amphibia: Anura: Phrynobatrachidae) from the Northern Mountains of Tanzania. Journal of Natural History 54 (1-4): 63-85, DOI: 10.1080/00222933.2020.1757171, URL: http://dx.doi.org/10.1080/00222933.2020.1757171

Naja (Boulengerina) melanoleuca Hallowell, 1857 Naia haie var. melanoleuca Hallowell, 1857, Proc. Acad. Nat. Sci. Philadelphia: 61. Type locality: Gabon, syntypes ANSP 6875���76, 6878���79. Naja haje var. leucosticta Fischer, 1885, Jahr. Hamburg. Wiss. Anst. 2: 115, pl. v, fig. 11. Type locality: Cameroon and Ogoou�� River, Gabon, syntypes ZMH 4280, 7048, 7299���7302. Naia melanoleuca; Boulenger, 1896: 376 (part, vars. A [a,b] & B [a,b]) Naja melanoleuca melanoleuca; Laurent, 1956: 290, pl. xxvi, fig. 2. Naja (Boulengerina) melanoleuca ; Wallach et al., 2009. Boulengerina melanoleuca; Wallach et al., 2014: 122. Aspidelaps bocagei Sauvage, 1884: 204 (type locality: Gabon and Majumba; holotype MNHN 1884.0015) has been listed as a synonym of N. melanoleuca (e.g., Broadley, 1983; Wallach et al., 2014; Cer��aco et al., 2017), but is in fact a synonym of Naja annulata, as is evident from Sauvage���s description, which notes approximately 20 dark double bands along the entire body length, and 21 mid-dorsal scale rows (see also Schmidt, 1923). Diagnosis: DOrsum blAck, OftEN with 1���3 sEmidividEd yEllOw crOssbANds ON thE NEck, thE first mAy bE AN OcEllus; vENtEr yEllOw with 4 tO 6 blAck bANds iN thE first 100 vENtrAls, thErEAftEr uNifOrm blAck. Variation: DOrsAl scAlE rOws ON NEck 19���27, At midbOdy 19 (vEry rArEly 17 Or 21); vENtrAls 209���230; subcAudAls 59���74 (TAblE 8). SuprAlAbiAls 7, thE third ANd fOurth ENtEriNg thE Orbit; iNfrAlAbiAls 8, thE first fOur iN cONtAct with thE ANtEriOr subliNguAls, NO cuNEAtE; prEOculAr 1; pOstOculArs 3 (vEry rArEly 2 Or 4); tEmpOrAls 1+2 Or 1+3; NuchAls bOrdEriNg tEmpOrAls 5���9, usuAlly 7. Colouration: HEAd brOwN, thE suprAlAbiAls bArrEd blAck ANd yEllOw, chiN yEllOw. BlAck AbOvE, sOmEtimEs A yEllOw mONOcEllAtE mArkiNg ON thE hOOd, Or 1���3 smAll yEllOw blOtchEs, thE dOrsAl scAlEs mAy bE tippEd with whitE iN juvENilEs (Fig. 8). YEllOw Or whitE bElOw with 4 tO 6 blAck bANds ON thE first 100 vENtrAls, usuAlly uNifOrm blAck thErEAftEr. Largest recorded: 2250+420 = 2670 mm, frOm MONiyA, IbAdAN, NigEriA (ButlEr, 1982: 110). Suggested common name: CENtrAl AfricAN fOrEst cObrA. Distribution: CENtErEd ON thE CONgO BAsiN, wEst tO sOuthwEstErN NigEriA ANd pOssibly sOuthErN BENiN, sOuth tO NOrthErN ANgOlA, NOt ExtENdiNg EAst Of thE AlbErtiNE Rift VAllEy, whErE it is rEplAcEd by N. subfulva. ThE wEstErN rANgE limits ArE pOOrly uNdErstOOd. POpulAtiONs frOm sOuthEAstErN NigEriA ArE clEArly AssigNAblE tO this fOrm. A fEw spEcimENs frOm LAmtA, sOuthErN BENiN (IRD 12.B, IRD 54.B, IRD 60.B), ANd GhANA (MNHN 1983.0663���64; NO furthEr lOcAlity iNfOrmAtiON) AlsO AppEAr tO bE AssigNAblE tO N. melanoleuca., Published as part of W��ster, Wolfgang, Chirio, Laurent, Trape, Jean-Fran��ois, Ineich, Ivan, Jackson, Kate, Greenbaum, Eli, Barron, Cesar, Kusamba, Chifundera, Nagy, Zolt��n T., Storey, Richard, Hall, Cara, W��ster, Catharine E., Barlow, Axel & Broadley, Donald G., 2018, Integration of nuclear and mitochondrial gene sequences and morphology reveals unexpected diversity in the forest cobra (Naja melanoleuca) species complex in Central and West Africa (Serpentes: Elapidae) in Zootaxa 4455 (1), DOI: 10.11646/zootaxa.4455.1.3, http://zenodo.org/record/1456971, {"references":["Hallowell, E. (1857) Notes of a collection of reptiles from the Gaboon country, West Africa, recently presented to the Academy of Natural Sciences of Philadelphia, by Dr. Henry A. Ford. Proceedings of Academy of Natural Sciences, Philadelphia, 9, 48 - 72.","Fischer, J. G. (1885) Ichthyologische und herpetologische Bemerkungen. V. Herpetologische Bemerkungen. Jahrbuch der Hamburgischen Wissenschaftlichen Anstalten, 2, 82 - 121.","Boulenger, G. A. (1896) Catalogue of the Snakes in the British Museum (Natural History) III. British Museum (Natural History), London, 727 pp. https: // doi. org / 10.5962 / bhl. title. 54273","Laurent, R. F. (1956) Contribution a l'herpetologie de la region des Grands Lacs de l'Afrique centrale. I. Generalites. II. Cheloniens. III. Ophidiens. Annales du Musee royal du Congo Belge, 48, 1 - 390.","Wallach, V., Wuster, W. & Broadley, D. G. (2009) In praise of subgenera: taxonomic status of cobras of the genus Naja Laurenti (Serpentes: Elapidae). Zootaxa, 2236, 26 - 36.","Wallach, V., Williams, K. L. & Boundy, J. (2014) Snakes of the World. A Catalogue of Living and Extinct Species. CRC Press, Boca Raton, 1227 pp. https: // doi. org / 10.1201 / b 16901","Sauvage, H. E. (1884) Notice sur une collection de reptiles et de poissons recueillie a Majumba, Congo. Bulletin de la Societe Zoologique de France, 9, 199 - 204.","Broadley, D. G. (1983) FitzSimons' Snakes of Southern Africa. Delta Books, Johannesburg, 376 pp.","Ceriaco, L. M. P., Marques, M. P., Schmitz, A. & Bauer, A. M. (2017) The \" Cobra-preta \" of Sao Tome Island, Gulf of Guinea, is a new species of Naja Laurenti, 1768 (Squamata: Elapidae). Zootaxa, 4324 (1), 121 - 141. https: // doi. org / 10.11646 / zootaxa. 4324.1","Schmidt, K. P. (1923) Contributions to the herpetology of the Belgian Congo based on the collection of the American Museum Congo Expedition, 1909 - 1915. Part II. Snakes. Bulletin of the American Museum of Natural History, 49, 1 - 146.","Butler, J. A. (1982) Capture au Nigeria d'un Naja noir et blanc de grande taille et notes sur les autres Najas de l'Ouest Africain. Notes Africaines, 176, 110 - 111."]}

Naja (Boulengerina) savannula sp. nov. Broadley, Trape, Chirio & W��ster Naia melanoleuca (not Hallowell) Boulenger, 1896: 376 (part, var. D). Naja ���banded form��� Hughes, 2013: 128. Naja sp. 1 cf. melanoleuca Hallowell, 1857 (yellow banded dorsum) Trape & Bald��, 2014:318. Naja sp. 1 cf. melanoleuca (banded savanna form) Trape & Bald��, 2014: 336. Holotype: MNHN 2018.0002, an adult male collected by Laurent Chirio on 31/03/ 2007 in Ni��ni��, W Biosphere Reserve, Benin, 11.05920 ��N, 2.20488 ��E, elevation 272 m (Fig. 7). Paratype: BMNH 1975.657, a male specimen collected on the grounds of Amadou Bello University Hospital, Zaria, Nigeria, by D. A. Warrell. Diagnosis: DistiNguishAblE frOm OthEr spEciEs Of thE N. melanoleuca cOmplEx by thE prEsENcE Of 3 tO 8 sEmidividEd yEllOwish Or whitish bANds ON thE ANtEriOr dOrsAl fOrEbOdy, bEcOmiNg uNifOrm blAck cAudAd; vENtEr yEllOw with 2 tO 8 blAck bANds. GENErAlly highEr mEAN subcAudAl scAlE cOuNts thAN thE OthEr spEciEs. GENEticAlly diAgNOsAblE thrOugh pOssEssiON Of uNiquE mitOchONdriAl hAplOtypEs (cytOchrOmE b: GENBANk MH337597 ���602; ND4: MH337403 ���408) ANd uNiquE PRLR ANd UBN1 hAplOtypEs (PRLR: MH337501 ���504; UBN1: MH337532 ��� 535). Description of holotype. DimENsiONs: A mAlE spEcimEN, sNOut���vENt lENgth 1815 mm, tAil lENgth 376 mm, tOtAl lENgth 2191 mm, rAtiO tOtAl lENgth: tAil lENgth 5.8. BOdy scAlAtiON: 25 scAlE rOws ArOuNd hOOd, 19 ArOuNd midbOdy, 15 ONE hEAd lENgth AhEAd Of thE vENt, All smOOth ANd ObliquE. DOrsAl scAlEs smOOth, ObliquE. VErtEbrAl rOw NOt ENlArgEd. 213 vENtrAls (ANd ONE iNcOmplEtE vENtrAl bEfOrE thE ANAl), 69 subcAudAls, All dividEd, ANAl siNglE. HEAd scAlAtiON: 7/7 suprAlAbiAls, 3rd & 4th cONtAct Orbit, 6th lArgEst; 8/8 iNfrAlAbiAls, first 4 cONtAct ANtEriOr chiN shiElds; pOstEriOr chiN shiElds iN NArrOw cONtAct At thEir ANtEriOr ENd, sEpArAtEd by mEdiAN scAlE AlONg mOst Of thEir lENgth; 1/1 prEOculAr, twicE As lONg As high; 3/3 pOstOculArs; 1/1 ANtEriOr tEmpOrAl; 3/3 pOstEriOr tEmpOrAls; rOstrAl brOAdEr thAN high, clEArly visiblE frOm AbOvE; 7 tEmpOrAls ANd NuchAls cONtActiNg thE pAriEtAls. PAttErN: uppEr sidE Of hEAd brOwN, lips ANd vENtrAl sidE Of thE hEAd pAlE crEAmy yEllOwish with EAch lAbiAl scAlE lAtErAlly bOrdErEd with blAck. NEck brOwN, dOrsum bEcOmiNg grAduAlly blAck tOwArds rEAr. TwO vEry distiNct pAlE dOrsAl crOssbANds At thE lEvEl Of vENtrAls (V) 24 tO 28, twO OthEr At lEvEl Of V 40 ��� 45, thrEE At lEvEl Of V 58 ��� 64, thrEE lEss distiNct At lEvEl Of V 77 ��� 83, ANd thrEE slightly mArkEd At lEvEl Of V 98 ��� 104. VENtrAls 1 tO 11, 13���19, 23��� 31, 38���47 ANd 57���66 pAlE crEAmy yEllOwish, V 20���22 ANd 32 ��� 37 blAck; V 12 ANd 48���56 yEllOwish spOttEd with blAck spOts. FrOm vENtrAl 67 tO thE vENt, vENtrAls spOttEd with blAck grAduAlly bEcOmE cOmplEtEly blAck. SubcAudAls blAck. Variation: DOrsAl scAlE rOws ON NEck 19���25, At midbOdy 19, bEfOrE vENt 12���15; vENtrAls 211���233, subcAudAls 63���77 (TAblE 8). DOrsAl sEmi-dividEd yEllOw bANds 3���8; vENtrAl priNcipAl blAck bANds 2���8 (Fig. 7). Etymology: ThE NAmE is dErivEd frOm thE cONtrActiON Of its sAvANNA hAbitAt ANd ANNulAtEd cOlOur pAttErN ANd wAs cOiNEd by BArry HughEs iN AN uNpublishEd 1968 mANuscript. WE hAvE rEtAiNEd this NAmE At thE rEquEst Of Our cOllEAguE BArry HughEs. Largest recorded: 1825+405 = 2230 mm, frOm MEdiNA DjikOyE, SENEgAl (IRD 6155.S). Suggested common name: WEst AfricAN bANdEd cObrA. Distribution: SENEgAl ANd GAmbiA EAst tO NOrthErN CAmErOON (Fig. 6). Naja savannula AppEArs tO bE rEstrictEd tO gAllEry fOrEst ArEAs iN AfricA iN GuiNEAN FOrEst/SAvANNA MOsAic, ExtENdiNg NOrthwArds iNtO WEst SudANiAN SAvANNA (ChiriO, 2003, 2013; MONAstEriO et al., 2016). Our rEcOrds ArE mOstly frOm lAtitudEs 10���14��N, ExcEpt iN thE DAhOmEy GAp iN EAstErN GhANA ANd BENiN, whErE thE spEciEs ApprOAchEs thE Gulf Of GuiNEA COAst. ThE EAstErN ExtENt Of thE rANgE is pOOrly uNdErstOOd. A spEcimEN frOm MArgui WANdAlA district, NOrthErN CAmErOON (ApprOx. 10.5��N 13.6��E; MNHN 1962.0022) AppEArs tO bE AssigNAblE tO this spEciEs, ANd ONE Of us (JFT) rEcENtly cOllEctEd A spEcimEN frOm MbOurA, 20 km SW BA��bOkOum, LOgONE OriENtAl PrOviNcE, ChAd (7.598��N, 15.596��E; IRD 2281.N), ANd thErE is A rEcENt rEcOrd frOm NEAr NiAmEy, NigEr (LC, uNpublishEd dAtA). It sEEms likEly thAt thE spEciEs hAs A widEr distributiON iN NOrthErN CAmErOON, ExtrEmE sOuthErN ChAd ANd pOssibly EvEN ExtrEmE NOrthwEstErN CENtrAl AfricAN REpublic (CAR)., Published as part of W��ster, Wolfgang, Chirio, Laurent, Trape, Jean-Fran��ois, Ineich, Ivan, Jackson, Kate, Greenbaum, Eli, Barron, Cesar, Kusamba, Chifundera, Nagy, Zolt��n T., Storey, Richard, Hall, Cara, W��ster, Catharine E., Barlow, Axel & Broadley, Donald G., 2018, Integration of nuclear and mitochondrial gene sequences and morphology reveals unexpected diversity in the forest cobra (Naja melanoleuca) species complex in Central and West Africa (Serpentes: Elapidae) in Zootaxa 4455 (1), DOI: 10.11646/zootaxa.4455.1.3, http://zenodo.org/record/1456971, {"references":["Boulenger, G. A. (1896) Catalogue of the Snakes in the British Museum (Natural History) III. British Museum (Natural History), London, 727 pp. https: // doi. org / 10.5962 / bhl. title. 54273","Hughes, B. (2013) Snakes of Benin, West Africa. Bulletin de la Societe Herpetologique de France, 144, 101 - 159.","Hallowell, E. (1857) Notes of a collection of reptiles from the Gaboon country, West Africa, recently presented to the Academy of Natural Sciences of Philadelphia, by Dr. Henry A. Ford. Proceedings of Academy of Natural Sciences, Philadelphia, 9, 48 - 72.","Trape, J. - F. & Balde, C. (2014) A checklist of the snake fauna of Guinea, with taxonomic changes in the genera Philothamnus and Dipsadoboa (Colubridae) and a comparison with the snake fauna of some other West African countries. Zootaxa, 3900 (3), 301 - 338. https: // doi. org / 10.11646 / zootaxa. 3900.3.1","Chirio, L. (2003) Inventaire des reptiles de la region de la Reserve de Biosphere Transfrontaliere du W (Niger / Benin / Burkina Faso: Afrique de l'Ouest). Bulletin de la Societe Herpetologique de France, 132, 13 - 41.","Chirio, L. (2013) Inventaire des reptiles de la region de Sangaredi (Guinee maritime). Bulletin de la Societe Herpetologique de France, 144, 67 - 100."]}

Naja (Boulengerina) subfulva Laurent, 1955 Naia melanoleuca (not Hallowell); Boulenger, 1896: 376 (part, vars A [c] & E). Naja melanoleuca subfulva Laurent, 1955, Revue Zool.��� Bot. Afr. 51: 132. Type locality: Lwiro, 1850 m, Kabare, Kivu, Belgian Congo [= Democratic Republic of Congo], holotype MRAC 17514. Naja melanoleuca aurata Stucki-Stirn, 1979, Snake Report 721: 617���620. Type locality: Bamenda, Cameroon. No type designated. Naja subfulva; Chirio & Ineich, 2006: 54. Naja melanoleuca subfulva; Chirio & LeBreton, 2007: 584. Naja (Boulengerina) melanoleuca subfulva ; Broadley & Blaylock, 2013: 139. Naja (Boulengerina) melanoleuca; Conradie et al., 2016: 28. Diagnosis. MidbOdy scAlE rOws 19, ExcEpt AlONg cOAstAl rEgiONs Of EAst AfricA (KENyA, TANzANiA), whErE mOst spEcimENs hAvE 17 rOws. PAttErN highly vAriAblE. Adults Of mOst pOpulAtiONs distiNguishAblE iN hAviNg A brOwN fOrEbOdy, OftEN with spOts, gENErAlly bEcOmiNg dArkEr Or blAckish pOstEriOrly. LAbiAl pAttErN mAy bE AttENuAtEd iN mANy Adults. VENtEr with sEvErAl blAck, dArk brOwN Or grEyish crOssbANds ON thE first 50 vENtrAls, grAduAlly bEcOmiNg uNifOrm blAck cAudAd iN sOmE pOpulAtiONs, but OftEN rEmAiNiNg ENtirEly light, OftEN with ExtENsivE dArkEr spOttiNg Or spEckliNg. WhErE prEsENt, thE light fOrEbOdy ANd/Or light pOstEriOr vENtEr ArE diAgNOstic fOr this spEciEs. GENErAlly fEwEr vENtrAl bANds ANd vENtrAl scAlEs thAN N. melanoleuca Or N. savannula ANd fEwEr subcAudAls thAN N. savannula (TAblE 8). GENEticAlly diAgNOsAblE thrOugh pOssEssiON Of uNiquE mitOchONdriAl hAplOtypEs (cyt b: GENBANk MH337603 ���633; ND4: MH337409 ���439) ANd uNiquE PRLR ANd UBN1 hAplOtypEs (PRLR: MH337441 ���471; UBN1: MH337531, MH337536 ���562, MH337564 ���566). Variation. DOrsAl scAlE rOws ON NEck 19���27, At midbOdy 19 (vEry rArEly 17 Or 21); vENtrAls 196���226; ANAl ENtirE; subcAudAls 55���71 (TAblE 8). SuprAlAbiAls 7 (vEry rArEly 5 Or 6), thE third ANd fOurth ENtEriNg Orbit; iNfrAlAbiAls 8 (rArEly 7), thE first fOur (rArEly thrEE) iN cONtAct with thE ANtEriOr subliNguAls, NO cuNEAtE (vEry rArEly ONE); prEOculAr 1; pOstOculArs 3; tEmpOrAl 1+2 Or 1+3; tEmpOrAl bOrdEriNg pAriEtAls 5���9, usuAlly 7. Colouration. HEAd brOwN, thE suprAlAbiAls usuAlly bArrEd blAck ANd yEllOw, chiN yEllOw. DOrsum uNifOrm blAck, Or with fAiNt whitE crOssbArs Or whitE-tippEd dOrsAl scAlEs, rEsultiNg iN A stippliNg EffEct, iN All juvENilEs. Adults frOm thE pEriphEry Of LAkE VictOriA ANd pArts Of thE CONgO BAsiN (Fig. 9) hAvE A uNifOrmly blAck dOrsum As Adults, whErEAs iN mOst Of thE rANgE, Adults bEcOmE brOwN ANtEriOrly. ThErE mAy bE 2 Or 3 fAiNt yEllOw dividEd crOssbArs ON thE NEck. VENtrAl bANdiNg OftEN fAiNt iN Adults, with grEy Or pAlE brOwN vENtrAl bANds thAt mAy bE vEry iNcONspicuOus, Or lAckiNg AltOgEthEr. NArrOw AccEssOry vENtrAl bANds usuAlly AbsENt. Largest recorded. 2016+380 = 2396 mm (tAil truNcAtEd) [JPT 1856 ��� ZiNAvE, MOzAmbiquE], but NMZB-UM 17594 frOm INyANgANi TEA EstAtEs, ZimbAbwE, hAd A tOtAl lENgth Of 2690 mm (ONly hEAd ANd NEck prEsErvEd). Suggested common name. BrOwN fOrEst cObrA. This suggEstiON rEflEcts thE fAct thAt iN mOst pOpulAtiONs, with thE ExcEptiON Of thE surrOuNdiNgs Of LAkE VictOriA ANd sOmE lOcAtiONs iN thE CONgO BAsiN, thE ANtEriOr bOdy Of Adults is brOwN rAthEr thAN blAck. WE NOtE thAt thE distributiON Of this spEciEs iNcludEs mANy NON-fOrEstEd hAbitAts, iNcludiNg sAvANNAs ANd OpEN wEtlANds (SpAwls et al., 2018). HOwEvEr, rEtENtiON Of thE wOrd ���fOrEst��� iN thE cOmmON NAmE cONvEys thE phylOgENEtic iNfOrmAtiON thAt this spEciEs is pArt Of thE Naja melanoleuca cOmplEx. Distribution. FOrEst/sAvANNA mOsAic, ENcircliNg thE CONgO BAsiN, frOm thE grAssfiElds Of wEstErN CAmErOON NOrthwArd, rEAchiNg 13.45��N At BOl, ChAd, ON thE NOrthErN shOrE Of LAkE ChAd (IRD 2802.N), EAst thrOugh thE CAR ANd SOuth SudAN tO wEstErN EthiOpiA, sOuth thrOugh UgANdA, wEstErN KENyA, EAstErN DRC, RwANdA, BuruNdi ANd wEstErN TANzANiA tO NOrthErN MAlAwi, wEst thrOugh ZAmbiA ANd KAtANgA tO ANgOlA ANd thE LOwEr CONgO rEgiON (Fig. 6). ThE distributiON ExtENds disjuNctly EAst ANd sOuth tO thE EAst AfricAN cOAst frOm KENyA tO NOrthErN KwAZulu-NAtAl (SOuth AfricA), iNcludiNg iNlANd lOcAtiONs iN wEstErN ZimbAbwE. ThE prEcisE distributiON Of this tAxON iN thE CONgO rEgiON rEmAiNs tO bE AscErtAiNEd: wE hAvE hErE prOvidEd mOlEculAr EvidENcE Of multiplE spEcimENs Of this spEciEs frOm thE EvErgrEEN fOrEst zONE Of NOrthErN REpublic Of CONgO (LikOuAlA FOrEst ��� JAcksON et al., 2007) ANd thE NOrthEAstErN DRC (NOrthwEstErN OriENtAlE PrOviNcE AlONg thE CONgO RivEr). SOmE Of thEsE spEcimENs ANd pOpulAtiONs ArE difficult tO diffErENtiAtE frOm sympAtric N. melanoleuca, NEcEssitAtiNg thE usE Of DNA bArcOdiNg ApprOAchEs (HEbErt et al., 2003; ClAusE et al., 2016) tO ENsurE rEliAblE idENtificAtiON. TwO spEcimENs frOm sOuthEAstErN NigEriA with lOw vENtrAl scAlE cOuNts ANd lAckiNg vENtrAl bANdiNg Or displAyiNg A siNglE fAdEd bANd mAy AlsO bE AssigNAblE tO this spEciEs (DEgEmA, RivErs StAtE: BMNH 1902.11.10.9, ANd ���Oil RivEr ���: BMNH 88.8.29.20), suggEstiNg A widEr distributiON iN EAstErN NigEriA. Comment. This spEciEs displAys cONsidErAblE gEOgrAphic vAriAtiON iN pAttErN ANd scAlAtiON, As wEll As iNdicAtiONs Of phylOgEOgrAphic structurE. ThE prEsENt study fOcusEd ON thE rElAtiONships AmONg thE fivE mitOchONdriAlly dEfiNEd cANdidAtE spEciEs, ANd did NOt ENcOmpAss thE full rANgE Of mOrphOlOgicAl vAriAtiON iN N. subfulva. It is pOssiblE thAt futurE wOrk mAy idENtify AdditiONAl cryptic divErsity withiN this widE-rANgiNg ANd vAriAblE spEciEs., Published as part of W��ster, Wolfgang, Chirio, Laurent, Trape, Jean-Fran��ois, Ineich, Ivan, Jackson, Kate, Greenbaum, Eli, Barron, Cesar, Kusamba, Chifundera, Nagy, Zolt��n T., Storey, Richard, Hall, Cara, W��ster, Catharine E., Barlow, Axel & Broadley, Donald G., 2018, Integration of nuclear and mitochondrial gene sequences and morphology reveals unexpected diversity in the forest cobra (Naja melanoleuca) species complex in Central and West Africa (Serpentes: Elapidae) in Zootaxa 4455 (1), DOI: 10.11646/zootaxa.4455.1.3, http://zenodo.org/record/1456971, {"references":["Laurent, R. F. (1955) Diagnoses preliminaires de quelques serpents venimeux. Revue de Zoologie et Botanique Africaine, 51, 127 - 139.","Boulenger, G. A. (1896) Catalogue of the Snakes in the British Museum (Natural History) III. British Museum (Natural History), London, 727 pp. https: // doi. org / 10.5962 / bhl. title. 54273","Stucki-Stirn, M. C. (1979) Snake Report 721. Herpeto-Verlag, Teuffenthal, 650 pp.","Chirio, L. & Ineich, I. (2006) Biogeography of the reptiles of the Central African Republic. African Journal of Herpetology, 55, 23 - 59. https: // doi. org / 10.1080 / 21564574.2006.9635538","Chirio, L. & LeBreton, M. (2007) Atlas des reptiles du Cameroun. Museum National d'Histoire Naturelle, Paris, 686 pp.","Broadley, D. G. & Blaylock, R. (2013) The Snakes of Zimbabwe and Botswana. Edition Chimaira, Frankfurt am Main, 387 pp.","Conradie, W., Bills, R. & Branch, W. R. (2016) The herpetofauna of the Cubango, Cuito, and lower Cuando river catchments of south-eastern Angola. Amphibian & Reptile Conservation, 10, 6 - 36."]}

Naja (Boulengerina) guineensis sp. nov. Broadley, Trape, Chirio, Ineich & W��ster Naia melanoleuca (not Hallowell) Boulenger, 1896: 376 (part, var. B [c,d], C). Naja sp. 2 cf. melanoleuca Hallowell, 1857 (blackish dorsum) Trape & Bald��, 2014: 318. Naja sp. 2 cf. melanoleuca (forest form). Trape & Bald��, 2014: 336. Holotype: MNHN 1921.0485, a male from N���Z��b��la, Macenta Prefecture, Nz��r��kor�� region of forested southeastern Guinea (8�� 05���N, 9�� 05���W), elev. 490 m, Coll. Paul Chabanaud (1876���1959) between 1919���1920 (Chabanaud, 1921: 471) (Fig. 5). Paratype: BMNH 1960.1.3.72, a male from Njala, Kori, Sierra Leone, Coll. C.T. Pyne. Diagnosis. Naja guineensis cAN bE distiNguishEd frOm thE pArtly sympAtric N. savannula sp. NOv. by lAckiNg ExtENdEd dOrsAl bANdiNg, OftEN hAviNg 17 rAthEr thAN 19 dOrsAl scAlE rOws At midbOdy, A gENErAlly lOwEr subcAudAl scAlE cOuNt, fEwEr vENtrAl bANds, A lEssEr pOstEriOr ExtENt Of thE vENtrAl bANdiNg, ANd A strONg tENdENcy tOwArds mElANism iN Adults. SpEcimENs with 19 midbOdy dOrsAl scAlE rOws cAN bE distiNguishEd frOm N. melanoleuca thrOugh thE rEducEd NumbEr Of vENtrAl bANds, lEssEr pOstEriOr ExtENt Of bANdiNg ANd tENdENcy Of ONtOgENEtic mElANism frOm N. subfulva iN lAckiNg A lightEr ANtEriOr dOrsum ANd thrOugh ONtOgENEtic mElANism, ANd frOm N. peroescobari iN hAviNg thE pOstEriOr chiN shiElds iN cONtAct. Description of holotype. DimENsiONs: A mAlE spEcimEN with dissEctEd tAil bAsE, sNOut���vENt lENgth 1850 mm, tAil lENgth 370 mm, tOtAl lENgth 2220 mm, rAtiO tOtAl lENgth: tAil lENgth 6:1. BOdy scAlAtiON: 23 scAlE rOws ArOuNd hOOd, 17 ArOuNd midbOdy, 13 ONE hEAd lENgth AhEAd Of thE vENt, All smOOth ANd ObliquE; 208 vENtrAls, 62 subcAudAls, All dividEd, ANAl siNglE. HEAd scAlAtiON: 7/7 suprAlAbiAls, 3rd & 4th cONtAct Orbit, 6th lArgEst; 8/8 iNfrAlAbiAls, first 4 cONtAct ANtEriOr chiN shiElds; ANtEriOr pAir Of chiN shiElds ANd iNfrAlAbiAls 3���4 with AN ANOmAly ON bOth sidEs, thEy ArE wArty ANd cOvErEd with circumvOlutiONs; 1/1 prEOculAr, twicE As lONg As high; 3/3 pOstOculArs; 1/1 ANtEriOr tEmpOrAl; pOstEriOr tEmpOrAls 3/3; rOstrAl brOAdEr thAN high ANd slightly dAmAgEd ANtEriOrly, clEArly visiblE frOm AbOvE; 7 tEmpOrAls ANd NuchAls cONtActiNg bOth pAriEtAls. PAttErN: uppEr sidE Of hEAd ANd uppEr tEmpOrAl rEgiON dArk brOwN iNcludiNg pAriEtAls ANd tEmpOrAls but blAck bEhiNd thEm, lips ANd vENtrAl sidE Of hEAd crEAmy yEllOwish withOut dArk mOttliNg ON chiN ANd ANtEriOr thrOAt; EAch pOstEriOr suprAlAbiAl EdgE brOAdly OutliNEd iN blAck frOm EyE lEvEl tO mOuth cOrNEr; tEmpOrAls uNifOrmly brOwNish. DOrsum uNifOrm brOwNish-blAck thrOughOut. VENtEr: ANtEriOr first 32 cm (14% Of tOtAl lENgth) Of vENtEr with AltErNAtiNg yEllOw ANd blAckish irrEgulAr bANds, thE pOstEriOr yEllOw bANds iNcrEAsiNgly suffusEd with dArk mOttliNg. ONly vENtrAl 1 is uNifOrm clEAr yEllOw, vENtrAls 2���10, 19, 28���29, 32���35 yEllOw but AlwAys iNcrEAsiNgly suffusEd with dArk mOttliNg, rEmAiNiNg vENtrAls ANd uNdErsidE Of tAil blAck (AftEr vENtrAl 35). Variation. MidbOdy dOrsAl scAlE rOw cOuNts Of 17 ANd 19 ArE ApprOximAtEly EquAlly cOmmON iN this spEciEs. IN lArgE Adults, light pAttErN ElEmENts ON thE hEAd ANd thrOAt OftEN bEcOmE hEAvily suffusEd with blAck pigmENt, lEAdiNg tO A virtuAlly ENtirEly mElANistic sNAkE. SOmE spEcimENs hAvE 1���4 gENErAlly fAiNt Or pOOrly dEfiNEd light bANds AcrOss thE NEck, ANd OccAsiONAlly AN OcEllAtE hOOd mArkiNg. Largest recorded: 1818+437 = 2255 mm, frOm BAllAssOu, GuiNEA (IRD 4213.G), but lArgEr spEcimENs hAvE bEEN ObsErvEd. BAsEd ON lOcAlity, MENziEs��� (1966) rEpOrt Of A spEcimEN mEAsuriNg ���Eight fEEt, Eight iNchEs��� (264 cm) frOm BO, SiErrA LEONE, is likEly tO rEfEr tO this spEciEs. Etymology. ThE spEcific EpithEt guineensis mEANs ���frOm GuiNEA ��� ANd is chOsEN tO rEflEct thE distributiON Of thE spEciEs iN thE UppEr GuiNEA fOrEsts Of WEst AfricA, pArt Of thE WEst AfricAN FOrEsts biOdivErsity hOtspOt (MyErs et al., 2000). Suggested common name. BlAck fOrEst cObrA. Distribution. ThE distributiON Of Naja guineensis AppEArs tO bE rEstrictEd tO thE UppEr GuiNEA FOrEsts Of wEstErN AfricA, frOm wEstErN TOgO tO LibEriA ANd GuiNEA (TrApE & BAld��, 2014) (Fig. 6). ThErE is A siNglE rEcOrd frOm CONtubOEl, GuiNEA BissAu (MBL 535). All OthEr rEcOrds 10��N Or lOwEr., Published as part of W��ster, Wolfgang, Chirio, Laurent, Trape, Jean-Fran��ois, Ineich, Ivan, Jackson, Kate, Greenbaum, Eli, Barron, Cesar, Kusamba, Chifundera, Nagy, Zolt��n T., Storey, Richard, Hall, Cara, W��ster, Catharine E., Barlow, Axel & Broadley, Donald G., 2018, Integration of nuclear and mitochondrial gene sequences and morphology reveals unexpected diversity in the forest cobra (Naja melanoleuca) species complex in Central and West Africa (Serpentes: Elapidae) in Zootaxa 4455 (1), DOI: 10.11646/zootaxa.4455.1.3, http://zenodo.org/record/1456971, {"references":["Boulenger, G. A. (1896) Catalogue of the Snakes in the British Museum (Natural History) III. British Museum (Natural History), London, 727 pp. https: // doi. org / 10.5962 / bhl. title. 54273","Hallowell, E. (1857) Notes of a collection of reptiles from the Gaboon country, West Africa, recently presented to the Academy of Natural Sciences of Philadelphia, by Dr. Henry A. Ford. Proceedings of Academy of Natural Sciences, Philadelphia, 9, 48 - 72.","Trape, J. - F. & Balde, C. (2014) A checklist of the snake fauna of Guinea, with taxonomic changes in the genera Philothamnus and Dipsadoboa (Colubridae) and a comparison with the snake fauna of some other West African countries. Zootaxa, 3900 (3), 301 - 338. https: // doi. org / 10.11646 / zootaxa. 3900.3.1"]}

Naja (Boulengerina) savannula sp. nov. Broadley, Trape, Chirio & Wüster Naia melanoleuca (not Hallowell) Boulenger, 1896: 376 (part, var. D). Naja “banded form” Hughes, 2013: 128. Naja sp. 1 cf. melanoleuca Hallowell, 1857 (yellow banded dorsum) Trape & Baldé, 2014:318. Naja sp. 1 cf. melanoleuca (banded savanna form) Trape & Baldé, 2014: 336. Holotype: MNHN 2018.0002, an adult male collected by Laurent Chirio on 31/03/ 2007 in Niénié, W Biosphere Reserve, Benin, 11.05920 °N, 2.20488 °E, elevation 272 m (Fig. 7). Paratype: BMNH 1975.657, a male specimen collected on the grounds of Amadou Bello University Hospital, Zaria, Nigeria, by D. A. Warrell. Diagnosis: DistiNguishAblE frOm OthEr spEciEs Of thE N. melanoleuca cOmplEx by thE prEsENcE Of 3 tO 8 sEmidividEd yEllOwish Or whitish bANds ON thE ANtEriOr dOrsAl fOrEbOdy, bEcOmiNg uNifOrm blAck cAudAd; vENtEr yEllOw with 2 tO 8 blAck bANds. GENErAlly highEr mEAN subcAudAl scAlE cOuNts thAN thE OthEr spEciEs. GENEticAlly diAgNOsAblE thrOugh pOssEssiON Of uNiquE mitOchONdriAl hAplOtypEs (cytOchrOmE b: GENBANk MH337597 –602; ND4: MH337403 –408) ANd uNiquE PRLR ANd UBN1 hAplOtypEs (PRLR: MH337501 –504; UBN1: MH337532 – 535). Description of holotype. DimENsiONs: A mAlE spEcimEN, sNOut–vENt lENgth 1815 mm, tAil lENgth 376 mm, tOtAl lENgth 2191 mm, rAtiO tOtAl lENgth: tAil lENgth 5.8. BOdy scAlAtiON: 25 scAlE rOws ArOuNd hOOd, 19 ArOuNd midbOdy, 15 ONE hEAd lENgth AhEAd Of thE vENt, All smOOth ANd ObliquE. DOrsAl scAlEs smOOth, ObliquE. VErtEbrAl rOw NOt ENlArgEd. 213 vENtrAls (ANd ONE iNcOmplEtE vENtrAl bEfOrE thE ANAl), 69 subcAudAls, All dividEd, ANAl siNglE. HEAd scAlAtiON: 7/7 suprAlAbiAls, 3rd & 4th cONtAct Orbit, 6th lArgEst; 8/8 iNfrAlAbiAls, first 4 cONtAct ANtEriOr chiN shiElds; pOstEriOr chiN shiElds iN NArrOw cONtAct At thEir ANtEriOr ENd, sEpArAtEd by mEdiAN scAlE AlONg mOst Of thEir lENgth; 1/1 prEOculAr, twicE As lONg As high; 3/3 pOstOculArs; 1/1 ANtEriOr tEmpOrAl; 3/3 pOstEriOr tEmpOrAls; rOstrAl brOAdEr thAN high, clEArly visiblE frOm AbOvE; 7 tEmpOrAls ANd NuchAls cONtActiNg thE pAriEtAls. PAttErN: uppEr sidE Of hEAd brOwN, lips ANd vENtrAl sidE Of thE hEAd pAlE crEAmy yEllOwish with EAch lAbiAl scAlE lAtErAlly bOrdErEd with blAck. NEck brOwN, dOrsum bEcOmiNg grAduAlly blAck tOwArds rEAr. TwO vEry distiNct pAlE dOrsAl crOssbANds At thE lEvEl Of vENtrAls (V) 24 tO 28, twO OthEr At lEvEl Of V 40 – 45, thrEE At lEvEl Of V 58 – 64, thrEE lEss distiNct At lEvEl Of V 77 – 83, ANd thrEE slightly mArkEd At lEvEl Of V 98 – 104. VENtrAls 1 tO 11, 13–19, 23– 31, 38–47 ANd 57–66 pAlE crEAmy yEllOwish, V 20–22 ANd 32 – 37 blAck; V 12 ANd 48–56 yEllOwish spOttEd with blAck spOts. FrOm vENtrAl 67 tO thE vENt, vENtrAls spOttEd with blAck grAduAlly bEcOmE cOmplEtEly blAck. SubcAudAls blAck. Variation: DOrsAl scAlE rOws ON NEck 19–25, At midbOdy 19, bEfOrE vENt 12–15; vENtrAls 211–233, subcAudAls 63–77 (TAblE 8). DOrsAl sEmi-dividEd yEllOw bANds 3–8; vENtrAl priNcipAl blAck bANds 2–8 (Fig. 7). Etymology: ThE NAmE is dErivEd frOm thE cONtrActiON Of its sAvANNA hAbitAt ANd ANNulAtEd cOlOur pAttErN ANd wAs cOiNEd by BArry HughEs iN AN uNpublishEd 1968 mANuscript. WE hAvE rEtAiNEd this NAmE At thE rEquEst Of Our cOllEAguE BArry HughEs. Largest recorded: 1825+405 = 2230 mm, frOm MEdiNA DjikOyE, SENEgAl (IRD 6155.S). Suggested common name: WEst AfricAN bANdEd cObrA. Distribution: SENEgAl ANd GAmbiA EAst tO NOrthErN CAmErOON (Fig. 6). Naja savannula AppEArs tO bE rEstrictEd tO gAllEry fOrEst ArEAs iN AfricA iN GuiNEAN FOrEst/SAvANNA MOsAic, ExtENdiNg NOrthwArds iNtO WEst SudANiAN SAvANNA (ChiriO, 2003, 2013; MONAstEriO et al., 2016). Our rEcOrds ArE mOstly frOm lAtitudEs 10–14°N, ExcEpt iN thE DAhOmEy GAp iN EAstErN GhANA ANd BENiN, whErE thE spEciEs ApprOAchEs thE Gulf Of GuiNEA COAst. ThE EAstErN ExtENt Of thE rANgE is pOOrly uNdErstOOd. A spEcimEN frOm MArgui WANdAlA district, NOrthErN CAmErOON (ApprOx. 10.5°N 13.6°E; MNHN 1962.0022) AppEArs tO bE AssigNAblE tO this spEciEs, ANd ONE Of us (JFT) rEcENtly cOllEctEd A spEcimEN frOm MbOurA, 20 km SW BAïbOkOum, LOgONE OriENtAl PrOviNcE, ChAd (7.598°N, 15.596°E; IRD 2281.N), ANd thErE is A rEcENt rEcOrd frOm NEAr NiAmEy, NigEr (LC, uNpublishEd dAtA). It sEEms likEly thAt thE spEciEs hAs A widEr distributiON iN NOrthErN CAmErOON, ExtrEmE sOuthErN ChAd ANd pOssibly EvEN ExtrEmE NOrthwEstErN CENtrAl AfricAN REpublic (CAR).

Wagner, Philipp, Greenbaum, Eli, Bauer, Aaron M., Kusamba, Chifundera, Leaché, Adam D. (2018): Lifting the blue-headed veil - integrative taxonomy of the Acanthocercus atricollis species complex (Squamata: Agamidae). Journal of Natural History 52 (13-16): 771-817, DOI: 10.1080/00222933.2018.1435833, URL: http://dx.doi.org/10.1080/00222933.2018.1435833

Acanthocercus minutus (Klausewitz, 1957: 170) (Figure 17) 1957 Agama atricollis minuta Klausewitz, Eidonomische Untersuchungen ��ber die Rassenkreise Agama cyanogaster und Agama atricollis. 2. Die Unterarten von Agama atricollis. Senckenbergiana biologica 38: 157���174. Holotype ZMB 29089, from ��� Dscheffedenza [=Ciaffedenza], Shoa���, Ethiopia (for comment see distribution). Description A small species of the genus with a total length below 290 mm (SVL: 93���117 mm, x = 108.2 mm, n = 3), tail only one fourth longer than SVL, shorter than in A. ugandaensis or A. kiwuensis. Both Loveridge (1920) and Klausewitz (1957) noted the small size of this species. The head seems to be only moderately broad because of the small size, but in relation to SVL length it is broader than in other species of the genus. Ear openings as large as eyes, with the tympanum visible. Occipital scale lacking. Nostril slightly below the canthus rostralis. No vertebral crest. Scales arranged in 104���114 (x = 109.7, n = 3) rows around midbody and 72���85 (x = 79.7, n = 3) longitudinal rows along the vertebrae. Matrix scales small, rhomboidal and keeled, scattered with enlarged scales. These larger scales are not as large and distinct as in A. gregorii. Vertebral scales keeled, enlarged scales intermixed with matrix scales but distinct to the lateral scalation, not arranged in distinct vertebral bands or framed by scales which are larger than the scales of the centre of the band. Gular scales small, smooth and flat. Ventral scales small and smooth, in 71���75 (x = 73.7, n = 3) longitudinal rows. Males usually with two (sometimes three) continuous rows of 14���17 precloacal pores, which are lacking in females. 1 = without locality [03.21��N, 28.52��E]; 2 = Okapi Faunal Reserve, Kayumaga stream; 3 = Avakubi, Ituri Forest; 4 = Epulu; 5 = Okapi Faunal Reserve; 6 = without locality [1.12��N, 29.25��E]; 7 = Bunia; 8 = Ndjuma Forest; 9 = Kisanzi [village near Mt. Teye]; 10 = Hakitengya; 11 = Byumba; 12 = Fort Portal; 13 = Semliki Forest; 14 = Nyakalengijo; 15 = Kyehundwe; 16 = Jambo Village; 17 = Kasese; 18 = Mount Ruwenzori; 19 = Butembo; 20 = Lake Edward; 21 = Kazinga channel; 22 = Kichwamba; 23 = Ibanda; 24 = Kabasha Escarpment; 25 = Kasese; 26 = Nyakabande; 27 = near Goma; 28 = Gisenyi [Rutshuru]; 29 = Buyumba; 30 = Isale; 31 = Upper Mulinga River, Idjwi Island; 32 = Kibuye; 33 = Bukavu; 34 = Cyamudongo; 35 = Bujumbura; 36 = Kikonda; 37 = Budongo Forest; 38 = Masinde; 39 = Kabulamuliro; 40 = Entebbe; 41 = Kampala; 42 = Mabira Forest; 43 = Jinja; 44 = Ntotoro; 45 = Osukuru; 46 = Elgonyi; 47 = Kitale; 48 = Kisere; 49 = Kamwega; 50 = Kakamega Forest; 51 = Buyangu Village; 52 = Kaimosi; 53 = Siaya; 54 = Ngiya; 55 = Kisumu; 56 = Lumbwa; 57 = Migori; 58 = Loita Plains; 59 = Nakuru; 60 = Nakuru National Park; 61 = Lake Naivasha; 62 = Thika; 63 = Mount Meru; 64 = Kitosh; 65 = without locality [1.52��N, 28.59��E]. There is no information about life coloration available. According to Klausewitz (1957) this species, like A. ugandaensis, is uniform brownish with a reticulated blue pattern on the throat. However, this description seems to be based on preserved specimens. Distribution A. minutus is mainly restricted to Ethiopia with few records from Kenya (e.g. adult female from Nairobi mentioned by Klausewitz 1957, Figure 14). The published type locality is Dscheffedenza in ��� Shoa���, which seems to be the German name for ��� Ciaffedenza���, a town east of Addis Ababa. Habitat and ecology Little information about habitat or ecology is available. Within a species account on A. atricollis, Largen and Spawls (2010) mentioned Ethiopian populations of small, frequently less colourful and ground-living specimens which probably refer to A. minutus. These populations are distributed within an altitudinal range of 1500���3000 m asl and inhabit dry savannah and woodland habitats. Largen and Spawls (2006, 2010) mentioned that they do not show affinity for trees and that the specimens were most frequently encountered in open environments in holes or beneath rocks. However, they only mentioned that there was uncertainty about which name to apply to these populations, but did not give citations or mention particular taxon names., Published as part of Wagner, Philipp, Greenbaum, Eli, Bauer, Aaron M., Kusamba, Chifundera & Leach��, Adam D., 2018, Lifting the blue-headed veil - integrative taxonomy of the Acanthocercus atricollis species complex (Squamata: Agamidae), pp. 771-817 in Journal of Natural History 52 (13 - 16) on pages 807-810, DOI: 10.1080/00222933.2018.1435833, http://zenodo.org/record/5174451, {"references":["Klausewitz W. 1957. Eidonomische Untersuchungen uber die Rassenkreise Agama cyanogaster und Agama atricollis. Die Unterarten von Agama atricollis [Eidonomic studies about the species complex of Agama cyanogaster and Agama atricollis. The subspecies of Agama atricollis]. Senckenb Biol. 38: 157 - 174.","Loveridge A. 1920. Notes on East African lizards collected 1915 - 1919, with the description of a new genus and species of skink and new subspecies of gecko. Proc Zool Soc London. 1920: 131 - 167.","Largen MJ, Spawls S. 2010. Amphibians and reptiles of ethiopia and eritrea. Frankfurt: Edition Chimaira; p. 694.","Largen MJ, Spawls S. 2006. Lizards of Ethiopia (Reptilia Sauria): an annotated checklist, bibliography, gazetteer and identification. Trop Zool. 19: 21 - 109."]}

Acanthocercus ugandaensis (Klausewitz, 1957: 169) (Figure 18) 1957 Agama atricollis ugandaensis Klausewitz, Eidonomische Untersuchungen über die Rassenkreise Agama cyanogaster und Agama atricollis. 2. Die Unterarten von Agama atricollis. Senckenbergiana biologica 38: 157–174. Holotype ZMB 11904, from ‘ Uganda’, East Africa. Description A medium sized species with a total length up to 350 mm in adult males (SVL: 90–124, x = 106 mm, n = 16), tail long, about 1.6 times longer than the SVL (fide Klausewitz 1957), longer than in A. gregorii and A. kiwuensis. Head broad (particularly in males) lacking the occipital scale. Ear openings slightly larger than eyes, with the tympanum visible. Nostril slightly below the canthus rostralis. No or very weakly developed nuchal crest. Scales arranged in 75–110 (x = 88.4, n = 16) rows around midbody and 54–69 (x = 60.2, n = 16) longitudinal rows along the vertebrate. Dorsal body scalation consists of small, smooth to feebly keeled, rhomboidal matrix scales, scattered with enlarged keeled scales. Vertebral scales strongly keeled, enlarged, usually larger than other enlarged body scales, forming a distinct vertebral band framed by scales which are larger than the scales of the centre of the band. Ventral scales small and smooth (sometimes very feebly keeled on the margins of the belly), in 56–80 (x = 65.7, n = 16) longitudinal rows. Gular scales small and keeled. Males usually with two (sometimes three) continuous rows of 12–17 precloacal pores, which are poorly developed in females. According to Klausewitz (1957) the coloration is brownish, without any pale or yellowish pattern, but this seems to be a description of preserved specimens. In nuptial males, the head, forelimbs and distal half of the tail are brilliant blue. Hind limbs and first half of the tail yellowish-greenish. On the upper surface of neck and body, the matrix scales are dark brown to black, with the enlarged scales yellow and blue on the flanks and blue along the vertebrate. Sometimes also some yellow spots on the head and the forelimbs. Females light brown, with a dark green to brown dorsolateral stripe on each side and a pale vertebral stripe. Distribution Acanthocercus ugandaensis was described from unspecified localities in Uganda, and since, the description has not been recorded elsewhere. Herein, we recognize it from Rwanda (Cyamudongo), Kenya (Kakamega, Nairobi) and Tanzania (Mount Meru) as well (Figure 15). Habitat and ecology Acanthocercus ugandaensis is a tree dweller and occurs in secondary dry forest, but also in plantations, large hedges along streets and on solitary trees in gardens (pers. observ. E. Greenbaum). There is no information about the social structure of the species, but solitary specimens only were trapped by the senior author around Kakamega forest, Kenya., Published as part of Wagner, Philipp, Greenbaum, Eli, Bauer, Aaron M., Kusamba, Chifundera & Leaché, Adam D., 2018, Lifting the blue-headed veil - integrative taxonomy of the Acanthocercus atricollis species complex (Squamata: Agamidae), pp. 771-817 in Journal of Natural History 52 (13 - 16) on pages 810-812, DOI: 10.1080/00222933.2018.1435833, http://zenodo.org/record/5174451, {"references":["Klausewitz W. 1957. Eidonomische Untersuchungen uber die Rassenkreise Agama cyanogaster und Agama atricollis. Die Unterarten von Agama atricollis [Eidonomic studies about the species complex of Agama cyanogaster and Agama atricollis. The subspecies of Agama atricollis]. Senckenb Biol. 38: 157 - 174."]}

Greenbaum, Eli, Sinsch, Ulrich, Lehr, Edgar, Valdez, Federico, Kusamba, Chifundera (2013): Phylogeography of the reed frog Hyperolius castaneus (Anura: Hyperoliidae) from the Albertine Rift of Central Africa: Implications for taxonomy, biogeography and conservation. Zootaxa 3731 (4): 473-494, DOI: http://dx.doi.org/10.11646/zootaxa.3731.4.3

Gekko petricolus group (Figure 6 A ���C) Species. G. badenii Nekrasova & Szczerbak, 1993 (Figure 6 B); G. canaensis Ngo & Gamble, 2011; G. grossmanni G��nther, 1994 (Figure 6 C); G. lauhachindaei Panitvong, Sumontha, Konlek & Kunya, 2010; G. petricolus Taylor, 1962 (Figure 6 A); G. russelltraini Ngo, Bauer, Wood & Grismer, 2009; G. takouensis Ngo & Gamble, 2010. Characters. 82.9���108.5 mm SVL; nares in contact with rostral; nasals 3; postmentals relatively large; dorsal tubercle rows 8���18; precloacal pores 8���15; postcloacal tubercles 1���3; no webbing between fingers and toes; fore and hind limbs without tubercles (but present on hind limbs of G. petricolus); lateral folds without tubercles; subcaudals enlarged, in a longitudinal row; head without pattern or with blotches or short stripes, but not forming a distinctive UU- or W-shaped pattern; back banded (G. badenii) or more or less symmetrically blotched (G. canaensis, G. grossmanni, G. lauhachindaei, G. petricolus, G. russelltraini, G. takouensis). Distribution. Laos, Tailand and Vietnam (Figure 3 A). Remarks. Based on morphological characters, G. russelltraini and G. takouensis are probable sister species, and both are closely related to G. canaensis and G. lauhachindaei. Futhermore G. grossmanni and G. petricolus are likely sister species. G. grossmanni has a similar pattern compared with G. p e t r i c o l u s, but the latter species is the sole member of the Gekko petricolus group that has tubercles on the hind limbs. G. badenii differs from the aforementiond taxa from southern Vietnam and Thailand in having a banded dorsal pattern without flecks between bands., Published as part of R��sler, Herbert, Bauer, Aaron M., Heinicke, Matthew P., Greenbaum, Eli, Jackman, Todd, Nguyen, Truong Quang & Ziegler, Thomas, 2011, Phylogeny, taxonomy, and zoogeography of the genus Gekko Laurenti, 1768 with the revalidation of G. reevesii Gray, 1831 (Sauria: Gekkonidae), pp. 1-50 in Zootaxa 2989 on page 18, DOI: 10.5281/zenodo.278393, {"references":["Nekrasova, O. D. & Szczerbak, N. N. (1993) The geckos from south of Vietnam. In: Programm and Abstracts of the 7 th Ordinary General Meeting Societas Europaea Herpetologica, Barcelona, 1993, 104.","Ngo, V. T. & Gamble, T. (2011) Gekko canaensis sp. nov. (Squamata: Gekkonidae), a new gecko from Southern Vietnam. Zootaxa, 2890, 53 - 64.","Gunther, R. (1994) Eine neue Art der Gattung Gekko (Reptilia, Squamata, Gekkonidae) aus dem Suden Vietnams. Zoologischer Anzeiger, 233, 57 - 67.","Panitvong, N., Sumontha, M., Konlek, K. & Kunya, K. (2010) Gekko lauhachindaei, a new cave-dwelling gecko (Reptilia: Gekkonidae) from Central Thailand. Zootaxa, 2671, 40 - 52.","Taylor, E. H. (1962) New Oriental reptiles. The University of Kansas Science Bulletin, Lawrence, 43, 209 - 263.","Ngo, V. T., Bauer, A. M., Wood, P. L. & Grismer, J. L. (2009) A new species of Gekko Laurenti, 1768 (Squamata: Gekkonidae) from Dong Nai Province, Southeastern Vietnam. Zootaxa, 2238, 33 - 42.","Ngo, V. T. & Gamble, T. (2010) A new species of Gekko (Squamata: Gekkonidae) from Ta Kou Nature Reserve, Binh Thuan Province, Southern Vietnam. Zootaxa, 2346, 17 - 28."]}

Gekko gecko group (Figure 2) Species. G. albofasciolatus G��nther, 1867; G. gecko (Linnaeus, 1758) (Figure 2 A); G. nutaphandi Bauer, Sumontha & Pauwels, 2008; G. siamensis Grossmann & Ulber, 1990 (Figure 2 B); G. smithii Gray, 1842 (Figure 2 C); G. verreauxi Tytler, 1864 (Figure 2 D); Gekko sp. ���Tioman���; Gekko sp. ���Sulawesi���; Gekko sp. ���Togian.��� Characters. 150.0���191.0 mm SVL; nares, except for G. verreauxi, not in contact with rostral; nasals 3���6; postmentals relatively slow (largest in G. siamensis), dorsal tubercle rows 10���19; precloacal pores 10���16; postcloacal tubercles 2���4 (rarely single); webbing between fingers and toes lacking; tubercles present on fore and hind limbs; lateral fold without tubercles; subcaudals enlarged, in two parallel rows; iris yellow, green, blue or brick red; Yshaped head pattern usually discernible; light (white), more or less transversally arranged, symmetrical dorsal and lateral blotches. Distribution. From India and Nepal to China, southwards to Indonesia (Indoaustralian Archipelago) (Figure 3 B); G. albofasciolatus is endemic to Borneo; G. verreauxi is endemic to the Andaman Islands; Gekko sp. "Tioman" is endemic to Tioman Island; Gekko sp. "Sulawesi" is endemic to Sulawesi; Gekko sp. "Togian" is endemic to Togian and Batudaka. Allochthonous populations of Gekko gecko exist in the Caribbean, in Belize, on Hawaii and in Florida (Kraus 2009). Remarks. In the original description of Gecko albo-fasciolatus (= Gekko albofasciolatus), G��nther (1867) only provided a vague type locality Polynesia with a question mark. Smith (1935) subsequently stated ��� type loc. unknown, probably Malay Archipelago.��� Later, Wermuth (1965) wrote ���Polynesia��� as the type locality with the addition (fide Smith 1935), that the species probably occurs in the Indo-Australian Archipelago. According to G��nther (1872), G. s m i t h i i occurs in the north of Borneo (Labuan, now Sabah) and G. albofasciolatus in the South (Banjermassin, Martapoura). We herein restrict the type locality of G. albofasciolatus to Banjermassin (= Banjarmasin), Kalimantan, Indonesia. While commenting on G. albofasciolatus, G��nther (1872: 589) stated that he received three specimens from Dr. Bleeker from Borneo under different names (Platydactylus pentonopus, Platydactylus borneensis, and Hemidactylus zosterophorus). From G��nther���s (1872) footnotes it is obvious that he allocated all of Bleeker���s specimens to G. albofasciolatus. Of Bleeker���s species, only Platydactylus borneensis had been recorded from Borneo. The names P. borne��nsis (= P. borneensis) and H. zosterophorus were introduced by Bleeker (1857) without formal species descriptions (see also Bleeker 1860). The type loaclity of Platydactylus borneensis (non Pentadactylus borneensis G��nther, 1864 = Aeluroscalabotes felinus G��nther, 1864; non Tarentola borneensis Gray, 1845 = Tarentola delalandii Dum��ril & Bibron, 1836) is ���Bandjermasin��� and the type locality of Hemidactylus zosterophorus is ���Padang (ook op Nias)���. In his critical review of Bleeker���s type specimens, Boulenger (1887) did not consider the taxa Platydactylus borneensis and Hemidactylus zosterophorus. According to Bauer (1994), both names are species inquirenda and Kluge (2001) listed them as nomina nuda (see also R��sler 2000). Therefore, the names Platydactylus borneensis Bleeker, 1857 and Hemidactylus zosterophorus Bleeker, 1857, which are according to G��nther���s (1872) statements younger, subjective synonyms of Gekko albofasciolatus G��nther, 1867, are not available. The same concerns G��nther���s (1872) name Platydactylus pentonopus. Boulenger (1885) synonymized G. albofasciolatus with G. s m i t h i i (see also Wermuth 1965; Kluge 1991, 1993; Bauer 1994; R��sler 2000). De Rooij (1915) obviously also followed Boulenger (1885), because she listed the distribution of G. albofasciolatus based on G��nther (1872) under the species G. smithii. Recently, Kluge (2001) revalidated G. albofasciolatus at the specific rank, but Malkmus et al. (2002) and Das (2004) did not consider G. albofasciolatus as part of the Bornean herpetofauna. According to its original description (G��nther 1867), G. albofasciolatus differs from G. s m i t h i i by a different coloration of the dorsum (reddish-olive, marbled with grayish in G. albofasciolatus) and the venter (lower parts uniform whitish in G. albofasciolatus). Both species correspond well with respect to size and characteristic scalation (e.g., nares not in contact with rostral; postmentals relatively short; number of supralabials and sublabials; dorsal tubercle rows; precloacal pores) as well as in head and back pattern. A distinguishing feature at present is the number of ventrals (26 in G. albofasciolatus versus 29���39 in G. smithii). However, further specimen-based studies are needed to confirm whether G. albofasciolatus in fact deserves specific recognition. Concerning G. s m i t h i i, Das (2004, 2007) depicted specimens from Borneo, which did not show a Y-shaped head pattern, as is characteristic for specimens from Thailand and Malaysia (see also figures in Chan-ard et al. 1999; Grossmann 2006). A specimen from eastern Kalimantan showed a brown vertebral stripe and very small white dorsal flecks (Grossmann 2006). Lim & Lim (1999) recorded G. s m i t h i i for the first time from Tioman Island, Malaysia (see Grossmann & Tillack 2004, 2005), which has been followed by Hien et al. (2001), and Grismer et al. (2002, 2004). However, the Pulau Tioman form differs from G. s m i t h i i sensu stricto according to Grossmann (2006) in eye coloration (turquous blue versus mossy green) and in ground coloration (gray brown versus green) and has been considered distinctive by Grismer (2006). In life, the form from Tioman more closely resembles G. v e r re a u x i, from which it differs in the lighter eye coloration (emerald green in G. verreauxi) and the position of the nostril. We therefore suspect the form from Tioman Island to represent a distinct species and thus refer to it as Gekko sp. ���Tioman���. Additionally, Koch et al. (2009) published the first records of G. smihii from Sulawesi and Pulau Togian as well as from Pulau Batudaka. These authors argued against human introduction and rather believed that a natural dispersal from the Sunda shelf had occurred. Slight differences of the forms from Sulawesi and Togian compared with G. smithii s. str. were explained by Koch et al. (2009) as incipient divergence stemming from isolation, but they could not exclude the possibility that a cryptic species complex exists. Accordingly, we herein treat these both forms tentatively under the names Gekko sp. "Sulawesi" and Gekko sp. "Togian". We herein consider the species of the Gekko gecko group a monophyletic unit. Species displaying green to blue or brick red iris (i.e., G. albofasciolatus, G. nutaphandi, G. siamensis, G. smithii, G. v e r re a u x i, Gekko sp. ���Tioman���, Gekko sp. ���Sulawesi���, Gekko sp. ���Togian���) and more-or-less green ground coloration seem to be closely allied. For remarks on Gekko gecko and congeners see below., Published as part of R��sler, Herbert, Bauer, Aaron M., Heinicke, Matthew P., Greenbaum, Eli, Jackman, Todd, Nguyen, Truong Quang & Ziegler, Thomas, 2011, Phylogeny, taxonomy, and zoogeography of the genus Gekko Laurenti, 1768 with the revalidation of G. reevesii Gray, 1831 (Sauria: Gekkonidae), pp. 1-50 in Zootaxa 2989 on pages 11-13, DOI: 10.5281/zenodo.278393, {"references":["Gunther, A. (1867) Additions to the knowledge of Australian reptiles and fishes. Annals and Magazine of Natural History, (4) 20, 45 - 68.","Linnaeus, C. (1758) Systema Naturae per Regna Tria Naturae, Secundum Classes, Ordines, Genera, Species, cum Characteribus, Differentiis, Synonymis, Locis. Edito decima. Tomus I. Laurenti Salvi [Lars Salvius], Holmiae [Stockholm]. [4] + 823 + [1] pp.","Bauer, A. M., Sumontha, M. & Pauwels, O. S. G. (2008) A new red-eyed Gekko (Reptilia: Gekkonidae) from Kanchanaburi Province, Thailand. Zootaxa, 1750, 32 - 42.","Grossmann, W. & Ulber, T. (1990) Ein neuer Gecko aus Zentral-Thailand: Gekko siamensis sp. nov. (Reptilia: Sauria: Gekkonidae). Sauria, 12 (3), 9 - 18.","Tytler, R. C. (1864) Observations on a few species of gecko alive in the possesion of the author. Journal of the Asiatic Society of Bengal, 33, 535 - 548.","Kraus, F. (2009) Alien Reptiles and Amphibians, a Scientific Compendium and Analysis. Springer Verlag, Dordrecht, xii + 567 pp., CD ROM.","Smith, M. A. (1935) The Fauna of British India, Including Ceylon and Burma. Reptilia and Amphibia. II. Sauria. Taylor & Francis Ltd., London, xiii + 440 pp., 1 pl., 2 maps.","Wermuth, H. (1965) Liste der rezenten Amphibien und Reptilien. Gekkonidae, Pygopodidae, Xantusiidae. Das Tierreich, 80, Walterv de Gruyter, Berlin, XXII + 246 pp.","Gunther, A. (1872) On the reptiles and amphibians of Borneo. Proceedings of the Zoological Society of London, 1872, 586 - 600.","Bleeker, P. (1857) Berigt omtrent eenige Reptilien van Sumatra, Borneo, Batjan en Boero. Natuurkundig Tijdschrift voor Nederlandsch Indie, (4) 13, 470 - 475.","Bleeker, P. (1860) Over de Reptilien-Fauna van Sumatra. Natuurkundig Tijdschrift voor Nederlandsch Indie, (4) 21, 284 - 298.","Gunther, A. (1864) The Reptiles of British India. The Ray Society London, xxvii + 452 pp., 26 pls.","Gray, J. E. (1845) Catalogue of the Specimens of the Lizards in the Collection of the British Museum. British Museum, London, xxiii + 289 pp.","Dumeril, A. M. C. & Bibron, G. (1836) Erpetologie Generale ou Histoire Naturelle Complete des Reptiles. Paris (Roret), 3, iv + 517 pp, errata, 2 folding tables, pls. 25 - 28, 33, 35, 43 - 48.","Boulenger, G. A. (1887) Note on some reptiles from Sumatra described by Bleeker in 1860. Annals and Magazine of Natural History, (5) 20, 152.","Bauer, A. M. (1994) Gekkonidae (Reptilia, Sauria), Part I Australia and Oceania. Das Tierreich 109 (Part). Walter de Gruyter, Berlin, XII + 306 pp.","Kluge, A. G. (2001) Gekkotan lizard taxonomy. Hamadryad, 26, 1 - 209.","Rosler, H. (2000) Kommentierte Liste der rezent, subrezent und fossil bekannten Gecko-Taxa (Reptilia: Gekkonomorpha). Gekkota, 2, 28 - 153.","Boulenger, J. A. (1885) Catalogue of the lizards in the British Museum (Natural History). I. Geckonidae, Eublepharidae, Uroplatidae, Pygopodidae, Agamidae. British Museum (Natural History), London, xii + 436 pp., 32 pls.","Kluge, A. G. (1991) Checklist of gekkonoid lizards. Smithsonian Herpetological Information Service, 85, 1 - 35.","Kluge, A. G. (1993) Gekkonoid Lizard Taxonomy. International Gecko Society, San Diego, 245 pp.","De Rooij, N. (1915) The Reptiles of the Indo-Australian Archipelago. I. Lacertilia, Chelonia, Emydosauria. E. J. Brill Ltd., Leiden, xiv + 384 pp.","Malkmus, R., Manthey, U., Vogel, G., Hoffmann, P. & Kosuch, J. (2002) Amphibians and Reptiles of Mount Kinabalu (North Borneo). A. R. G. Gantner Verlag Kommanditgesellschaft, Ruggell, 424 pp.","Chan-ard, T., Grossmann, W., Gumprecht, A., Schulz, K. D. (1999). Amphibians and reptiles of Peninsular Malaysia and Thailand. Bushmaster Publications, Wuerselen, 240 pp.","Grossmann, W. (2006) Grunaugengeckos Gekko smithii & Gekko siamensis. Natur und Tier - Verlag GmbH, Munster, 62 pp.","Lim, K. K. P. & Lim, L. J. (1999) The terrestrial herpetofauna of Pulau Tioman, Peninsular Malaysia. The Raffles Bulletin of Zoology, 6. 131 - 155.","Grossmann, W. & Tillack, F. (2004) Pulau Tioman - Perle im Sudchinesischen Meer (Teil 1). Reptilia, 9 (6), 42 - 49.","Grossmann, W. & Tillack, F. (2005) Pulau Tioman - Perle im Sudchinesischen Meer (Teil 2). Reptilia, 10 (1), 56 - 64.","Hien, P., Grossmann, W. & Schafer, C. (2001) Beitrag zur Kenntnis der landbewohnenden Reptilienfauna von Pulau Tioman, West Malaysia. Sauria, 23 (4), 11 - 28.","Grismer, L. L., McGuire, J. A., Sosa, R. A. & Kaiser, H. (2002) Revised checklist and comments on the terrestrial herpetofauna of Pulau Tioman, Peninsular Malaysia. Herpetological Review, 33, 26 - 29.","Grismer, J. L., Grismer, L. L., Das, I., Yaakob, N. S., Liat, L. B., Leong, T. M., Youmans, T. M. & Kaiser, H. (2004) Species diversity and checklist of the herpetofauna of Pulau Tioman, Peninsular Malaysia, with a preliminary overview of habitat utilization. Asiatic Herpetological Research, 10, 247 - 279.","Grismer, L. L. (2006) Amphibians and Reptiles of the Tioman Archipelago, Malaysia. Forestry Department Peninsular Malaysia, Kuala Lumpur, 215 pp.","Koch, A., McGuire, J. A., Arida, E., Riyanto, A. & Hamidy, A. (2009) First record of Gekko smithii Gray, 1842 (Reptilia: Gekkonidae) from Sulawesi, Indonesia. Salamandra, 45, 172 - 176."]}

Gekko vittatus group (Figure 7) Species. G. vittatus Houttuyn, 1782 (Figure 7 A���B); Gekko sp. ��� Belau ���; Gekko sp. ���Malaupaina.��� Characters. Maximum SVL 140.0 mm; nares in contact with rostral; nasals 3���4; postmentals relatively small; dorsal tubercle rows 12���14; precloacal pores 14���58; postcloacal tubercles 1���3; no webbing between fingers and toes; fore and hind limbs with tubercles; lateral folds with tubercles; subcaudals not enlarged; head unicolored, without pattern; nominate form with white, anteriorly bifurcated dorsal stripe. Distribution. G. vittatus is known from Indonesia (from Java to New Guinea), Papua New Guinea, the Solomon Islands and northern Vanuatu (Torres and Banks Islands, Malakula Island); Gekko sp.��� Belau ��� is currently recorded from the Republic of Palau; Gekko sp. ���Malaupaina��� is currently reported from Malaupaina Island (Olu Malau Islands), Solomons (Crombie & Pregill 1999; McCoy 2006). Remarks. Gekko vittatus was described by Houttuyn (1782) with the locality data ���zekerlyk uit de India.��� Fitzinger (1826) referred the species to the genus Platydactylus Goldfuss, 1820 (see also Schlegel 1827; Wagler 1830; Dum��ril & Bibron 1836), which was subsequently split into three subgenera by Fitzinger (1843), of which vittatus belonged to his new subgenus Scelotretus. Lacerta unistriata, which was described by Shaw (1792), proved to be a synonym of G. vittatus (e.g., Daudin 1802; Gray 1845; Boulenger 1885; Wermuth 1965; Bauer 1994; Kluge 2001). Likewise Platydactylus bivittatus, which was described by Dum��ril & Bibron (1836) from New Guinea and Waigeo Island, Indonesia, was synonymized with G. vittatus (Loveridge 1948). Fitzinger���s (1843) Platydactylus (Scelotretus) bivittatus and Boulenger���s (1885) Gecko vittatus var. bivittatus referred to the same taxon. Gecko trachylaemus, described by Peters (1873), was also synonymized with G. vittatus (Peters & Doria 1878). The number of preanofemoral pores recorded in G. vittatus varies considerably according to different authors (Boulenger 1885: 50���59, De Rooij 1915: 14���58, Loveridge 1948: 21���57). Such variation of preanofemoral pores has not been described for any other Gekko species and likely points to the existence of cryptic species. However, in the series of G. vittatus from New Guinea and Ambon Island which was studied by us, we could not find any significant differences (see Table 1). With respect to pattern, we observed only slight variation in the number of light tail bands (4���6) and in the shape of the vertebral stripe on the tail and its bifurcation in the shoulder region (bifurcated branches may reach only as far as the back of head or may reach the eyes). Gekko sp. ���Malaupaina��� is known from the eastern Solomons (e.g., Malaupaina Island). This form differs from G. vittatus s. str. in the coloration of the iris (gray versus light brown), ground coloration (black olive versus brown olive) and in pattern (light vertebral stripe that bifurcates on the neck lacking in G. v i t t a t u s s. str.), see McCoy (1980: Pl. 2, fig. D, Pl. 6, fg. A) and McCoy (2006: Pl. 15). Mertens (1934: plate 2, fig. 7) pointed to a similarly patternless G. vittatus s. lat. from Nissan Atoll (Green Island Group, Papua New Guinea). The taxonomy of the southern Malaupaina form clearly deserves further research. Gekko vittatus from the Belau Islands (Republic of Palau) was mentioned as a distinct species by Crombie & Pregill (1999), however, it remains undescribed. The obscurely striped Gekko sp. ��� Belau ��� was occasionally referred to Gecko vittatus bivittatus (e.g., Boulenger 1885; Dryden & Taylor 1969), but Crombie & Pregill (1999) refer to significant differences between Gekko sp. ��� Belau ��� and the unstriped Gekko vittatus from New Guinea and the Solomons. The Belau form of G. vittatus shows some genetic divergence from G. vittatus across the remainder of the range (see above). The G. vittatus of Vanuatu shares its greatest molecular affinity with the form from the southern Solomons (see Figure 1)., Published as part of R��sler, Herbert, Bauer, Aaron M., Heinicke, Matthew P., Greenbaum, Eli, Jackman, Todd, Nguyen, Truong Quang & Ziegler, Thomas, 2011, Phylogeny, taxonomy, and zoogeography of the genus Gekko Laurenti, 1768 with the revalidation of G. reevesii Gray, 1831 (Sauria: Gekkonidae), pp. 1-50 in Zootaxa 2989 on pages 19-21, DOI: 10.5281/zenodo.278393, {"references":["Houttuyn, M. (1782) Het onderschied der salamanderen van de haagdissen in ' t algemeen, en van de gekkos in ' t byz'onder, aangetood. Verhandelingen Uitgegeven door het Genootschap der Wetenschappen te Vlissingen, 9, 305 - 336, 1 pl.","Crombie, R. I. & Pregill, G. K. (1999) A checklist of the herpetofauna of the Palau Islands (Republic of Belau), Oceania. Herpetological Monographs, 13, 29 - 80.","McCoy, M. (2006) Reptiles of the Solomon Islands. Pensoft Publishers, Sofia, 147 pp.","Fitzinger, L. I. (1826) Neue Classification der Reptilien nach ihren naturlichen Verwandtschaften. Ver l ag J. G. He ub ner, Wi en, 66 pp.","Schlegel, H. (1827) Herpetologische Nachrichten. Isis von Oken, 20, columns 281 - 294.","Wagler, J. G. (1830) Naturliches System der Amphibien mit vorangegangener Classification der Saugethiere und Vogel. J. G. Cotta´sche Buchhandlung, Munchen, vi + 354 pp., 2 pls.","Dumeril, A. M. C. & Bibron, G. (1836) Erpetologie Generale ou Histoire Naturelle Complete des Reptiles. Paris (Roret), 3, iv + 517 pp, errata, 2 folding tables, pls. 25 - 28, 33, 35, 43 - 48.","Fitzinger, L. I. (1843) Systema reptilium. Amblyglossae. Braumuller & Seidel, Vindobonae, 106 pp.","Shaw, G. (1792) The single-striped lizard. Naturalist's Miscellany, Vol. 3, Nodder & Co., London, 2 pp., pl. 89.","Daudin, F. M. (1802) Histoire Naturelle, Generale et Particuliere des Reptiles. Vol. 4. F. Dufart, Paris, 397 pp, 13 pls.","Gray, J. E. (1845) Catalogue of the Specimens of the Lizards in the Collection of the British Museum. British Museum, London, xxiii + 289 pp.","Boulenger, J. A. (1885) Catalogue of the lizards in the British Museum (Natural History). I. Geckonidae, Eublepharidae, Uroplatidae, Pygopodidae, Agamidae. British Museum (Natural History), London, xii + 436 pp., 32 pls.","Wermuth, H. (1965) Liste der rezenten Amphibien und Reptilien. Gekkonidae, Pygopodidae, Xantusiidae. Das Tierreich, 80, Walterv de Gruyter, Berlin, XXII + 246 pp.","Bauer, A. M. (1994) Gekkonidae (Reptilia, Sauria), Part I Australia and Oceania. Das Tierreich 109 (Part). Walter de Gruyter, Berlin, XII + 306 pp.","Kluge, A. G. (2001) Gekkotan lizard taxonomy. Hamadryad, 26, 1 - 209.","Loveridge, A. (1948) New Guinean reptiles and amphibians in the Museum of Comparative Zoology and United States National Museum. Bulletin of the Museum of Comparative Zoology, 101, 305 - 430.","Peters, W. C. H. (1873) Uber eine, zwei neue Gattungen enthaltende, Sammlung von Batrachiern des Hrn. Dr. O. Wucherer aus Bahia, so wie uber einige neue oder weniger bekannte Saurier. 2. Uber neue oder wenig bekannte Sauria. Monatsberichte der koniglich Akademie der Wissenschaften zu Berlin, 1872, 774 - 776.","Peters, W. C. H. & Doria, G. (1878) Catalogo dei rettili e dei batraci raccolti da O. Beccari, L. M. D'Albertis e A. A. Bruijn nella Sotto-Regione Austro-Malese. Annali del Museo Civico di Storia Naturale di Genova, 13, 323 - 450, pls. I - VII.","De Rooij, N. (1915) The Reptiles of the Indo-Australian Archipelago. I. Lacertilia, Chelonia, Emydosauria. E. J. Brill Ltd., Leiden, xiv + 384 pp.","McCoy, M. (1980) Reptiles of the Solomon Islands. Handbook No. 7, Wau Ecology Institute, Wau, Papua New Guinea, 80 pp.","Mertens, R. (1934) Die Inselreptilien, ihre Ausbreitung, Variation und Artbildung. Zoologica, 32, 1 - 209.","Dryden, G. L. & Taylor, E. H. (1969) Reptiles from the Mariana and Caroline Islands. University of Kansas Science Bulletin, 48, 269 - 279."]}

Gekko japonicus group (Figures 4, 5A) Species. G. auriverrucosus Zhou & Liu 1982; G. canhi R��sler, Nguyen, Doan, Ho & Ziegler 2010; G. chinensis Gray, 1842; G. hokouensis Pope, 1928 (Figure 4 B); G. japonicus (Schlegel, 1836) (Figure 4 A); G. l i b o e n s i s Zaho & Li, 1982; G. melli Vogt, 1922; G. palmatus Boulenger, 1907 (Figure 4 C); G. scabridus Liu & Zhou, 1982; G. s c i e n - tiadventura R��sler, Ziegler, Vu, Herrmann & B��hme, 2004 (Figure 4 D); G. shibatai Toda, Sengoku, Hikida & Ota, 2008; G. similignum Smith, 1923; G. subpalmatus G��nther, 1864; G. swinhonis G��nther, 1864; G. taibaiensis Song, 1985; G. t a w a e n s i s Okada, 1956; G. vertebralis Toda, Sengoku, Hikida & Ota, 2008 (Figure 5 A); G. wenxianensis Zhou & Wang, 2008; G. yakuensis Matsui & Okada, 1968. Characters. 58.9���99.2 mm SVL; nares in contact with rostral (except for G. auriverrucosus); nasals 3 (rarely 2 in G. c h i n e n s i s); postmentals relatively small (e.g., G. similignum), largest in G. c a n h i, G. c h i n e n s i s, G. palmatus, G. scientiadventura; 0���21 dorsal tubercle rows; 0���32 precloacal pores; postcloacal tubercles 1���4; webbing between fingers and toes weakly developed to extensive (G. chinensis, G. m e l l i, G. palmatus, G. scientiadventura, G. similignum, G. subpalmatus); tubercles present on fore and hind limbs, hind limbs only, or lacking all together; lateral fold without tubercles; subcaudals enlarged, in a longitudinal row (in G. yakuensis medially subdivided); head pattern present or not, without figure-shape (UU- to W-shaped in G. m e l l i and W-shaped in G. scientiadventura); vertebral region with relatively large, light flecks, blotches or bands. Distribution. China, Japan, Korea, Taiwan, and Vietnam including offshore islands; possibly Laos (Figure 3 A). Remarks. The Gekko japonicus group is very complex. Different morphological features of the species of this group (e. g., nares in contact with rostral or not, broad webbing between fingers and toes or not, dorsal tubercles absent or present, tubercles on limbs absent or present, see Table 2) demonstrate that this genetically well-defined clade is highly morphologically variable (see Figure 1). Numerous authors have listed Gekko chinensis as a representative of Vietnam���s herpetofauna (e.g., Bourret 1937 b, 1940 a, 1943; Zhao & Adler 1993; Welch 1994; Bobrov 1995 a; R��sler et al. 2005). According to Nguyen et al. (2005), G. chinensis is widely distributed in the north of the country, whereas G. palmatus is only recorded from the two northern provinces of Lang Son and Vinh Phuc (see also Nguyen & Ho 1996). However, according to Ziegler et al. (2006) and based on new locality records (see Appendix 1) the distribution of G. palmatus in Vietnam in fact extends from several northern provinces southward to central Vietnam in Quang Binh Province. Taking into account the records given for G. c h i n e n s i s by Nguyen & Ho (1996), the latter species must occur in sympatry with G. palmatus at least in the northern Vietnamese provinces of Bac Giang, Bac Kan, Cao Bang, Lang Son, Quang Ninh, and Vinh Phuc. However, all Gekko studied by us from these provinces proved to be G. palmatus and none had the diagnostic characters of G. c h i n e n s i s. This also makes the records of G. chinensis from Hai Duong and Ninh Binh provinces (see Nguyen & Ho 1996) questionable and they are probably based on misidentifications of G. palmatus. Bobrov & Semenov (2008) listed all previous identifications of G. chinensis from Vietnam as G. palmatus. Ota et al. (1995) believed G. c h i n e n s i s and G. palmatus to represent allopatric taxa (see also Matsui & Ota 1995). Concordant with the latter authors we herein suggest that the known occurrence of G. chinensis is limited to China (southern provinces of Fujian, Guangdong, Guangxi, Guizhou, Hong Kong, Sichuan, and Yunnan) whereas that of G. palmatus is only in northern and central Vietnam (Bac Giang, Bac Kan, Cao Bang, Hai Duong, Hai Phong, Lang Son, Ninh Binh, Quang Binh, Quang Ninh, and Vinh Phuc provinces). Further collections or study of museum specimens are needed to clarify the identification of populations along the China-Vietnam border and whether G. chinensis and G. palmatus occur in sympatry. The distinguishing character of broad toe webbing in G. palmatus versus a narrower webbing in G. c h i n e n s i s, which can regularly be found in the older literature (e.g., Boulenger 1907; Smith 1935; Bourret 1937 b), may have conspicuously added to the above listed confusion and misidentifications, because it represents an ambiguous diagnostic character (Ota et al. 1995). Smith (1935: 114) even concluded that G. palmatus is ���possibly a racial form of G. c h i n e n s i s.��� However, G. palmatus and G. chinensis can be easily distinguished by the size of the internasals compared to the nasorostrals (internasals same size or larger than nasorostrals in G. chinensis versus always smaller in G. palmatus) and statistically significant differences in the number of midbody scales and the number of lamellae on the first, third and fifth fingers as well as the first toe (Ota et al. 1995). A key character that allows for proper identification of both aforementioned species is the presence of limb tubercles. These are present in G. c h i n e n s i s, but are completely lacking in G. palmatus (our data). Ota et al. (1995) indicated shank tubercles for G. chinensis and the specimen depicted in Zhao & Adler (1993: pl. 21, fig. G.) has tubercles both on the fore and hind limbs; but the specimen of G. palmatus depicted by Nguyen et al. (2009: fig. 314) completely lacks such tubercles. The taxonomic history of G. similignum is controversial. The type locality of this taxon is Hainan Island (Smith 1923). Stejneger (1932) synonymized G. similignum with G. chinensis (see also Zhao et al. 1999). However, Ota et al. (1995) and Kluge (2001) listed it as valid at the specific rank. Gekko chinensis and G. smilignum differ in maximum SVL, in the number of scales around midbody and in the number of lamellae on the first and fourth toes (see Table 1). Gekko liboensis was described by Zhou & Li (1982) based on an adult female specimen collected in 1979. Three further specimens (two females and a juvenile) were collected in 1984 (Zhao et al. 1999). Zhao & Adler (1993) synonymized G. liboensis with G. hokouensis (see also G��nther 1994; Kluge 2001; Henkel & Schmidt 2003). In contrast, Kluge (1993), Bauer (1994), Welch (1994), Matsui & Ota (1995), Zhao et al. (1999), R��sler (2000), R��sler et al. (2005, 2006) treated G. liboensis as a valid species. Gekko liboensis differs from G. hokouensis by a larger SVL (85 versus 70 mm), absence of internasals, more interorbitals (40 versus 30���33), flat dorsal tubercles, fewer dorsal tubercle rows (10 versus 12���18) and by the number of lamellae below the fourth toe (9 versus 15���18). However, as adult males are still lacking for G. l i b o e n s i s, important diagnostic characters and possibly additional distinguishing features such as the number of precloacal pores are still missing. The nares of G. auriverrucosus are not in contact with the first supralabial (Zhou & Liu 1982), which seems to represent a derived character of the Gekko gecko group (except for G. verreauxi). Zhao & Adler (1993) synonymized G. scabridus with G. chinensis (see also Kluge 2001), but Matsui & Ota (1995) and Zhao et al. (1999) again evaluated the former taxon as valid species. Liu & Zhou (1982) pointed to the phenotypic similarity between G. scabridus and G. japonicus. According to Liu & Zhou (1982), G. scabridus differs from G. japonicus by having tubercles on its back and hind limbs and by a higher count of precloacal pores. From Table 1 it is obvious that there exist additional distinguishing features between the two species. Compared with G. japonicus, G. scabridus has fewer interorbitals (30 versus 32���35), fewer lamellae below the first toe (6���9 versus 10���12), and fourth toe (7���9 versus 14���16), more dorsal tubercle rows (17���21 versus 9���14) and a higher number of precloacal pores (10���15 versus 6���9). In comparsion to G. chinensis it has only slightly developed webbing between the fingers and toes, fewer precloacal pores, and more postcloacal tubercles. We, therefore, accept the specific status of G. scabridus and regard it as a valid member of the Gekko japonicus group. Vogt (1922) originally described G. m e l l i based on a large series of which now only two specimens still exist (Bauer & G��nther 1991). Stejneger (1932) listed G. m e l l i as a subspecies of G. subpalmatus and Pope (1935) finally synonymized the former species with G. subpalmatus. R��sler et al. (2005) revalidated G. melli as a distinct species (see also R��sler & Tiedemann 2007). With respect to G. subpalmatus it is interesting to note that the back of the holotype is slightly banded and that the vertebral region shows four indistinct light blotches (R��sler et al. 2005: figs. 11���12). However, the ground coloration of the live specimen depicted in Zhao & Adler (1993: their Pl. 32, fig. B) is blackish-olive (Black 3 U). This specimen, in addition, has a repeatedly interrupted orange (722 U) vertebral stripe and tail blotches of the same color; a banded pattern as observable from the holotype is not discernible (see also fig. in Huang & Zong 1998: pl. 29). Whether this reflects intraspecific variation or a persistent confusion over the identity of this species remains to be investigated., Published as part of R��sler, Herbert, Bauer, Aaron M., Heinicke, Matthew P., Greenbaum, Eli, Jackman, Todd, Nguyen, Truong Quang & Ziegler, Thomas, 2011, Phylogeny, taxonomy, and zoogeography of the genus Gekko Laurenti, 1768 with the revalidation of G. reevesii Gray, 1831 (Sauria: Gekkonidae), pp. 1-50 in Zootaxa 2989 on pages 13-17, DOI: 10.5281/zenodo.278393, {"references":["Rosler, H., Nguyen, Q. T., Doan, V. K., Ho, T. C., Nguyen, T. T. & Ziegler, T. (2010) A new species of the genus Gekko Laurenti, 1768 (Squamata: Sauria: Gekkonidae) from Vietnam with remarks on G. japonicus (Schlegel, 1836). Zootaxa, 2329, 56 - 68.","Schlegel, H. (1836) Le Platydactyle Monarque. Platydactylus Monarchus. In: Dumeril, A. M. C. & Bibron, G. (1836), Erpetologie Generale ou Histoire Naturelle Complete des Reptiles. Vol. 3. Roret, Paris, pp. 335 - 336.","Vogt, T. (1922) Zur Reptilien- und Amphibienfauna Sudchinas. Archiv fur Naturgeschichte, 10, 135 - 146.","Boulenger, G. A. (1907) Description of new lizard in the British Museum. Annals and Magazine of Natural History, 19, 486 - 489.","Toda, M., Sengoku, S., Hikida, T. & Ota, H. (2008) Description of two new species of the genus Gekko (Squamata: Gekkonidae) from the Tokara and Amami Island Group in the Ryukyu Archipelago, Japan. Copeia, 2008, 452 - 466.","Smith, M. A. (1923) On a collection of reptiles and batrachians from the island of Hainan. Journal of the Natural History Society of Siam, 6, 195 - 212.","Gunther, A. (1864) The Reptiles of British India. The Ray Society London, xxvii + 452 pp., 26 pls.","Song, M. (1985) A new species of Gekko from Shaanxi. Acta Herpetologica Sinica, 4, 329 - 330.","Okada, Y. (1956) A new species of gekko from Shikoku, Japan. Annotationes Zoologicae Japonensis, 29, 239 - 241.","Zhou, K. & Wang, Q. (2008) New species of Gekko (Squamata: Sauria: Gekkonidae) from China: morphological and molecular evidence. Zootaxa, 1778, 59 - 68.","Matsui, T. & Okada, Y. (1968) A new species of Gekko found in Yakushima, one of the small island south of Kyushu. Acta Herpetologica Japonica, 3, 1 - 4.","Bourret, R. (1937 b) Notes herpetologiques sur l'Indochine francaise. XV. Lezards et Serpents recus au Laboratoire des Sciences Naturelles de l'Universite au cours de l'annee 1937. Descriptions de deux especes et de deux varietes nouvelles. Bulletin general de l'Instruction publique, 17 (4), annexe, 57 - 80.","Bourret, R. (1940 a \" 1939 \") Notes herpetologiques sur l'Indochine francaise. XVIII. Reptiles et Batraciens recus au Laboratoire des Sciences Naturelles de l'Universite au cours de l'annee 1939. Descriptions de quatre especes et d'une variete nouvelles. Bulletin general de l'Instruction publique, 18 (4), annexe, 5 - 39, 1 pl.","Bourret, R. (1943) Notes herpetologiques sur l'Indochine francaise. XXIV. Reptiles et Batraciens recus au Laboratoire des Sciences Naturelles de l'Universite au cours de l'annee 1942. Notes et travaux de l'Ecole Superieure des sciences de l'Universite Indochinoise, 2, 1 - 17, 2 pls.","Zhao, E. & Adler, K. (1993) Herpetology of China. Society for the Study of Amphibians and Reptiles, Oxford, Ohio, 522 pp., 48 pls., 1 foldout map.","Welch, K. R. G. (1994) Lizards of the World. 1. Geckos. R & A Research and Information Limited, Taunton, 165 pp.","Bobrov, V. V. (1995 a) Checklist and bibliography of the Lizards of Vietnam. Smithsonian Herpetological Information Service, 105, 1 - 28.","Nguyen, V. S., Ho, T. C. & Nguyen, Q. T. (2005) A Checklist of Amphibians and Reptiles of Vietnam [in Vietnamese]. Agriculture Publishing House, Hanoi, 180 pp.","Nguyen, V. S. & Ho, T. C. (1996) A Checklist of the Reptiles and Amphibians of Vietnam [in Vietnamese]. Science and Technique Publishing House, Hanoi, 264 pp.","Ziegler, T., Ohler, A., Vu, N. T., Le, K. Q., Nguyen, X. T., Dinh, H. T. & Bui, N. T. (2006) Review of the amphibian and reptile diversity of Phong Nha-Ke Bang National Park and adjacent areas, central Truong Son, Vietnam. In: Vences, M., Kohler, J., Ziegler, T. & Bohme, W. (eds), Herpetologia Bonnensis II. Proceedings of the 13 th Congress of the Societas Europaea Herpetologica, Bonn, pp. 247 - 262.","Bobrov, V. V. & Semenov, D. V. (2008) Jastschjerizy Vietnama [Lizards of Vietnam]. Towarristschestowo nautschnyx isdanii KMK, Moscow, 225 pp.","Matsui, M. & Ota, H. (1995) On Chinese herpetology. Herpetologica, 51 (2), 234 - 250.","Smith, M. A. (1935) The Fauna of British India, Including Ceylon and Burma. Reptilia and Amphibia. II. Sauria. Taylor & Francis Ltd., London, xiii + 440 pp., 1 pl., 2 maps.","Nguyen, V. S., Ho, T. C. & Nguyen, Q. T. (2009) Herpetofauna of Vietnam. Edition Chimaira, Frankfurt am Main, 768 pp.","Stejneger, L. (1932) The Chinese lizards of the genus Gekko. Proceedings of the United States National Museum, 132, 1 - 6.","Zhao, E., Zhao, K. & Zhou, K. (1999) Fauna Sinica. Reptilia Vol. 2. Squamata, Lacertilia [in Chinese]. Science Press, Beijing, xi + 394 pp., 8 pls.","Kluge, A. G. (2001) Gekkotan lizard taxonomy. Hamadryad, 26, 1 - 209.","Gunther, R. (1994) Eine neue Art der Gattung Gekko (Reptilia, Squamata, Gekkonidae) aus dem Suden Vietnams. Zoologischer Anzeiger, 233, 57 - 67.","Henkel, F. W. & Schmidt, W. (2003) Praxis Ratgeber Geckos. Edition Chimaira, Frankfurt am Main, 159 pp.","Kluge, A. G. (1993) Gekkonoid Lizard Taxonomy. International Gecko Society, San Diego, 245 pp.","Bauer, A. M. (1994) Gekkonidae (Reptilia, Sauria), Part I Australia and Oceania. Das Tierreich 109 (Part). Walter de Gruyter, Berlin, XII + 306 pp.","Rosler, H. (2000) Kommentierte Liste der rezent, subrezent und fossil bekannten Gecko-Taxa (Reptilia: Gekkonomorpha). Gekkota, 2, 28 - 153.","Bauer, A. M. & Gunther, R. (1991) An annotated type catalogue of the geckos (Reptilia: Gekkonidae) in the Zoological Museum, Berlin. Mitteilungen aus dem Zoologischen Museum in Berlin, 67, 279 - 310.","Pope, C. H. (1935) The Reptiles of China. Turtles, Crocodilians, Snakes, Lizards. The American Museum of Natural History, New York, lii + 604 pp.","Rosler, H. & Tiedemann, F. (2007) Gekko melli Vogt, 1922 and its types (Reptilia, Sauria, Gekkonidae). Mitteilungen aus dem Museum fur Naturkunde in Berlin, Zoologische Reihe, 83, 105 - 108.","Huang, Z. Y. & Zong, Y. (1998) Zhongguo Techan de Paxing Dongwu. Fudan University, Shanghai, 116 pp."]}

Gekko reevesii Gray, 1831 1831 Gecko reevesii Gray in Griffith & Pidgeon, Anim. Kingdom Cuvier., 9 Synops. Spec.: 48. Diagnosis. A large species of Gekko, with SVL> 150 mm; head distinct from body; body somewhat flattened; tail round in section, not thickened at base; supralabials 10���14; sublabials 9���13; nares in contact with rostral; nasals 3��� 6; internasal 0���1; interorbitals 17���29; dorsal tubercle rows 12���18; scales between mental and cloacal slit 133���141; midbody scales 82���103; subdigital lamellae below first toe 13���19; subdigital lamellae below fourth toe 18���24; no extensive webbing between fingers and toes; forearm and hind limbs with tubercles; precloacal pores 13���20; postcloacal tubercles 1���4; tail tubercles present; subcaudals not enlarged; grayish brown to grayish green ground coloration, with reddish brown dorsal blotches; dorsal pattern tends to consist of transverse bands and short stripes. Holotype: BMNH 1946.8. 2598 ���male, ad., China, collector unknown (Figures 8, 9). SVL 130.0 mm; TL 110.0 mm; LT 28.5 mm; HL 36.4 mm; HW 26.2 mm; HH 14.0 mm; SE 11.5 mm; EE 12.4 mm. SVL/TL 1.18; SVL/LT 4.56; SVL/HL 3.57; HL/HW 1.39; HL/HH 2.60; SE/EE 0.93. Rostral concave above, wider than high (RW 4.80 mm, RH 2.70 mm, RW/RH 1.78), wider than mental (RW/ MW 1.71); 13 / 13 supralabials; 22 / 18 snout scales adjacent to supralabials; nares not in contact with rostral (Figure 9 A); 5 / 4 nasals; nasorostrals ca. double the size of supranasals and five times larger than postnasals; internasal single, large (Figure 9 B); snout medially with flat, elongate depression; lateral snout scales round to oval, curved, next to each other; ca. 12���13 large scales between postnasals and orbital cavity; 61 scales between seventh supralabials; medial snout scales barely smaller than lateral snout scales; dorsal ciliary scales 1.5 times higher than medial snout scales; temporal scales larger than lateral snout scales; ear opening vertical, oval; 20 interorbitals, granular, in orbital region twice as large as medially; back of head and neck granular; granular scales slightly larger than medial interorbitals; neck tubercles round, concial; mental pentagonal, longer than wide (MW 2.8 mm, ML 3.0 mm, MW/ ML 0.93), not distinctly larger than first sublabials (Figure 9 C); 11 / 11 sublabials; 17 / 15 gular scales adjacent to sublabials; 2 small postmentals, hexagonal, 1.5 times longer than wide, anteriorly in contact with mental and first sublabials; 5 gular scales adjacent to postmentals; outer gular scales ca. 1.5 times larger than inner ones; gular scales as large as medial snout scales, round, slightly curved, smooth, next to each other, arranged in more or less regular transverse rows; dorsals as large as medial snout scales, round flat, next to each other, arranged in more or less regular transversal rows; dorsal tubercles 2���3 times as large as adjacent dorsals (DTL 2.1 mm, DTW 1.5 mm, DTL/DTW 1.40, DTLx 100 /SVL 1.62, DTWx 100 /SVL 1.15), oval, asymmetrically conical (Figure 9 D), arranged in 14 more or less regular longitudinal rows, encircled by 9 dorsals; lateral tubercles smaller than dorsal ones (LTL 1.8 mm, LTW 1.2 mm, LTL/LTW 1.50, LTLx 100 /SVL 1.39, LTWx 100 /SVL 0.92); lateral folds slightly developed; ventrals flat, smooth, subimbricate, and larger than dorsals; 102 scales around midbody; limbs without tubercles; arm and forearm scales flat, smooth, subimbricate; shank with curved tubercles; dorsal and ventral thigh scales anteriorly flat, imbricate, posteriorly smaller, curved; enlarged femoral scales lacking; 9 / 10 tubercles on thigh; fingers and toes 1���4 basally with narrow webbing; 19 / 19 subdigital lamellae below first finger and 21 / 23 below fourth finger; 18 / 18 subdigital lamellae below first toe and 23 / 24 below fourth toe; 19 precloacal pores in a continuous, angular-shaped row (Figure 9 E); ca. five rows of enlarged scales behind precloacal pores; 1 / 1 blunt, conical postcloacal tubercles; tail not thickened at base, with whorls; dorsal tail scales larger than scales of body dorsum, more or less squarish, slightly curved, subimbricate, arranged in relatively regular transverse rows; six dorsal, medial scale rows in the third tail whorl; anterior tail tubercles asymmetrically conical, posterior ones curved; six tubercles in the third and fifth tail whorl, in a transverse row in posterior whorl region; four scale rows between tubercle rows 2���3 and 3 scale rows between tubercle rows 3���4; subcaudals larger than dorsal tail scales, flat, smooth, subimbricate, arranged in two parallel rows, 2���3 rows per whorl (Figure 9 F). Dorsum olive brown (4655 U); head with indistinct brown (478 U) stripes and blotches; back with dark brown bands (469 U); back tubercles light gray (Cool Gray 1 U) as well as encircling dorsals, resulting in a light flecked dorsal pattern; those light flecks displaced, in ca. 8 transverse rows; limbs marbled, with scattered light tubercles; six dark and six light tail bands, of which the dark ones are broader (Figure 8 A); venter gray (400 U); throat light gray, with indistinct pattern; medially stronger pigmented belly, with few brown (4635 U) blotches (Figure 8 B); anterior tail slightly banded, posterior parts with more distinct bands; tail tip dark. Further specimens: We list the characters of 42 additional specimens from China and Vietnam that proved to be morphologically assignable to G. reevesii (see Appendix 1): 70.0���173.0 mm SVL; 58.0���138.0 mm TL; 15.4��� 35.2 mm LT; 20.8���46.5 mm HL; 16.0��� 37.5 mm HW; 9.6���22.5 mm HH; 9.6���16.4 mm SE; 8.9���17.7 mm EE. Body proportions: SVL/TL 1.03���1.26; SVL/LT 4.25���5.34; SVL/HL 3.02���3.75; HL/HW 1.25���1.60; HL/HH 1.91���2.85; SE/EE 0.73���1.12. Scalation: RW 3.8���6.3 mm; RH 1.90���3.70 mm (RW/RH 1.41���2.20); MW 2.10���4.60 mm; ML 1.80 ���4.00 mm (MW/ML 0.85���1.50); RW/MW 1.20���2.14; 10���14 SPL; 9���13 SBL; 3���6 N; 0���1 IN; 17���29 IO; 2 PM; 3���6 GP; 12��� 18 DTR; 5���10 GSDT; 133���141 SMC; 86���102 SR; 28���32 V; 15���19 LF 1; 18���24 LF 4; 13���19 LT 1; 18���24 LT 4; 2���24 TT; 13���20 PP; 0���18 PS; 1���4 PAT; 6 T 1 W; 6 T 5 W; 3���5 scales between second and third and third and fourth caudal tubercle rows. reevesii; for abbreviations see Material and methods. continued next page Comparisons: Comparisons with subspecies of G. gecko: Characters of different geographic populations of G. gecko are in part overlapping. Mean values of supralabials, sublabials, dorsal tubercle rows, lamellae below fingers and toes change clinally from north to south (R��sler 2005 a). Gekko reevesii significantly differs (p G. g. gecko sensu stricto from Java, by having a smaller SVL, a proportionally smaller head height, fewer supralabials, interorbitals, femoral tubercles, and postcloacal tubercles, as well as by having greater number of tubercles rows, precloacal pores, lamellae below first finger, lamellae below first toe, and lamellae below fourth toe (see Tables 3��� 4). The ground coloration of G. reevesii is grayish brown to grayish green (Figure 10) versus ultramarine gray to bluish gray in G. g. gecko. The dorsal blotches of G. reevesii are reddish brown to dark brown versus cinnober red in the nominate form. Furthermore, the dorsal pattern of G. reevesii tends to comprise transverse bands and short stripes versus relatively irregularly arranged, distinct single blotches (see figures in Zhao & Adler 1993; R��sler 1995; Ziegler 2002; Grossmann 2004 a; Nguyen et al. 2009). Gekko reevesii does not co-occur with G. g. azhari. Because the latter was described based on two females only (Mertens 1955), and because no further specimens have been collected since then, comparisons and relations with adjoining Indian populations of G. gecko (e.g., West Bengal, Dhaka) are barely known and require further investigation (Peterson 1980; Ahmed & Dasgupta 1992; Tikader & Sharma 1992). Based on current knowledge, G. reevesii can be morphologically distinguished from G. g. azhari by its larger SVL and domed versus flat dorsal tubercles (see Tables 3���4). see Material and methods, except for T 3 W = tubercles in the third tail whorl; SB 2 / 3 = scales between second and third tail tubercle rows; SB 3 / 4 = scales between third and fourth tail tubercle rows. continued next page Our genetic samples did not include representatives of G. reevesii. However, Qin et al. (2007) found significant differences between geckos from as close as 85 km in Guangxi (red tokay from Nanning versus black tokays from Chongzuo and Qinzhou). Such differences (> 8 %) among probably parapatric, if not sympatric, taxa are certainly taxonomically noteworthy. In addition, further taxonomic subdivisions may also be present in Gekko gecko. Our molecular phylogeny revealed two relatively deeply divergent evolutionary lineages: a western clade (Myanmar, Thailand) and an eastern clade (China, Cambodia, Indonesia, Malaysia), within which close relationships of Chinese and Indonesian G. gecko exist (Figure 1). As extensive human-mediated movement of G. gecko occurs in association with the traditional medicine trade throughout the range of the species (Bauer 2009), and G. gecko is common around human habitations, it is possible that the observed relationships may be confounded by translocations. Certainly, much deeper sampling of G. g e c k o across its entire range is needed. Comparisons of the western G. gecko with G. g. azhari are particularly crucial, as is, of course, the evaluation of G. reevesii. Possible further evidence of multiple species in G. gecko comes from chromosomal studies. All G. gecko karyotyped to date have a diploid number of 38 and a fundamental number of 48 (Cohen et al. 1967; Wu & Zhao 1984; Solleder & Schmid 1984; Sharma & Kasid 1992; Du et al. 2002), however, Solleder & Schmid (1984) found heteromorphic sex chromosomes in males, while these were not reported by other workers, and some researchers have reported acrocentric pairs (Cohen et al. 1967), whereas others explicitly state that none are present (Sharma & Kasid 1992). Comparisons with remaining Gekko species: G. reevesii differs from all other Gekko species by its large SVL (> 170 mm), in combination with narial contact with rostral only, more than ten lamellae below fourth toe, subcaudals not enlarged, and a variable gray ground coloration (see identification key below). Distribution. Gekko reevesii is currently known from southern China (provinces of Fujian, Guangdong, Guangxi, and Yunnan) and northern Vietnam (southwards to Quang Binh Province). According Lue et al. (1999) and Shang (2001), several early 20 th century records of G. reevesii from Taiwan represent individuals artificially transported from the Chinese mainland. At least within Guangxi both red and black tokays occur in close proximity. Indeed, based on the sampling of Qin et al. (2007), red tokay (G. gecko gecko) populations from Nanning, Ningming County are virtually surrounded by black tokay (G. reevesii) sites. Both forms also occur in northern Vietnam. The exact boundaries of G. reevesii thus remain uncertain., Published as part of R��sler, Herbert, Bauer, Aaron M., Heinicke, Matthew P., Greenbaum, Eli, Jackman, Todd, Nguyen, Truong Quang & Ziegler, Thomas, 2011, Phylogeny, taxonomy, and zoogeography of the genus Gekko Laurenti, 1768 with the revalidation of G. reevesii Gray, 1831 (Sauria: Gekkonidae), pp. 1-50 in Zootaxa 2989 on pages 22-30, DOI: 10.5281/zenodo.278393, {"references":["Gray, J. E. (1831) A synopsis of the species of the class Reptilia. In Griffith, E. & Pidgeon, E. (eds.), The Animal Kingdom Arranged in Conformity with its Organization, by the Baron Cuvier. Vol. 9. The Class Reptilia Arranged by the Baron Cuvier, with Specific Descriptions, Whittaker, Treacher & Co., London, pp. 1 - 110.","Rosler, H. (2005 a) Studien am Tokeh: 2. Intraspezifische Variation der sudostasiatischen Populationen von Gekko gecko (Linnaeus, 1758) (Sauria: Gekkonidae). Gekkota, 5, 65 - 149.","Zhao, E. & Adler, K. (1993) Herpetology of China. Society for the Study of Amphibians and Reptiles, Oxford, Ohio, 522 pp., 48 pls., 1 foldout map.","Rosler, H. (1995) Geckos der Welt - Alle Gattungen. Urania Verlag, Leipzig, 256 pp.","Ziegler, T. (2002) Die Amphibien und Reptilien eines Tieflandfeuchtwald-Schutzgebietes in Vietnam. Natur und Tier-Verlag GmbH, Munster, 342 pp.","Grossmann, W. (2004 a) Der Tokeh, Gekko gecko. Natur und Tier-Verlag GmbH, Munster, 61 pp.","Nguyen, V. S., Ho, T. C. & Nguyen, Q. T. (2009) Herpetofauna of Vietnam. Edition Chimaira, Frankfurt am Main, 768 pp.","Mertens, R. (1955) Uber eine eigenartige Rasse des Tokehs (Gekko gecko) aus Ost-Pakistan. Senckenbergiana Biologica, 36, 21 - 24.","Peterson, T. (1980) Bangladesh 1979. Snoken, Uppsala, 10 (4), 84 - 101.","Ahmed, S. & Dasgupta, G. (1992) Reptilia. In: Ghosh, A. K. (ed.), Fauna of West Bengal. Part 2 (Reptilia, Amphibia, Fishes, Hemichordata and Archaeozoology). Zoological Survey of India, Calcutta, pp. 1 - 65.","Tikader, B. K. & Sharma, R. C. (1992) Handbook Indian lizards. Zoological Survey of India, Calcutta, xv + 250 pp.","Qin, X., Zeng, Z. & Liang, Y. (2007) Genetic variation and differentiation of Gekko gecko from different population [in Chinese]. Zoological Research, 28, 286 - 290.","Cohen, M. M., Huang, C. C. & Clark, H. F. (1967) The somatic chromosomes of 3 lizard species: Gekko gecko, Iguana iguana, and Crotaphytus collaris. Experientia, 23, 769 - 771.","Wu, G. & Zhao, E. (1984) Study on karyotypes of Gekko gecko and Gekko subpalmatus [in Chinese]. Acta Herpetologica Sinica, 3, 61 - 64.","Solleder, E. & Schmid, M. (1984) XX / XY-sex chromosomes in Gekko gecko (Sauria: Reptilia). Amphibia-Reptilia, 5, 339 - 345.","Sharma, G. P. & Kasid, U. N. (1992) A chromosomal survey of four species of gekkonid lizards. Cytobios, 69, 187 - 197.","Du, S., Cheng, L. & Liu, D. (2002) Advances of systematic research of the genus Gekko (Sauria: Gekkonidae) [in Chinese]. Sichuan Journal of Zoology, 21, 200 - 204.","Lue, K. Y., Tu, M. C. & Shang, G. S. (1999) A Field Guide to Amphibians and Reptiles in Taiwan. Society for Wildlife and Nature, Taipei, 343 pp.","Shang, G. (2001) Natural Portraits of Lizards of Taiwan [in Chinese]. Bigtrees Press, Taipei, 173 pp."]}

Gekko monarchus group (Figure 5 B���D) Species. G. carusadensis Linkem, Siler, Diesmos, Sy & Brown, 2010; G. e r n s t k e l l e r i R��sler, Siler, Brown, Demegillo & Gaulke, 2006 (Figure 5 C); G. kikuchii Oshima, 1912; G. mindorensis Taylor, 1919 (Figure 5 D); G. m o n a rchus (Schlegel, 1836) (Figure 5 B); G. palawanensis Taylor, 1925. Characters. 63.0���100.0 mm SVL; nares in contact with rostral; nasals 3 (rarely 4 in G. e r n s t k e l l e r i); postmentals relatively large; dorsal tubercle rows 10���24; precloacal pores 32���70; postcloacal tubercles 2���3 (1 in G. c a r u - sadensis and G. palawanensis); no webbing between fingers and toes; tubercles on fore and hind limbs (not on fore limbs of G. ernstkelleri); lateral folds without tubercles; subcaudals mostly enlarged, in a longitudinal row; more or less distinct, W-shaped head pattern; light (white to gray), mostly asymmetrically arranged dorsal and lateral flecks. Distribution. G. monarchus is known from the Philippines westwards to Myanmar and to Indonesia (Aru Islands, Kei Islands, New Guinea) in the south (Figure 3 A); G. kikuchii is endemic to Lanyu Island, Taiwan; remaining group members are endemic to the Philippines. Remarks. G. kikuchii was described by Oshima (1912) from Botel Tobago (Lanyu) near Taiwan. Many subsequently published taxonomic papers refer to Oshima (1912) but do not provide further insights in the form of a redescription of the species or new material (e.g., Horikawa 1927; Wang & Wang 1956; Lue et al. 1988); see also relevant comments in Ota (1989 b). In describing G. mindorensis, Taylor (1919) compared the new species with G. kikuchii and G. monarchus. According to Taylor (1919), G. mindorensis differs from the former species by its longer limbs, more preanofemoral pores and by lacking webbing between toes in both sexes; G. mindorensis differs from G. m o n a rc h u s by a relatively larger ear opening, a larger eye diameter, a smaller interorbital distance, more flattened dorsal tubercles, more ventrals and preanofemoral pores, and a distinctly different dorsal pattern (see also Taylor 1922 a). However, Wang (1962) pointed to the similarity of G. kikuchii with G. monarchus and G. mindorensis. Brown & Alcala (1978) redescribed G. mindorensis based on new specimens, but only provided thorough comparisons with G. monarchus, but not with G. kikuchii. Based on the new variation in size and scalation reported in the redescription provided by Brown & Alcala (1978), G. mindorensis was in fact no longer distinguishable from G. kikuchii based on the data on the latter species provided by Taylor (1919), as is also obvious from Table 1. Crombie (in Bauer 1994) thus had doubts on the distinct status of G. mindorensis and stated ���probably a junior synonym of Gekko kikuchii.��� Finally, Ferner et al. (2001) and Kluge (2001) listed G. mindorensis as a synonym of G. kikuchii. However, there might be differences in pattern between G. kikuchii and G. mindorensis. Figures of different specimens of G. kikuchii in Ota (1989 b, 1991), Lue et al. (1999), Shang (2001), and Henkel & Schmidt (2003) show a brown ground coloration with a nearly identical dorsal pattern, consisting of two parallel dark brown pairs of blotches. In contrast, the ground coloration of G. mindorensis seems to be gray, with transverse rows (see Gaulke 2003: fig. 8) or rows dissolved into four to six blotches each (see R��sler et al. 2006: fig. 13). In addition, the light dorsal blotches in G. mindorensis appear to be more distinct than in G. kikuchii. We must add that the detailed figures provided by Okada (1936: fig. 4) most probably belong to a Hemidactylus species (maybe H. stejnegeri), but certainly not G. kikuchii. The synonymy of G. mindorensis with G. kikuchii is also contradicted on zoogeographic grounds. The amphibian and lizard fauna of Lanyu Island shows, according to Okada (1936) and Ota et al. (1988), greater affinities to Taiwan than to the Philippines, and G. mindorensis does not occur on the northern Philippine Islands (e.g., Luzon, Batanes) that are closest to Taiwan (Brown & Alcala 1970, 1978; Alcala 1986; Gaulke & Altenbach 2006). Kuntz & Ming (1970) listed G. kikuchii for Penghu Lieh-tao (Pescador Island), but the species is endemic to Lanyu Island (���Botel Tobago ���) (see Ota 1989 b). Further taxonomic research is required on more specimens, especially of G. kikuchii, to clarify whether G. mindorensis is a junior synonym of G. kikuchii, or whether two closely related cryptic species are involved., Published as part of R��sler, Herbert, Bauer, Aaron M., Heinicke, Matthew P., Greenbaum, Eli, Jackman, Todd, Nguyen, Truong Quang & Ziegler, Thomas, 2011, Phylogeny, taxonomy, and zoogeography of the genus Gekko Laurenti, 1768 with the revalidation of G. reevesii Gray, 1831 (Sauria: Gekkonidae), pp. 1-50 in Zootaxa 2989 on page 17, DOI: 10.5281/zenodo.278393, {"references":["Linkem, C. W., Siler, C. D., Diesmos, A. C., Sy, E. & Brown, R. M. (2010) A new species of Gekko (Squamata: Gekkonidae) from central Luzon Island, Philippines. Zootaxa, 2396, 37 - 49","Rosler, H., Siler, C. D., Brown, R. M., Demegillo, AD. & Gaulke, M. (2006) Gekko ernstkelleri sp. n. - a new gekkonid lizard from Panay Island, Philippines. Salamandra, 42, 197 - 211.","Oshima, M. (1912) Description of a new gecko from Botel Tobago Island. Philippine Journal of Science, 7, 241 - 242.","Taylor, E. H. (1919) New or rare Philippine reptiles. The Philippine Journal of Science, 14, 105 - 125.","Schlegel, H. (1836) Le Platydactyle Monarque. Platydactylus Monarchus. In: Dumeril, A. M. C. & Bibron, G. (1836), Erpetologie Generale ou Histoire Naturelle Complete des Reptiles. Vol. 3. Roret, Paris, pp. 335 - 336.","Taylor, E. H. (1925) Additions to the herpetological fauna of the Philippine Islands. IV. The Philippine Journal of Science, 26, 97 - 111.","Horikawa, Y. (1927) Animals collected on the island of Kotosho by Mr. Sasaki. Transactions of the Natural History Society of Formosa, 17, 295 - 296.","Wang, C. S. & Wang, Y. M. (1956) The reptiles of Taiwan. Quarterly Journal of the Taiwan Museum, 9, 1 - 86.","Ota, H. (1989 b) A review of the geckos (Lacertilia: Reptilia) of the Ryukyu Archipelago and Taiwan. In: Matsui, M. Hikida, T. & Goris, R. C. (eds.), Current Herpetology in East Asia, Herpetological Society of Japan, Kyoto, pp. 222 - 261.","Taylor, E. H. (1922 a) The Lizards of the Philippine Islands. Philippine Bureau of Science, Manila, 269 pp., 23 pls.","Wang, C. S. (1962) The reptiles of Botel-Tobago. Quarterly Journal of the Taiwan Museum, 5, 141 - 190.","Brown, W. C. & Alcala, A. C. (1978) Philippine lizards of the family Gekkonidae. Silliman University Natural Science Monographs, 1, 1 - 146.","Bauer, A. M. (1994) Gekkonidae (Reptilia, Sauria), Part I Australia and Oceania. Das Tierreich 109 (Part). Walter de Gruyter, Berlin, XII + 306 pp.","Ferner, J. W., Brown, R. M., Sison, R. V. & Kennedy, R. S. (2001) The amphibians and reptiles of Panay Island, Philippines. Asiatic Herpetological Research, 9, 34 - 70.","Kluge, A. G. (2001) Gekkotan lizard taxonomy. Hamadryad, 26, 1 - 209.","Lue, K. Y., Tu, M. C. & Shang, G. S. (1999) A Field Guide to Amphibians and Reptiles in Taiwan. Society for Wildlife and Nature, Taipei, 343 pp.","Shang, G. (2001) Natural Portraits of Lizards of Taiwan [in Chinese]. Bigtrees Press, Taipei, 173 pp.","Henkel, F. W. & Schmidt, W. (2003) Praxis Ratgeber Geckos. Edition Chimaira, Frankfurt am Main, 159 pp.","Gaulke, M. (2003) Ubersicht der philippinischen Gecko-Fauna. Gekkota, 4, 3 - 24.","Okada, Y. (1936) Studies on the lizards of Japan. Contribution. I. Gekkonidae. Science Reports of the Tokyo Bunrika Daigaku Section B, 42, 233 - 289.","Ota, H., Crombie, R. I., Lue, K. - Y. & Chen, S. - H. (1988) A record of Gekko hokouensis (Reptilia: Sauria) from Lanyu Island, Taiwan, with notes on its variation. Journal of Taiwan Museum, 41, 7 - 14.","Brown, W. C. & Alcala, A. C. (1970) The zoogeography of the herpetofauna of the Philippine Islands, a fringing archipelago. Proceedings of the California Academy of Sciences, 4 (38), 105 - 130.","Alcala, A. C. (1986) Guide to Philippine flora and fauna. X. Reptiles. JMC Press Incorporated, Quezon, xiv + 195 pp.","Gaulke, M. & Altenbach, A. V. (2006) Waldgeckos im Haus, zur Geckofauna von Panay. Reptilia, 11 (5), 64 - 71.","Kuntz, R. E. & Ming, D. Z. (1970) Vertebrates of Taiwan taken for parasitological and biomedical studies by U. S. Naval Medical Research Unit No. 2, Taipei, Taiwan, Republic of China. Quarterly Journal of the Taiwan Museum, 23, 1 - 37."]}

Greenbaum, Eli, Villanueva, Cesar O., Kusamba, Chifundera, Aristote, Mwenebatu M., Branch, William R. (2011): A molecular phylogeny of Equatorial African Lacertidae, with the description of a new genus and species from eastern Democratic Republic of the Congo. Zoological Journal of the Linnean Society 163 (3): 913-942, DOI: 10.1111/j.1096-3642.2011.00732.x, URL: http://dx.doi.org/10.1111/j.1096-3642.2011.00732.x

Greenbaum, Eli, Bauer, Aaron M., Jackman, Todd R., Vences, Miguel, Glaw, Frank (2007): A phylogeny of the enigmatic Madagascan geckos of the genus Uroplatus (Squamata: Gekkonidae). Zootaxa 1493 (1): 41-51, DOI: 10.11646/zootaxa.1493.1.2, URL: https://biotaxa.org/Zootaxa/article/view/zootaxa.1493.1.2

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