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Redescription of external morphology Based on a re-examination of the holotype and on the examination of the recently collected specimens RBINS 20301–20302, shown on Figures 3–6. Raw measurements and scale counts of the three specimens are provided in Table 2. Snout-vent length to 76 mm; total length> 229 mm (holotype). Tail 2.5 times SVL (based on RBINS 20302, the only specimen with a complete, original tail; the tail tip of RBINS 20301 is missing and is not healed). Body moderately depressed. Head distinct from neck, narrow (HL/HW ratio 1.8–2.1), long (HL/SVL ratio 0.25–0.30, proportionally longest in the subadult), depressed (HH/HL ratio 0.41–0.49, least depressed in the subadult). Head covered with symmetric plates. Rostral well visible in dorsal view. Frontonasal rounded anteriorly, slightly wider than long. A line of small tubercles along the posterior borders of the internasals, frontonasal and prefrontals (also along the lateral borders of the prefrontals in the subadult), “en forme de perles” (i.e., pearl-shaped) as described by Monard (1940). Suture between internasals subequal to suture between prefrontals (respectively 0.8 and 0.9 mm in RBINS 16301). Pupil round. Lower eyelid scaly. Canthus rostralis rounded. Lores near-vertical. Nostril opening in contact with 1 st supralabial, the anterior nasal and the two posterior, superposed, nasals. Four supraorbital scales, the anterior- and posteriormost small, separated from the supraciliaries by a continuous row of granular scales. In the holotype the anterior- and posteriormost supraoculars are entire, while in RBINS 20301 the anterior supraoculars are divided into two (left) or three (right) fragments and the posterior ones into three (left) or two (right) fragments. In RBINS 20302 the anterior supraoculars are divided into two fragments on each side, and the posterior supraoculars are unfragmented. Tympanic opening large, rounded, surrounded by smooth scales (i.e., no auricular denticulation), including a crescentic scale bordering the antero-dorsal limit of the tympanum. Seven or eight supralabials; one of them much enlarged and bordering the orbit, separating the five anterior supralabials (sometimes four, as on the left side of the holotype) from the two posterior ones. Frontal hexagonal, surrounded anteriorly by the two prefrontals, laterally by three supraoculars on each side, and posteriorly by the two frontoparietals. Anterior border of frontal pointing forward. Frontal narrow posteriorly. Length of frontal slightly smaller than distance between frontal and snout tip (respectively 4.1 and 5.1 mm in RBINS 16301). Frontoparietals pentagonal. Parietals large, nearly as long as frontal, separated by the interparietal scale and a small occipital. Pineal eye visible through the interparietal scale. Scales on the upper surface of the head smooth, except the pearl-shaped tubercles. Temporal area covered by an elongate, thin temporal plate along the parietal, and by smaller to granular scales. Six (in a single case seven) infralabials. Mental followed by four pairs of sublinguals, the first three in contact on the midline, the fourth pair separated from each other. Sublinguals progressively increasing in size posteriorly. Gular collar present and distinct, ventrally including five distinctly enlarged scales. The right profile and ventral view of the head, which had never been illustrated for the holotype so far, are shown on Figure 3. Mediodorsal scale rows not widened. Dorsal scales granular, in 67–70 longitudinal rows at midbody, those on the lower flanks nearly smooth, those on upper flanks and dorsum with a median, single, longitudinal keel. About 40 dorsal scales between legs. Six longitudinal rows of parallelepipedal or trapezoidal, widened, smooth, ventral scales; the two medioventral rows narrower than the lateral ones. Between the gular collar and the line of porebearing scales, 27 or 28 transversal rows of ventrals. A distinctly enlarged preanal plate, bordered laterally by a row of a few small preanals on each side. Femoral pores in a continuous row of 17–20. The left and right series of pores of RBINS 20301 and RBINS 20302 are separated by respectively three and two poreless scales (two in the holotype according to the Figure 3 in the original description, but actually three according to our observations). Subdigital lamellae of fingers and toes with two keels each. First finger shortest. Second finger longer than 5 th. Third and 4 th fingers longest, of subequal length. RBINS 20301 and RBINS 20302 both show 16/16 subdigital lamellae under the 4 th finger. When the leg is stretched alongside the body it extends anteriorly beyond the gular collar. Length of the feet comparable to head length. Toes without lateral denticulate fringes. First toe shortest. Second and 5 th toe of subequal length, shorter than the 3 rd and much shorter than the 4th. Subdigital lamellae under 4 th toe 23 to 26 (23 in the holotype, 25 or 26 in the two other specimens). Supracaudals much larger than dorsal scales. All supracaudal scales rectangular, presenting a strong median keel, each keel bearing a small tubercle at its posterior extremity. Subcaudals at the base of the tail smooth and rounded posteriorly, quickly becoming rectangular with a medial strong keel, but without a terminal tubercle on the keel. The tail of the holotype is broken. Monard (1940) mentioned 87 subcaudals, but the tail tip shows a uniform color contrasting with the anterior part of the tail, indicating that it is regenerated. The tail of the subadult RBINS 20302 is original, and shows 143 subcaudals. Coloration in life. Based on original description, RBINS 20301–20302 and Figures 3–7. The dorsal surface of the head is uniformly dark brown. The ventral surface of the head, the body and of the base of the tail is uniformly white. The background color of the sides of the head above the mouth line is dark brown, progressively darkening posteriorly to become black as the background color of the flanks and the first half of the dorsum. There is an alignment of white spots on the upper and lower lips and another on the temporal area, in continuity with four similar, irregular alignments of white spots along the whole length of the flanks. Four continuous, parallel white dorsal stripes extend from just behind the parietals till the posterior part of the dorsum where they fade and disappear (Figures 4–5 and 7). In the subadult these white stripes are irregular and discontinuous (Figure 6). From half-length of the trunk, the dorsal background color turns to reddish-brown (similar to the color of laterite), continuing to the tail. The upper surfaces of the proximal parts of the arms are black with white spots, turning to reddish-brown with irregular lighter spots on the distal parts of the arms. The upper surfaces of the legs show a reddish-brown background color with irregular lighter spots and markings. The lower surface of the tail progressively reaches posteriorly the uniform reddish-brown color uniformly covering the dorsal and lateral surfaces of the tail until its tip. The lower surfaces of the arms and legs are grayish-white, the palms are reddish-brown. Cranial osteology. Based on subadult male RBINS 20302. Snout and palatomaxillary bones The premaxilla is a single, dentigerous bone (Figure 8A–D). The alveolar shelf carries probably seven teeth (counting the sockets) and juts out caudad as two triangular processes touching the maxilla. There is a narrow, tapering, dorsocaudally directed nasal process that is almost as long as the nasals themselves and wedges the tapering tip between the nasals. The nasals are paired, almost flat elements that form a straight suture between themselves and an interdigitating suture with the frontal (Figure 8A, D). The anterior tips are pointed and diverge from each other to make room for the intercalating premaxillary nasal process. The nasals are the widest at about the mid-length where they project out laterally into a shark fin-shaped, anteriorly embayed protuberance slightly overlapping the maxillary facial process. The maxilla has a high facial process with a triangular posterodorsal process reaching the frontal and another small posterolateral process below it touching the prefrontal (Figure 8A–B). There is a semilunar embayment between these two processes. The alveolar border bears 15 pleurodont teeth. Right above the teeth, a palatal shelf medially expands (Figure 8C). The premaxillary process is short and slightly upturned and bifurcated into an anterolateral and an anteromedial process, as is common in many lacertids (Villa & Delfino 2019). The palatine is an edentulous, ventrally concave bone that overlaps the palatal shelf of maxilla laterally with a maxillary process (Figure 8C). This process also establishes contact with the prefrontal dorsolaterally. The squarish vomerine process overlaps the palatine process of the vomer. Between these two processes, namely the maxillary and the palatine, there is an anterior embayment and a ventral concavity corresponding to choana. Posteriorly the palatine ends in three little triangular protuberances of which the medial two overlap the pterygoid, and the lateral one laterally articulates to that bone. The palatines do not contact each other medially. The pterygoid is an edentulous, triradiate bone (Figure 8C). The anteromedial process of the pterygoid, which articulates with the palatine, is longer than the anterolateral process of the same that articulates with the ectopterygoid. The quadrate process of the pterygoid is slender, lateromedially compressed and is directed posterolateral. The dorsolateral surface of the quadrate process bears a small facet for the epipterygoid. The medial surface of this process bears a longitudinal groove for the attachment of the pterygomandibularis muscle (Daza et al. 2011; Das & Pramanick 2019). The ectopterygoid articulates medially to the ectopterygoid articular facet on the pterygoid anterolateral process (Figure 8C). Anteriorly it overlaps the palatal shelf of the maxilla and just contacts the palatine. The vomer forms the casing of the vomeronasal organ together with the septomaxilla (Figure 8C). Vomers contact each other medially except from their posterior medial margins. Anteriorly, they contact the maxilla. Posteriorly vomers touch the palatine. Anterolaterally vomers are narrow to create an opening for the vomeronasal fenestra. Behind this, vomers expand laterally, only to taper laterally again to for choana. The septomaxillae form the dorsal encasing of the vomeronasal organ (Figure 8A, D). The septomaxillae are concave ventrally. Their medial edges are turned dorsad. Anteroventrally the septomaxilla ends with two small, pointed processes. Except a small protuberance, there is no clear posteromedial process as in some Palaearctic lacertids (Villa & Delfino 2019). Chondrocranial braincase bones The braincase bones show some degree of fusion, although sutures are detectable, at least partially, between the otic capsule elements and the ventral braincase elements, namely the parabasisphenoid and the basioccipital. Prootic is a prominent element housing cochlea, anterior and horizontal (partly) semicircular canals and their ampullae (Figure 8B–C). The anterior semicircular canal forms a prominent bulge on the anterolateral surface, just behind the crista alaris. In this species, the crista alaris is a narrow semilunar projection immediately rostrad and somewhat medial to the anterior semicircular canal bulge (Figure 8B). Immediately below this bulge, begins a rather weakly developed crista prootica that runs caudad from this point. Ventrad to the crista prootica, the anterior margin of the prootic is embayed by the incisura prootica (Figure 8B). Ventrad to the incisura prootica notch, prootic projects rostrad into an obtusely triangular anterior inferior process. The bulge along the horizontal semicircular canal continues posteriorly as a prootic process to reach the anterior surface of the paroccipital process. On the medial surface of the prootic, there are two auditory nerve foramina. The prootic articulates (in this specimen, fused) with the otoccipital along the former’s posterior lateral margin. The paired otoccipital forms the occipital condyle with the basioccipital (Figure 8A, E). The otoccipitals are composite (of opisthotic and exoccipital), hollow, bulbous bones that encase the internal ear. Dorsally the otoccipitals project out into a prominent, posterolaterally directed, axe head-shaped (in posterior view) paroccipital process that is approximately one-third the length of the quadrate. The supratemporal attaches to the anterior surface of the lateral end of this process. Ventrad to the supratemporal facet, there is a facet for the quadrate. Dorsad to the basal tubera, the otoccipital has a very prominent, deep embayment of the recessus scalae tympani which is bordered anterodorsally by the crista interfenestralis. Posterodorsal to the lateral opening of the recessus scalae tympani, there is a vagus foramen. The position of the hypoglossal foramina could not be detected clearly in the scan. One of the two single elements of the ventral braincase is the parabasisphenoid (Figure 8C). The bone has an elongated, narrow parasphenoid rostrum. Behind this parasphenoid element, the basisphenoid begins to expand. On both sides of the base of the parasphenoid rostrum, a truncated looking trabecula cranii is present. Between the trabeculae, on the dorsal surface is situated the sella turcica.Two very small internal carotid foramina open within the sella turcica. Caudad to the sella turcica, the crista sellaris is present transversely. Two short, anteroventrally directed basipterygoid processes project out from the ventrolateral base of the basisphenoid (Figure 8C). These processes expand at their end. Dorsally and medially, the anterior vidian foramen pierces the basipterygoid process. The basioccipital is the ventral element of the braincase participating in the formation of the occipital condyle (Figure 8A, C, E). The bone is hexagonal, dorsally concave and in this specimen, partly fused with the otoccipital, the prootic and the parabasisphenoid. Ventrad to the lateral opening of the recessus scalae tympani, the basioccipital has a small protuberance, the basal tubera. The basioccipital forms the floor of the recessus scalae tympani. The supraoccipital consists of a dorsomedian roof for the foramen magnum and two expanded and hollowed lateroventral wings partly encasing the inner ear (Figure 8A). In this species the processus ascendens is a very small, anteriorly truncated protuberance which does not reach the parietal (Figure 8E). This bone articulates with the prootic and the otoccipital. Dermal skull roofing bones The frontal bone is a single (fused in this specimen, condition in hatchlings is not known), elongated, skull roof element, being almost twice as long as the parietal in this species (Figure 8A, D). The frontal is wide at the rostral end and very wide at the caudal end and relatively narrower in the middle. A few digitiform processes from the frontals form interdigitating sutures with the nasals and the maxillae. The frontal articulates with the prefrontal along the anterior one-third of the former’s lateral margin. On the ventral surface of the frontal, along its outer margin, there is a crest, the crista cranii. The caudal end of the frontal expands markedly into two posterolateral processes with a squarish end and articulates with the parietal and the postfrontal. This particular specimen does not show any rugosity on the dorsal surface of the frontal. The parietal is a squarish, short, wide skull roof bone (Figure 8A–B). There are no anterolateral processes. However, there is a narrow, tapering posterolateral process on each side that articulates with the supratemporal and is just separated from the squamosal. These processes do not reach the paroccipital process of the otoccipital. There is a pineal foramen (Ledesma & Scarpetta 2018) piercing the parietal. There is a fossa parietalis (Oelrich 1956) at the midpoint of the embayed posterior margin of the parietal, though the processus ascendens from the supraoccipital is very weakly developed in this species and does not reach the fossa. Circumorbital bones The prefrontals are cavernous (in anteromedial view) bones that meet the frontals laterally, the maxillary facial process anteriorly and anterodorsally, the palatines ventrally and the lacrimals ventrolaterally (Figure 8A–B). The ventrolateral border of the orbitonasal flange of the parietal has a deep lacrimal notch. The orbitonasal flange projects ventrally into a posteroventral process medial to the lacrimal notch. Dorsally, the flange projects into a caudally directed process along the crista cranii. There is rather weak palpebral crest. The jugals are paired bones consisting of an anterior and a posterodorsal process, the former being more robust, especially close to the junction between the two, than the latter (Figure 8A–C). The anterior process tapers anteriorly and meets the lacrimal.This process articulates with the frontal anteroventrally and the ectopterygoid ventromedially. There is a faintly developed, caudally directed protuberance, the quadratojugal process, at the junction between the anterior and the posterodorsal processes. The slender posterodorsal process touches the postorbital. The lacrimal is a small sliver of a bone overlying the junction of the prefrontal, jugal and the maxilla (the prefrontal is just separated from the jugal), immediately caudad to the prefrontal lacrimal notch (Figure 8B). The postfrontal is a triradiate bone with a very slender anterolateral process articulating to the posterior lateral margin of the frontal, a weakly developed, triangular protuberance articulating to the anterodorsal margin of the postorbital and a robust, dagger-like posterior process wedged between the frontal and the parietal medially and the postorbital laterally (Figure 8A–B). The postorbital is situated below the postfrontal (Figure 8A–B). It consists of a short, triangular anteroventral process that contacts the posterodorsal process of the jugal and a longer posterior process touching the postfrontal and the squamosal, but the resolution of the scan did not permit determination of the exact end point of this process. In this specimen, the postfrontal and the postorbital do not extend more than one-third the length of the squamosal and thus, leave a prominent upper temporal fenestra open. There are prominent supraorbital ossifications above and scleral ring within the orbit (Figure 8A–B). Suspensorial and palatoquadrate derived bones The squamosals are a pair of J-shaped suspensorial bones (Figure 8A–B). Rostrally the squamosal is tapered and articulates medially to the postorbital. Posteriorly the bone curves ventrad and contacts the supratemporal dorsomedially and the quadrate ventrally. The ventrally curved caudal end of this bone is squarish. The supratemporals are small, curved bones with a tapered rostral and widened caudoventral end (Figure 8A–B). The caudoventral end bears an articulatory facet for the quadrate. Anterolaterally and anteromedially the supratemporal articulates with the squamosal and the posterolateral process of the parietal, respectively. The epipterygoid is a rather nondescript rod-like bone that articulates to the pterygoid right behind t, Published as part of Pauwels, Olivier S. G., Das, Sunandan, Camara, Lewei Boyo, Chirio, Laurent, Doumbia, Joseph, D'Acoz, Cédric D'Udekem, Dufour, Sylvain, Margraf, Nicolas & Sonet, Gontran, 2023, Rediscovery, range extension, phylogenetic relationships and updated diagnosis of the Ornate Long-tailed Lizard Latastia ornata Monard, 1940 (Squamata: Lacertidae), pp. 501-524 in Zootaxa 5296 (4) on pages 507-517, DOI: 10.11646/zootaxa.5296.4.1, http://zenodo.org/record/7984314, {"references":["Monard, A. (1940) Resultats de la mission scientifique du Dr. Monard en Guinee portugaise, 1937 - 1938. VIII. Reptiles. Arquivos do Museu Bocage, 11, 147 - 182, pl.","Villa, A. & Delfino, M. (2019) A comparative atlas of the skull osteology of European lizards (Reptilia: Squamata). Zoological Journal of the Linnean Society, 187 (3), 829 - 928. https: // doi. org / 10.1093 / zoolinnean / zlz 035","Daza, J. D., Diogo, R., Johnston, P. & Abdala, V. (2011) Jaw adductor muscles across lepidosaurs: a reappraisal. The Anatomical Record, 294 (10), 1765 - 1782. https: // doi. org / 10.1002 / ar. 21467","Das, S. & Pramanick, K. (2019) Comparative anatomy and homology of jaw adductor muscles of some South Asian colubroid snakes (Serpentes: Colubroidea). Vertebrate Zoology, 69 (1), 93 - 102. https: // doi. org / 10.26049 / VZ 69 - 1 - 2019 - 04","Ledesma, D. T. & Scarpetta, S. G. (2018) The skull of the gerrhonotine lizard Elgaria panamintina (Squamata: Anguidae). PloS ONE, 13 (6), e 0199584. https: // doi. org / 10.1371 / journal. pone. 0199584","Oelrich, T. M. (1956) The anatomy of the head of Ctenosaura pectinata (Iguanidae). University of Michigan Museum of Zoology Miscellaneous Publications, 94, 1 - 122.","Good, D. A. (1987) A phylogenetic analysis of cranial osteology in the gerrhonotine lizards. Journal of Herpetology, 21, 285 - 297. https: // doi. org / 10.2307 / 1563970"]}

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."]}

David, Patrick, Campbell, Patrick D., Deuti, Kaushik, Hauser, Sjon, Luu, Vinh Quang, Nguyen, Truong Quang, Orlov, Nikolai, Pauwels, Olivier S. G., Scheinberg, Lau- Ren, Sethy, Priyadarsi Girija Sankar, Smits, Ton, Teynié, Alexandre, Vogel, Gernot (2022): On the distribution of Gonyosoma prasinum (Blyth, 1854) and Gonyosoma coeruleum Liu, Hou, Ye Htet Lwin, Wang & Rao, 2021, with a note on the status of Gonyosoma gramineum Günther, 1864 (Squamata: Serpentes: Colubridae). Zootaxa 5154 (2): 175-197, DOI: https://doi.org/10.11646/zootaxa.5154.2.4

Gehyra wongchan sp. nov. (Figures 1–6) Gehyra cf. angusticaudata –– Meesook et al. 2021: 322. Holotype. CUMZ-R-2598 (field no. MS 740), adult male caught on 20 October 2020 in Tham (= Cave) Khao Chan (14°58’35.2”N, 101°18’11.2”E), Tha Luang District, Lopburi Province, central Thailand, by W. Meesook, N. Donbundit and M. Sumontha. Paratypes (4). CUMZ-R-2599 and CUMZ-R-2613 (field nos. MS 261 and MS 262, respectively), adult male and juvenile female collected on 26 April 2008 at Wat Khao Wong (ca. 14°57’51.5”N 100°41’58.3”E), Kok Samrong District, Lopburi Province, by K. Kunya and M. Sumontha. CUMZ-R-2611 and CUMZ-R-2612 (field nos. MS 738 and MS 739, respectively), juvenile male and adult female collected on 26 March 2021; same locality and collectors as holotype. Diagnosis. Gehyra wongchan sp. nov. can be distinguished from all other congeneric species by the combination of its maximal known SVL of 52.4 mm, 8–10 supralabials, 76–80 dorsal and 48–50 ventral scale rows around midbody, absence of skin folds on limbs, 17 or 18 preanofemoral pores in males in a continuous series extending to mid-length of femur (pores absent in females), tail not- to moderately widened behind vent in adults, a single row of widened subcaudals (about 1/3 of the width of tail in its anterior part, progressively occupying the whole width of the tail towards the tip), digits and toes unwebbed, 7 or 8 divided subdigital lamellae on 4th toe, and a dorsal pattern with white spots as large or larger than adjacent crescent-shaped black markings on a beige to light brown background. Description of holotype. Adult male (Figures 1–3 and Table 1). SVL 45.2 mm. Head long (HeadL/SVL 0.32), relatively broad (HeadW/HeadL ratio 0.63), somewhat depressed (HeadD/HeadL ratio 0.34), poorly distinct from neck. Lores and interorbital region slightly inflated; prefrontal region concave; canthus rostralis smoothly rounded. Snout moderate (SnOrb/HeadL ratio 0.33), less than 1.5 times eye diameter (OrbD/SnOrb ratio 0.75); scales on rostrum, lores, top of head and occiput small, granular, lacking enlarged tubercles; scales on snout much larger than those on interorbital region. Eye relatively large (OrbD/HeadL ratio 0.25). Pupil vertical with crenelated margins. Supraciliaries short. Ear opening oval, moderate (EarL/HeadL ratio 0.08); eye to ear distance slightly shorter than diameter of orbit (OrbEar/OrbD ratio 0.89). Rostral more than two times wider (2.7 mm) than deep (1.2 mm). No rostral groove present; two much enlarged supranasals separated by two small, antero-posteriorly aligned, granular scales. Rostral in contact with supralabial I, nostrils, supranasals and a granular scale separating the supranasals. Nostrils oval, each surrounded by rostral, supranasal, two postnasals and first supralabial. Interorbital scale rows across narrowest point of frontal 13. Mental triangular, wider (2.6 mm) than deep (1.8 mm), much deeper than infralabials; mental in contact with four scales: laterally with first infralabials and posteriorly with a pair of greatly enlarged and elongate inner postmentals meeting behind the mental. Each postmental bordered anteriorly by first infralabial, medially by mental, laterally by an enlarged chin shield (outer postmental), and posteriorly by undifferentiated granular gular scales. Including the two postmentals, there are two pairs of enlarged chin shields. Enlarged supralabials to midpoint of orbit 8 (left and right); supralabials to angle of jaws 10 (left and right); enlarged infralabials 10 (left and right). Gular scales small, subimbricate, grading posteriorly into slightly larger, subimbricate pectoral scales, which grade posteriorly into larger, subimbricate ventrals. Body robust, trunk relatively long (TrunkL/SVL ratio 0.63), dorsoventrally depressed in cross-section, with poorly distinct ventrolateral folds. Dorsal scales small, granular to subimbricate, without tubercles. Ventral scales slightly larger than dorsals. Midbody scale rows across belly to ventrolateral folds 48. No enlarged, precloacal scales. Seventeen pore-bearing precloacal scales, in a continuous row, extending to mid-length of the femur. Scales on palm and sole smooth, flat, rounded. Scales on dorsal aspects of hind limbs homogeneous, granular. Scales on dorsal surface of forelimb homogeneous, granular, flat to slightly conical. Fore- and hind limbs moderately long, stout; forearm and tibia moderately long (FAL/SVL ratio 0.13; TibL/SVL 0.17). No skin folds (sensu Oliver et al. 2016) on fore and hindlimbs. Digits relatively short; digit I, both manus and pes, clawless; all remaining digits strongly clawed; distal portions of digits II-V strongly curved, arising from distal portion of expanded subdigital pad. Scansors beneath each toe divided; scansors 5-6-6-7-7 (left manus), 6-7-7-6-6 (right manus), 6-7-7-8-8 (left pes), 6-7-8-8-7 (right pes). Relative length of digits of manus: IV>V=III>II>I. No webbing between digits or toes. Tail original, depressed, slightly longer than head and body (TailL/SVL ratio 1.10); dorsal surface of tail covered with small, squarish, juxtaposed granules forming more or less regular transverse rows. Median row of strongly enlarged subcaudal plates extending about 1/3 across the width of tail in its anterior part, progressively occupying the whole width of the tail towards the tip (Figure 2). A single, moderate, post-cloacal tubercle on each side of tail base. Coloration in life. Background color of dorsal surface of head, dorsum, and dorsal surfaces of members and tail beige (Figure 1). Poorly contrasted and incomplete preorbital darkish stripe. Supraorbital region bluish. A few irregular black and white spots on the dorsal surface of head. Discontinuous black nuchal collar from one orbit to the other, posteriorly bordered on nape by four white spots. A pair of large white paravertebral spots on neck each anteriorly bordered by a black crescent, each white spot of a diameter of about seven dorsal granular scales. Five similar pairs of large white paravertebral spots between limb insertions, not symmetrically arranged. A few white small spots on lower flanks. A black crescent above sacrum. Dorsal surface of tail showing eight blackish bands, each bordered posteriorly by a whitish thinner band. No bands or spots on the dorsal surfaces of limbs, hands and feet. Throat and belly beige without spots, darkening towards posterior abdomen; pore-bearing scales lighter. Lower surface of limbs, hands, feet and tail brown. In preservative the general color darkens, and the dorsal pattern becomes less contrasting (Figures 2 and 3). Variation. Main morphometric and meristic characters of the type series are provided in Table 1. Morphological characters and color pattern of the paratypes agree in most respects with the holotype. The females lack preanofemoral pores. As with the holotype, all paratypes have an original tail. Females show slightly shorter original tails than males (TailL/SVL ratio 0.87–0.96 vs. 1.09–1.10). The white spots on dorsum are proportionally smaller in juveniles than in adults (Figures 1, 2, 4 and 6). Distribution and natural history. Wat Khao Wong is located about 65 km W of Tham Khao Chan (Figure 14). Both sites lie on limestone hills surrounded by cultivated areas. These hills belong to the “ Saraburi Group Limestones” in the south-west margin of the Khorat Plateau (Ponta et al. 2013; Warren et al. 2014). Nothing is known about the diet or the reproduction of the new species. The holotype was found while it was foraging on a tree near the entrance of Khao Chan Cave (Figures 1 and 15). At the type-locality, individuals were observed inside the cave, on limestone and trees near the cave entrance, and on nearby buildings. The species is locally common. Within Khao Chan Cave Gehyra wongchan sp. nov. individuals were observed in syntopy with Dixonius siamensis Boulenger, Gekko pradapdao and the ubiquitous G. gecko (Linnaeus), Hemidactylus frenatus Duméril & Bibron and H. platyurus (Schneider) (Gekkonidae). We found several individuals of Lycodon capucinus (Boie) in and around the cave, and numerous shed skins of L. davisonii (Blanford) (Colubridae) within the cave. Etymology. The specific epithet wongchan is a name in apposition, invariable, based on the contraction of the localities of the paratypes (Wat Khao Wong) and of the holotype (Tham Khao Chan). In Thai wongchan also means the Moon, in reference to the typical crescent-shaped marks on the nape and dorsum of the new species. We suggest the following common names: จิ้งจกหินวงจันทร์ (Djing-djok-hin wongchan; Thai); Lunulate four-clawed gecko (English), and Gehyra lunulée (French). Comparison to other species. Based on its scalation and dorsal pattern, Gehyra wongchan sp. nov. is readily distinguished from the four other Gehyra species found in mainland Southeast Asia; their main diagnostic characters are compared in Table 2. Gehyra wongchan sp. nov. differs from the Thai endemic G. angusticaudata by its much higher VentR (48–50 vs. 35), much lower PrePo number (17 or 18 extending to mid-length of the femur vs. 31–37 extending the whole length of the femur), absence of webbing on fingers and toes (vs. basal webbing), and its dorsal pattern (large white spots adjacent to black crescents vs. small and nearly indistinct white spots among small black spots, see Figures 7 and 8). The holotype of Gehyra fehlmanni (Figure 9) was collected at ‘‘ 4 km. NW [of] Kanchanaburi, Kanchanaburi Province’’, western Thailand. In the original description, Taylor (1962: 223) noted that the scales on dorsal and lateral surfaces of the holotype were irregular, ‘‘suggesting the possibility that the entire tail has been regenerated’’, but also that the subcaudal scales are widened, ‘‘appearing to be normal and not reproduced’’. The latter fact, combined with the presence on the dorsal surface of the tail of a color pattern pursuing the one seen on the dorsum, and of a lateral fringe of small denticulate scales, seems to indicate that the tail is original. Figure 10 is the first published image of the single paratype of Gehyra fehlmanni, collected at ‘‘Tonka Harbour Tin Mine, Ronpibon, Nakhon Si Thammarat Province’’, southern peninsular Thailand, i.e., at approximately 500 airline km south of the type-locality. Its tail has been lost and was not available to Edward Taylor for the species’ description. Gehyra wongchan sp. nov. can be distinguished from G. fehlmanni by its higher VentR (48–50 vs. 42), lower PrePo number (17 or 18 vs. 22), absence of webbing on fingers and toes (vs. basal webbing), and by its dorsal pattern (transversally elongate black spots with smaller white spots in G. fehlmanni, see Figures 9–11). If the tail of the holotype of Gehyra fehlmanni is indeed original, it is shorter than the tails of males of G. wongchan sp. nov. (TailL/ SVL ratio 0.89 vs. 1.09–1.10). A population of Gehyra cf. fehlmanni has been reported by Grismer et al. (2007) from eastern Cardamom Mountains in Cambodia, exhibiting a dorsal color pattern similar to that of G. fehlmanni but differing in several scalation characters. This Cambodian population differs from Gehyra wongchan sp. nov. by its dorsal pattern (in particular by its transversally elongate black spots, much larger than its white spots) and its 37 PrePo (vs. 17 or 18). Gehyra wongchan sp. nov. can be differentiated from G. lacerata by its lower SL number (8–10 vs. 12), lower PrePo number (17 or 18 vs. 20), its transversely enlarged subcaudals (vs. not enlarged), a tail not- to moderately widened posterior to vent (vs. strongly widened), and by its dorsal pattern (large white spots adjacent to black crescents vs. large white spots, as large or larger than non-adjacent rounded black spots in G. lacerata). Adult Gehyra lacerata individuals have a much more robust habitus than adults of G. wongchan sp. nov. and of the other Thai representatives of the genus (Figure 12). From Gehyra mutilata (and its Malayan subjective synonyms Gehyra butleri Boulenger, 1900, described from Kuala Lumpur, and Peropus packardii Cope, 1869, described from Penang; see synonymies by Taylor 1963 and Grismer 2011), Gehyra wongchan sp. nov. can be distinguished based on its smaller SVL (52.4 vs. 61 mm), higher VentR (48–50 vs. 35–44), lower PrePo number (17 or 18 vs. 24–44), absence of webbing on fingers and toes (vs. basal webbing), its dorsal pattern (large white spots adjacent to black crescents vs. white spots often poorly to not visible in G. mutilata, when present small or not larger than black spots), and a tail not- to moderately widened behind vent (vs. moderately to strongly widened) (Figure 13).

Trimeresurus kuiburi sp. nov. (Figures 1–6) Trimeresurus sp. — Sumontha et al. 2017a: 561. Holotype. QSMI 1500 (field number MS 703); adult male collected by Montri Sumontha, Nirut Chomngam, Parinya Phawangkhanant and Prapanth Iamwiriyakul on 27 November 2016 near Ban Thung Noi (ca. 12.084500 N, 99.948556 E), Khao Daeng Sub-district, Kui Buri District, Prachuap Khiri Khan Province, Peninsular Thailand. Paratypes (5). AUP-02005 (field number MS 701), PSUZC-R 734 (MS 704) and PSUZC-R 735 (MS 731), adult males collected on 9 August 2015, 20 June 2014 and 3 October 2020, respectively (all preserved with everted hemipenes). QSMI 1501 (MS 702) and AUP-02006 (MS 732), adult females collected on 20 June 2014 and 26 June 2020, respectively. Same locality and collectors as the holotype. Diagnosis. Trimeresurus kuiburi sp. nov. can be distinguished from all other congeneric species by the combination of its red/purple bands on a green dorsum; white concave suborbital stripe in males; white, spaced vertebral dots in males; pale green belly lacking dark dots or stripe on the lateral sides of the ventrals; partially fused 1 st supralabial and nasal scale; 19 dorsal scale rows at midbody; 164–171 ventrals; 63–65 subcaudals in males, 51–53 in females; maximal known SVL of 451 mm; and long, papillose hemipenes. Description of holotype. Adult male (Figures 1–2). Body cylindrical, long and thin; SVL 376.4 mm; tail length 88.2 mm; total length 464.6 mm. Pupil vertically elliptical. Loreal pit present. Head triangular in dorsal view, elongate, clearly distinct from the neck; head length 18.0 mm; head width 13.4 mm; HW/HL 0.74; distance between nostrils 3.7 mm. Snout elongate, 34% of HL, 1.78 times as long as horizontal diameter of eye, obliquely truncated when seen from lateral side, flattened and rounded when seen from above, with a distinct canthus rostralis. Distance between eye and nostril 4.4 mm on both sides. Rostral slightly visible from above, triangular; rostral width 2.9 mm, rostral height 2.0 mm. Nostril completely enclosed in nasal scale; nasal scale partially fused with first supralabial. Shield bordering anterior edge of pit fused with second supralabial, lacking any small scales between it and nasal. Long, thin, crescent-like subocular scale, in contact with third supralabial, separated from the 4 th and 5 th supralabials by one row of scales, separated from the 6 th supralabial by 1/2 scales. Two upper preoculars above the loreal pit, elongated, and in contact with the single loreal which separates the upper preoculars from the nasal; lower preocular forming the lower margin of the loreal pit; 1/1 postocular. Eleven/ten supralabials, 3 rd supralabial largest; 13/12 infralabials, those of the first pair in contact with each other behind the mental, the first three pairs in contact with the single pair of chin shields. Six pairs of gulars aligned between the chin shields and the preventral. One pair of enlarged internasals, in contact above the rostral. Length of the single, unfragmented supraocular 4.1/ 4.1 mm; width of supraocular 1.3/ 1.3 mm; ratio SOL/SOW 3.15. Supraoculars slightly indented on their inner margin by the upper head scales. At least nine scales between the supraoculars. Scales on snout and in the interorbital region smooth, irregular, subimbricate; temporal and occipital scales moderately keeled. Dorsal scales in 21-19-15 rows. Scale row reduction from 21 to 19 rows resulted from the fusion of the 4 th and 5 th rows above ventrals 30/31; reduction from 19 to 17 from the fusion of the 4 th and 5 th rows above ventral 108; and reduction to 15 rows by fusion of the 5 th and 6 th rows above ventral 121 on the left side, and of the 4 th and 5 th rows above ventral 122 on the right side. Dorsal scales all moderately keeled, except the first row which is unkeeled. One preventral + 166 ventrals. Anal scale single; 65 subcaudals, all divided. Hemipenes long, papillose (holotype preserved with everted hemipenes). Tail distinctly prehensile. Coloration in life. Dorsal surface of head dark green with many scales also partly or entirely dark red, especially on the snout, and the interorbital, temporal and occipital regions. A contrasting white stripe begins under the loreal pit, forms a concave curve under the eye, then continues straight from the 7 th to the last supralabial where it stops. Black vertical pupil; iris copper. The background color of the dorsum is dark green, similar to that of the head, with about 62 irregular, dark red crossbands. These red bands are about two dorsal scales long mediodorsally, but narrower on the lower flanks where they are about one dorsal scale long. At the level of the vertebral row, the red bands are generally separated by one dorsal scale. The vertebral row exhibits an interrupted line of white dots, spaced by five or six dorsal scales. Each scale of the lowest dorsal scale row shows a horizontally elongate white spot, forming a continuous white line from the neck to the end of the tail. Most of the scales of the first row also exhibit a smaller red dot, forming a dotted red line. Above the white stripe of the first dorsal scale row, the tail is red, except some thin lighter bands. The infralabials are green and white; the underside of the head is mostly white. The belly is uniformly pale green, of a lighter tone than the background color of the dorsum. In preservative the background dorsal color turned to brownish, with poorly contrasted darker marks on the head and bands on the dorsum; the belly color became greenish grey. The suborbital concave white stripe remained contrasted, as well as the white stripe on the first dorsal scale row. Variation. The main meristic, morphometric and chromatical characters of the type-series of Trimeresurus kuiburi sp. nov. are presented in Table 3. All paratypes and other specimens show a nasal scale partially fused with the first supralabial, similarly to the holotype (Figures 1, 2A, 5A & 5B, 6A & 6B). Scale row reduction from 21 to 19 rows and from 19 to 17 rows generally result from the fusion of the 4 th and 5 th dorsal rows, less frequently from the fusion of the 5 th and 6 th rows. All individuals, similarly to the holotype, show a pale green belly and lack a dark stripe or a dark dot on the lateral tips of the ventrals. Trimeresurus kuiburi sp. nov. shows a strong sexual dimorphism in the number of subcaudals (63–65 in males vs. 51–53 in females). All males exhibit a concave white suborbital stripe (Figures 1, 2A, 5A & 5B), while in females this stripe is straight, and less contrasted or sometimes not visible (Figures 1, 6A & 6B). The spaced white vertebral dots are present in all males, and absent in all females; they are regularly spaced by 5 to 6 vertebral scales. The number of dark bands on dorsum varies from 59 to 68 among the preserved types, without apparent sexual dimorphism; the highest number we observed was 70 in an adult female that we caught and released at the type-locality. All paratypes lost most of their dorsal pattern once in preservative; such a drastic color fading was also observed after a short time in preservative for Trimeresurus kanburiensis and T. venustus (Malhotra & Thorpe 2004b). Distribution and natural history. The type-locality of Trimeresurus kuiburi sp. nov. (Figure 7) is situated along the border with the southern tip of Khao Sam Roi Yot National Park where we also observed the new species. It is locally an abundant snake. Adults are found foraging at night on karst boulders (Figure 8). Young individuals are generally found on small bushes growing among the boulders. Several individuals in pre-shedding phase were found perched high on taller bushes. This pitviper is not aggressive if gently approached or handled. Nothing is known about its diet in the wild, except that one adult individual regurgitated a Cyrtodactylus samroiyot just after capture. Trimeresurus kuiburi sp. nov. is probably a generalist predator, as captive individuals ate geckos (Hemidactylus murrayi Gleadow, H. frenatus Duméril & Bibron and H. platyurus (Schneider)), frogs (Fejervarya limnocharis (Gravenhorst) and Polypedates leucomystax (Gravenhorst)) and mice. Mating was observed in captivity in the morning of 4 October 2020, after a rain shower. The preserved female QSMI 1501 contains nine eggs. We found the following squamates in direct proximity to the new pitviper species: Calotes versicolor (Daudin) (Agamidae), Cnemaspis lineogularis Wood, Grismer, Aowphol, Aguilar, Cato, Grismer, Murdoch & Sites, Cyrtodactylus samroiyot, Dixonius kaweesaki and D. siamensis (Boulenger), Gekko gecko (Linnaeus), Gehyra fehlmanni (Taylor), Hemidactylus frenatus and H. platyurus (Gekkonidae), Ahaetulla sp., Dendrelaphis subocularis (Boulenger), Lycodon capucinus (Boie) and Lycodon davisonii (Blanford) (Colubridae), Calliophis cf. maculiceps (Günther) (Elapidae) and Indotyphlops braminus (Daudin) (Typhlopidae). Etymology. The specific epithet is a noun in apposition, invariable, in reference to the administrative district Kui Buri in which the type-locality lies. We suggest the following common names: งูหางแห้มกุย (Ngu Hang Ham Kui, Thai), Kui Buri Pitviper (English), Trimérésure de Kui Buri (French), and Kui Buri Bambusotter (German). Comparison to other species. The possession of a long papillose hemipenis and a partially fused first supralabial and nasal scale makes of Trimeresurus kuiburi sp. nov. a member of the Trimeresurus group sensu stricto (Cryptelytrops group sensu Malhotra & Thorpe 2004a), which currently includes the following 20 species: Trimeresurus albolabris (Gray), T. andersonii Theobald, T. cantori (Blyth), T. cardamomensis Malhotra, Thorpe, Mrinalini & Stuart, T. caudornatus Chen, Ding, Vogel & Shi, T. davidi Chandramouli, Campbell & Vogel, T. erythrurus (Cantor), T. fasciatus (Boulenger), T. guoi Chen, Shi, Vogel & Ding, T. honsonensis (Grismer, Ngo & Grismer), T. insularis Kramer, T. kanburiensis Smith, T. labialis Fitzinger in Steindachner, T. macrops Kramer, T. mutabilis Stoliczka, T. purpureomaculatus (Gray), T. rubeus Malhotra, Thorpe, Mrinalini & Stuart, T. salazar Mirza, Bhosale, Phansalkar, Sawant, Gowande & Patel, T. septentrionalis Kramer and T. venustus Vogel (David & Vogel 2015; Chandramouli et al. 2020; Chen et al. 2020a –b; Mirza et al. 2020). Among these species, Trimeresurus kuiburi sp. nov. is readily distinguished from the widespread Trimeresurus albolabris, the Andaman and Nicobar Islands endemic T. andersonii, the Nicobar Archipelago endemics T. cantori, T. davidi, T. labialis and T. mutabilis, the Cardamom Mounts endemic T. cardamomensis, the Chinese T. caudornatus, the Indo-Burmese T. erythrurus, the Indonesian Tanahdjampea Island endemic T. fasciatus, the widespread T. guoi, the Vietnamese Hon Son Island endemic T. honsonensis, the Indonesian T. insularis, the Southeast Asian T. macrops, the southern Vietnamese-Cambodian T. rubeus, the Indian T. salazar and the Himalayan T. septentrionalis by its dorsal pattern made of purple-red bands on a green background (David & Vogel 2000; David et al. 2003; Grismer et al. 2008; Malhotra et al. 2011; Vogel et al. 2014; Chandramouli et al. 2020; Chen et al. 2020 a –b; Mirza et al. 2020). Its 19 MSR further separate it from Trimeresurus andersonii (21 MSR), T. cantori (27, 29 or 31), T. cardamomensis (21), T. caudornatus (21), T. davidi (21 or 23), T. erythrurus (23 or 25), T. fasciatus (21), T. guoi (21), T. honsonensis (21), T. insularis (21), T. labialis (21 or 23), T. macrops (21), T. mutabilis (21), T. purpureomaculatus (23–29), T. rubeus (21), T. septentrionalis (21) and most specimens of T. venustus (21, rarely 19). Compared to all members of the Cryptelytrops group, Trimeresurus kuiburi sp. nov. is superficially similar only to T. kanburiensis and T. venustus, two geographically-restricted endemics of western Thailand and the Thai-Malay Peninsula, respectively. The northernmost published record of Trimeresurus venustus was mentioned by Pauwels et al. (2013; Figure 9B) who listed the reptile taxa co-occurring at the type-locality of Cyrtodactylus sanook in Muang District, Chumphon Province. At that time we were not aware of the existence of the Kui Buri population and we did not morphologically examine in detail what we believed to be not more than a northern range extension of Trimeresurus venustus. The males of the Chumphon population, contrary to Trimeresurus kuiburi sp. nov., do not show a concave but a straight suborbital stripe, and they show a white vertebral dot every 2 to 4 vertebral scales, while male T. kuiburi sp. nov. show a dot every 5 or 6 scales, and typical male T. venustus do not show white dots (Figure 9A). In any case, Trimeresurus kuiburi sp. nov. is distinct from Trimeresurus venustus sensu auctorum. A striking pattern difference is the concave shape of the suborbital white stripe in male Trimeresurus kuiburi sp. nov. It is straight and bordering the orbit in males of the northern and southern populations of T. venustus (less contrasted or sometimes not visible in females), as can be clearly seen in photographs published in the original description of T. venustus showing specimens from Thung Song, as well as in Manthey & Grossmann (1997: 411, one individual from ‘Süd-Thailand’), Chan-ard et al. (1999: 197–198, an individual from Tai Rom Yen National Park in Surat Thani Prov., and another from Surat Thani Province without locality details, identified as T. kanburiensis), Gumprecht & Ryabov (2002: 37, a female from Kanchanadit, Surat Thani Prov., as T. kanburiensis), Vogel (2006: 122–123, five individuals from ‘South Thailand’), Figures 10–11, and in the museum material we examined (see Appendix). Many other photographs and some drawings of Trimeresurus venustus s.l. were published without any locality information, but none of them showed a convex suborbital stripe, and as far as we know, none of them was illustrating individuals from Prachuap Khiri Khan Province. This includes among others the individuals illustrated by Thumwipat & Nutphand (1982: 140, identified as T. purpureomaculatus), Kundert (1984: Fig. 106, as T. sumatranus), Lim & Lee (1989: 107 top, as T. purpureomaculatus), Nabhitabhata (1989: 176, implicitly as T. kanburiensis), Cox (1991: 398, as T. kanburiensis), Jintakune & Chanhome (1995: 129, as T. kanburiensis), Cox et al. (1998: 22, as T. kanburiensis), Jintakune (2000: 151, as T. kanburiensis), Nutphand (2001: 298–299, as T. purpureomaculatus; see David et al. 2004a), Chan-ard (2002: 127, as T. kanburiensis), Sanders et al. (2006: Supplementary material: Fig. 2b), Anonymous (2010: 3, as T. kanburiensis), Chanhome et al. (2011: 326), O’Shea (2013: 18), Chan-ard et al. (2015: 287; individuals from Surat Thani or Nakhon Si Thammarat), Suteparuk & Vasaruchapong (2016: Fig. 2), Visser (2015: Figs 610, 612–613, 615–622), Oliveri et al. (2016: Fig. 7 A-D) and Fuchs et al. (2019). The latter publication described the medical consequences of the bite by a F2 captive individual of unknown geographic origin; it is stated that this species (T. venustus) has 21-21-15 DSR, but these numbers actually come from the internet (http://www.toxinology.com/) and were not verified on the individual in question (J. Fuchs, pers. comm. to OSGP, Oct. 2020). *for the specimens with 21 MSR; for the two specimens with 19 MSR for which the SRR was noted it happened at V 22 and 84 (specimens PSGV 600 & PSGV 662; see David et al. 2004). Besides the differences in the suborbital stripe in males (concave vs. straight) and the lower number of midbody scale rows (19 vs. 21, exceptionally 19), Trimeresurus kuiburi sp. nov. differs from T. venustus by the absence (vs. presence) in both sexes of red dots on the lateral sides of the ventrals, the presence of white vertebral dots every 5 or 6 in males (vs. absence), a shorter tail in females (0.133 –0.135 vs. 0.137 –0.148), and the lower number of ventrals in males (164–166 vs. 166–181) (see Table 4). While the internasals are always in contact in Trimeresurus kuiburi sp. nov., they are in contact in only 45% of cases in T. venustus (David et al. 2004b). Trimeresurus kuiburi sp. nov. differs from T. kanburiensis by its smaller maximal SVL (451 vs. 582 mm), internasals always in contact (vs. always separated), generally wider supraocular, higher IOS (9–11 vs. 7–9), lower number of ventrals in males (164–166 vs. 170–178), more subcaudals in males (63–65 vs. 59), more subcaudals in females (51–53 vs. 41–51), distinct dorsal color (red/purple bands on bottle green background vs. dark olive-brown bands on grayish-olive background), concave (vs. straight) suborbital stripe in males, and pale green vs. creamy white belly (see Table 4). White vertebral dots in males are more spaced in Trimeresurus kuiburi sp. nov. than in T. kanburiensis (5 or 6 vs. 3–5 scales apart). Geographically, Trimeresurus kuiburi sp. nov. is well separated from both T. kanburiensis and T. venustus. Trimeresurus kanburiensis is restricted to the area of Sai Yok in Kanchanaburi Province, a part of the Tenasserim Range showing a high endemism in squamates (Sumontha et al. 2017b), located about 300 km N-NW of the typelocality of T. kuiburi sp. nov. Trimeresurus venustus is separated from T. kuiburi sp. nov. by a gap of at least 180 km (population of Chumphon Province, whose status is not yet established, but similar to T. venustus in pattern; see Figures 9A & 9B) or at least 350 km (other venustus -like populations). The analysis of cytochrome b revealed that mean interspecific p -distances between Trimeresurus kuiburi sp. nov. and related Trimeresurus species ranged from 4.57 % to 18.10 %, with the minimum of 4.57 % ± 0.91 % being to T. macrops and the maximum of 18.10 % ± 1.68 % being to P. mucrosquamatus (Table 5). Moreover, 16S rRNA gene was analyzed and showed the interspecific p -distances between T. kuiburi sp. nov. and related Trimeresurus, Published as part of Sumontha, Montri, Suntrarachun, Sunutcha, Pauwels, Olivier S. G., Ngkhanant, Parinya Pawa-, Chomngam, Nirut, Iamwiriyakul, Prapanth & Chanhome, Lawan, 2021, A new karst-dwelling, colorful pitviper (Viperidae: Trimeresurus) from northern Peninsular Thailand, pp. 307-332 in Zootaxa 4974 (2) on pages 309-326, DOI: 10.11646/zootaxa.4974.2.4, http://zenodo.org/record/4775625, {"references":["Sumontha, M., Chomngam, N., Phanamphon, E., Pawangkhanant, P., Viriyapanon, C., Thanaprayotsak, W. & Pauwels, O. S. G. (2017 a) A new limestone-dwelling leaf-toed gecko (Gekkonidae: Dixonius) from Khao Sam Roi Yot massif, peninsular Thailand. Zootaxa, 4247 (5), 556 - 568. https: // doi. org / 10.11646 / zootaxa. 4247.5.2","Malhotra, A. & Thorpe, R. S. (2004 b) Reassessment of the validity and diagnosis of the pitviper Trimeresurus venustus Vogel, 1991. Herpetological Journal, 14, 21 - 33.","Malhotra, A. & Thorpe, R. S. (2004 a) A phylogeny of four mitochondrial gene regions suggests a revised taxonomy for Asian pit vipers Trimeresurus and Ovophis. Molecular Phylogenetics and Evolution, 32, 83 - 100. https: // doi. org / 10.1016 / j. ympev. 2004.02.008","David, P. & Vogel, G. (2015) An updated list of Asian pitvipers and a selection of recent publications. In: Visser, D. (Ed.), Asian Pitvipers. Breeding experience & wildlife. Edition Chimaira, Frankfurt am Main, pp. 545 - 565.","Chandramouli, S. R., Campbell, P. D. & Vogel, G. (2020) A new species of green pit viper of the genus Trimeresurus Lacepede, 1804 (Reptilia: Serpentes: Viperidae) from the Nicobar Archipelago, Indian Ocean. Amphibian & Reptile Conservation, 14 (3), Taxonomy Section, e 264, 169 - 176.","Chen, Z., Shi, S., Gao, J., Vogel, G., Song, Z., Ding, L. & Dai, R. (2020 a) A new species of Trimeresurus Lacepede, 1804 (Squamata: Viperidae) from Southwestern China, Vietnam, Thailand and Myanmar. Asian Herpetological Research, 11 (4), 1 - 11.","Mirza, Z. A., Bhosale, H. S., Phansalkar, P. U., Sawant, M., Gowande, G. G. & Patel, H. (2020) A new species of green pit vipers of the genus Trimeresurus Lacepede, 1804 (Reptilia, Serpentes, Viperidae) from western Arunachal Pradesh, India. Zoosystematics and Evolution, 96 (1), 123 - 138. https: // doi. org / 10.3897 / zse. 96.48431","David, P. & Vogel, G. (2000) On the occurrence of Trimeresurus albolabris (Gray 1842) on Sumatra Island, Indonesia. Senckenbergiana biologica, 80 (1 / 2), 225 - 232.","David, P., Vogel, G. & Vidal, N. (2003) On Trimeresurus fasciatus (Boulenger, 1896) (Serpentes: Crotalidae), with a discussion on its relationships based on morphological and molecular data. The Raffles Bulletin of Zoology, 51 (1), 149 - 157.","Grismer, L. L., Ngo, V. T. & Grismer, J. L. (2008) A new species of insular pitviper of the genus Cryptelytrops (Squamata: Viperidae) from southern Vietnam. Zootaxa, 1715 (1), 57 - 68. https: // doi. org / 10.11646 / zootaxa. 1715.1.4","Malhotra, A., Thorpe, R. S., Mrinalini & Stuart, B. L. (2011) Two new species of pitviper of the genus Cryptelytrops Cope 1860 (Squamata: Viperidae: Crotalinae) from Southeast Asia. Zootaxa, 2757, 1 - 23. https: // doi. org / 10.11646 / zootaxa. 2757.1.1","Vogel, G., David, P. & Chandramouli, S. R. (2014) On the systematics of Trimeresurus labialis Fitzinger in Steindachner, 1867, a pitviper from the Nicobar Islands (India), with revalidation of Trimeresurus mutabilis Stoliczka, 1870 (Squamata, Viperidae, Crotalinae). Zootaxa, 3786 (5), 557 - 573. https: // doi. org / 10.11646 / zootaxa. 3786.5.4","Pauwels, O. S. G., Sumontha, M., Latinne, A. & Grismer, L. L. (2013) Cyrtodactylus sanook (Squamata: Gekkonidae), a new cave-dwelling gecko from Chumphon Province, southern Thailand. Zootaxa, 3635 (3), 275 - 285. https: // doi. org / 10.11646 / zootaxa. 3635.3.7","Manthey, U. & Grossmann, W. (1997) Amphibien & Reptilien Sudostasiens. Natur und Tier - Verlag, Munster, 512 pp.","Gumprecht, A. & Ryabov, S. (2002) Die Gattung Trimeresurus Lacepede, 1804 - Zu Kenntnisstand der Forschung. Draco, 12 (3), 31 - 44.","Vogel, G. (2006) Venomous snakes of Asia. Giftschlangen Asiens. Terralog 14, Edition Chimaira, Frankfurt am Main, 148 pp.","Thumwipat, B. & Nutphand, W. (1982) Treatment of patients bitten by venomous snakes and venomous snakes of Thailand. Thai Zoological Center, Bangkok, viii + 162 pp. [in Thai]","Kundert, F. (1984) De mysterieuze wereld van de slangen. Van Holema & Warendorf, Weesp, 39 pp., 132 pls.","Lim, F. L. K. & Lee, M. T. - M. (1989) Fascinating snakes of Southeast Asia - an introduction. Tropical Press Sdn. Bhd., Kuala Lumpur, xvii + 124 pp.","Nabhitabhata, J. (1989) Species diversity of Thai herpetofauna. In: Biodiversity in Thailand. Biological Science Seminar 7. Chiang Mai University & Science Society of Thailand, Bangkok, pp. 169 - 204. [in Thai, English summary]","Cox, M. J. (1991) The snakes of Thailand and their husbandry. Krieger Publishing Company, Malabar, xxxviii + 526 pp.","Jintakune, P. & Chanhome, L. (1995) Ngoophit nai Prathet Thai [Venomous snakes of Thailand]. Queen Saovabha Memorial Institute, Thai Red Cross, Bangkok, 175 pp. [in Thai]","Cox, M. J., van Dijk, P. P., Nabhitabhata, J. & Thirakhupt, K. (1998) A photographic guide to snakes and other reptiles of Thailand and Southeast Asia. Asia Books, Bangkok, 144 pp.","Jintakune, P. (2000) Ngoophit nai Prathet Thai [Venomous snakes of Thailand]. Samnakphimmatichon, Bangkok, 176 pp. [in Thai]","Nutphand, W. (2001) Patterns of the snakes in Thailand. Amarin Printing and Publishing Public Co. Ltd., Bangkok, 319 pp. [in Thai]","David, P., Cox, M. J., Pauwels, O. S. G., Chanhome, L. & Thirakhupt, K. (2004 a) When a book review is not sufficient to say all: an in-depth analysis of a recent book on the snakes of Thailand, with an updated checklist of the snakes of the Kingdom. The Natural History Journal of Chulalongkorn University, 4 (1), 47 - 80.","Chan-ard, T. (2002) Kheeowphetchakhat [Green killers]. Advanced Thailand Geographic, 7 (51), 114 - 132. [in Thai]","Sanders, K. L., Malhotra, A. & Thorpe, R. S. (2006) Evidence for a Mullerian mimetic radiation in Asian pitvipers. Proceedings of the Royal Society, B, 273, 1135 - 1141. https: // doi. org / 10.1098 / rspb. 2005.3418","Anonymous (2010) Venomous snakes of Thailand and the Red Cross Snake Farm & Institute in Bangkok. ACT Herpetological Association Newsletter, August - September, 2 - 3.","Chanhome, L., Cox, M. J., Vasaruchapong, T., Chaiyabutr, N. & Sitprija, V. (2011) Characterization of venomous snakes of Thailand. Asian Biomedicine, 5 (3), 311 - 328.","O'Shea, M. (2013) The trouble with Trims! The Herptile, 38 (1), 18 - 23.","Suteparuk, S. & Vasaruchapong, T. (2016) A report of a confirmed case of Beautiful pit viper (Trimeresurus venustus) bite. 15 th Asia Pacific Association of Medical Toxicology International Scientific Conference, Tan Tock Seng Hospital, Singapore, 17 - 20 November 2016, poster.","Visser, D. (2015) Asian Pitvipers. Breeding experience & wildlife. Edition Chimaira, Frankfurt am Main, 576 pp.","Oliveri, M., Cermakova, E. & Knotek, Z. (2016) The viper fangs: clinical anatomy, principles of physical examination and therapy (a review). Acta Veterinaria Brno, 85, 247 - 250. https: // doi. org / 10.2754 / avb 201685030247","Fuchs, J., Bessire, K. & Weiler, S. (2019) A confirmed bite by a Beautiful Pit Viper (Trimeresurus venustus) resulting in local symptoms. Toxicon, 163, 44 - 47. https: // doi. org / 10.1016 / j. toxicon. 2019.03.019","David, P., Vogel, G., Sumontha, M., Pauwels, O. S. G. & Chanhome, L. (2004 b) Expanded description of the poorly known pitviper Trimeresurus kanburiensis Smith, 1943, with confirmation of the validity of Trimeresurus venustus Vogel, 1991 (Reptilia: Serpentes: Crotalidae). Russian Journal of Herpetology, 11 (2), 81 - 90.","Sumontha, M., Kunya, K., Dangsri, S. & Pauwels, O. S. G. (2017 b) Oligodon saiyok, a new limestone-dwelling kukri snake (Serpentes: Colubridae) from Kanchanaburi Province, western Thailand. Zootaxa, 4294 (3), 316 - 328. https: // doi. org / 10.11646 / zootaxa. 4294.3.2"]}

Dixonius mekongensis sp. nov. (Figures 1–5) Holotype. AUP02007 (field no. MS 561),adult male caught on 2 July 2009 on a sandstone platform (ca. 15°27’19.1”N, 105°34’12.0”E) by T. Kaewmanee, Na Pho Klang Sub-district, Khong Chiam District, Ubon Ratchathani Province, eastern Thailand. Paratypes (2). PSUZC-R 736 (field no. MS 624), adult male, and AUP 02008 (field no. MS 562), adult female. Same locality, collecting date and collector as holotype. Diagnosis. Dixonius mekongensis sp. nov. can be distinguished from all other congeneric species by the combination of its maximal known SVL of 51.2 mm; 16 longitudinal rows of dorsal tubercles; 32 to 34 paravertebral scales; 22 to 24 longitudinal rows of ventrals across the abdomen; seven precloacal pores in males, no pores in females; a marked canthal stripe; and a uniform or spotted dorsal pattern. Description of holotype. Adult male (Figures 1–3). SVL 48.6 mm. Head relatively long (HL/SVL ratio 0.32), wide (HW/HL ratio 0.67), not markedly depressed (HD/HL ratio 0.44), distinct from neck. Lores and interorbital region weakly inflated. Canthus rostralis relatively prominent. Snout moderately short (SnOrb/HL ratio 0.36), rounded, slightly longer than orbit diameter (OrbD/SnOrb ratio 0.61). Scales on snout and forehead small, hexagonal to rounded, flattened, with smooth or slightly rugose surface. Scales on snout larger than those on occipital region. Eye of moderate size (OrbD/HL ratio 0.22). Pupil vertical with crenelated margins. Supraciliaries short, without spines. Ear opening oval, moderate (EarL/HL ratio 0.06); orbit to ear distance greater than orbit diameter. Rostral about twice wider than high, dorsally incompletely divided by a median cleft. Two enlarged supranasals in broad contact. Rostral in contact with supralabial I on each side, nostrils and both supranasals. Nostrils round, each surrounded by supranasal, rostral, supralabial I and two postnasals. Mental triangular, about as long as deep. Two pairs of enlarged postmentals, anteriormost approximately four times larger than posterior. Each anterior postmental bordered anteriorly by mental, medially by the other anterior postmental, anterolaterally by infralabial I, posterolaterally by the second postmental; the pair collectively bordered posteromedially by a row of six throat scales. Supralabials to mid-orbital position 7/6; enlarged supralabials to angle of jaws 8/9. Infralabials 6/7. Interorbital scales eight. Body slender, elongate (TrunkL/SVL ratio 0.44), without ventrolateral folds. Dorsal scales heterogeneous, small, irregular, flattened to conical, distributed among large, strongly keeled tubercles arranged in 16 regular longitudinal rows at midbody. Flanks covered with irregular, smooth to slightly conical scales. Gular region with relatively homogeneous, granular scales. Ventral scales smooth, imbricate, their free margin rounded. Ventrals increasing in size from throat to chest to abdomen. Midbody scale rows across belly to lowest rows of tubercles 24. Seven precloacal pores in a continuous series. Pore-bearing scales not enlarged relative to adjacent scale rows. No femoral pores or enlarged femoral scales. Fore- and hind limbs short, slender (FAL/SVL ratio 0.14; TibL/SVL ratio 0.15). Digits slender, dilated distally, all bearing robust, slightly recurved claws. Basal subdigital lamellae narrow, without scansorial surfaces (6- 8-10-11-10 right manus; 12-15-14-8-7 right pes); setae-bearing lamellae restricted to enlarged, distal, ‘‘leaf-like’’ scansors. Scales on palm and sole small, smooth, rounded to oval. Interdigital webbing absent. Relative length of digits: III>IV>V>II>I (manus), IV>V>III>II>I (pes). Tail length 60.8 mm of which the last 23.7 mm are regenerated. Supracaudals markedly keeled in the anterior portion of the tail. Ventral tail scales of the original portion of the tail enlarged into transverse plates. *partly regenerated. Coloration in life. Dorsal surface of head gray with numerous small and irregular black blotches. On each side of the head a black canthal stripe runs from the nostril through the eye and extends to the ear; it is bordered below and above by a light gray thin area. After an interruption at the level of the tympanum, the black stripe continues till above the shoulder. On the snout, at about mid-length between the eyes and the tip of the snout, a transversal bars links the left and right canthal stripes. The supralabials and infralabials are whitish. Similarly to the dorsal surface of the head, the neck, the dorsum and the dorsal surface of the original portion of the tail show a gray background color with numerous, irregularly disposed, black spots (Figure 1). Flanks lighter than the dorsum and less punctuated. Dorsal surfaces of members gray with small black spots. Ventral surfaces of head, body, members and tail whitish. In preservative the colors strongly fade and become less contrasted (Figures 2 and 3). Variation. The main morphometric and meristic characters of the type series are provided in Table 1. Morphological characters of the paratypes agree in most respects with the holotype. The last 22.1 mm of the tail of the male paratype are regenerated. The female paratype has an original tail and shows a TailL/SVL ratio of 1.17. Similarly to the holotype, the male paratype has a continuous series of pores. Precloacal pores are absent in the female. Depending on the individuals, the background color of the body is gray to brown, and the pattern varies from spotted to uniform (Figures 1, 4 and 5). The dorsal pattern does not seem to show a clear sexual dimorphism, although all fully spotted individuals observed were males, and totally uniform patterns were found only in females. Distribution and natural history. All individuals of the new species were found active at night on large sandstone platforms and boulders at the type-locality. We also observed numerous individuals in Pha Taem National Park at Soi Sawan Waterfall (15°27’38.4”N, 105°34’40.5”E) and surrounding sandstone outcrops (Figures 6 and 7). In each of these locations the vegetation is sparse, and includes patches of grass, bamboos and shrubs. Dixonius mekongensis sp. nov. is locally abundant. We found a single other squamate species in strict syntopy in all sites, Gekko petricolus Taylor, another sandstone obligate gecko. Etymology. The specific epithet refers to the Mekong River and the Greater Mekong Subregion. A cliff bordering the type-locality offers a spectacular view of this majestic river. We suggest the following common names: จิ้งจกดินแม่โขง (Djing-djok din Mekong, Thai), Mekong leaf-toed gecko (English), Dixonius du Mékong (French), and Mekong Blattfingergecko (German). Comparison to other species. The main diagnostic morphological and chromatical characters of Dixonius species are presented in Table 2. Dixonius mekongensis sp. nov. is distinguished from the Vietnamese D. aaronbaueri Ngo & Ziegler, 2009 based on its sensibly larger SVL (51.2 vs. 38.6 mm), higher Ven number (22–24 vs. 18 or 19), much higher DTR number (16 vs. 11), much lower PV number (32–34 vs. 45–50), much lower PV’ number (20–24 vs. 29–32), and its higher PrePo number (7 vs. 5). It differs from the southern Thai Dixonius dulayaphitakorum Sumontha & Pauwels, 2020 by its much lower DTR number (16 vs. 22), its lower ICS number (25–27 vs. 30–33), its marked (vs. unmarked) canthal stripe, and a very distinct dorsal pattern with smaller dorsal spots. It can be separated from the western Thai Dixonius hangseesom Bauer, Sumontha, Grossmann, Pauwels & Vogel, 2004 by its larger SVL (51.2 vs. 42.1 mm), higher DTR number (16 vs. 12–14), lower InterOrbS number (8 or 9 vs. 10) and a distinct dorsal pattern (U/Sp vs. Ba/Bl). It differs from the peninsular Thai Dixonius kaweesaki Sumontha, Chomngam, Phanamphon, Pawangkhanant, Viriyapanon, Thanaprayotsak & Pauwels, 2017 by its larger SVL (51.2 vs. 41.6 mm), shorter original tail in females (TailL/SVL ratio 1.17 vs. 1.36–1.55), higher DTR number (16 vs. 12 or 13), higher InterOrbS number (8 or 9 vs. 6 or 7), lower SL number (7–9 vs. 10 or 11), lower PrePo number (7 vs. 9–11) and by its dorsal pattern (Sp or U vs. St). It differs from the Laotian Dixonius lao Nguyen, Sitthivong, Ngo, Luu, Nguyen, Le & Ziegler, 2020 by its much lower DTR number (16 vs. 20–23), much lower PV number (32–34 vs. 40–43), lower PrePo number (7 vs. 8), and by its marked (vs. unmarked) canthal stripe. It can be separated from Dixonius melanostictus Taylor, 1962 by its much higher DTR number (16 vs. 10 or 11), lower PrePo number (7 vs. 9), and by its dorsal pattern (U/Sp vs. St). It is distinguished from the Vietnamese Dixonius minhlei Ziegler, Botov, Nguyen, Bauer, Brennan, Ngo & Nguyen, 2016 by its higher DTR number (16 vs. 14 or 15), lower PV number (32–34 vs. 38–44), and smaller black dorsal spots. It differs from the peninsular Thai Dixonius pawangkhananti Pauwels, Chomngam, Larsen & Sumontha, 2020 by its larger SVL (51.2 vs. 42.6 mm), shorter original tail in females (TailL/SVL ratio 1.17 vs. 1.33), much higher Ven number (22–24 vs. 16), higher InterOrbS number (8 or 9 vs. 7), higher PrePo number (7 vs. 6), its continuous series of pores in males (vs. a series medially interrupted by a poreless scale), and by its dorsal pattern (U/Sp vs. Ba/Bl). It differs from Dixonius siamensis by its smaller SVL (51.2 vs. 57.0 mm), its higher DTR number (16 vs. 10–14) and its marked (vs. unmarked) canthal stripe. Dixonius mekongensis sp. nov. can be distinguished from the Laotian D. somchanhae Nguyen, Luu, Sitthivong, Ngo, Nguyen, Le & Ziegler by its lower DTR number (16 vs. 19–21), lower PV number (32–34 vs. 35–40), and higher PrePo number (7 vs. 5 or 6). It can be separated from the Vietnamese Dixonius taoi Botov, Phung, Nguyen, Bauer, Brennan & Ziegler, 2015 by its larger SVL (51.2 vs. 43.9 mm), much higher DTR number (16 vs. 11 or 12), higher PrePo number (7 vs. 5 or 6), and by its dorsal pattern (U/Sp vs. Bl). From Dixonius vietnamensis Das, 2004, it differs by its larger SVL (51.2 vs. 42.4 mm), higher Ven number (22–24 vs. 15–21), and its lower PV number (32–34 vs. 36). The type-localities of all currently recognized Dixonius species recorded in Thailand are mapped on Figure 8.

Gekko pradapdao sp. nov. (Figures 1–6) Holotype. CUMZ-R 2590 (field nr. MS 733), adult male caught on 20 October 2020 in Tham (= Cave) Khao Chan (14°58’35.2”N, 101°18’11.2”E), Tha Luang District, Lopburi Province, by W. Meesook, M. Sumontha and N. Donbundit. Paratypes (3). AUP-02009 and PSUZC-R 738 (field nr. MS 735 and MS 736, respectively), adult males, and CUMZ-R 2591 (field nr. MS 734), adult female. Same locality, collecting date and collector as holotype. Diagnosis. Gekko pradapdao sp. nov. can be distinguished from all other congeneric species by the combination of its maximal known SVL of 127.1 mm, lack of contact between nostrils and rostral, 24–28 interorbital scales between supraciliaries, 89–91 scale rows around midbody, 16–18 dorsal tubercle rows at midbody, 30–34 ventral scale rows at midbody, 11–13 precloacal pores in males, a single postcloacal tubercle on each side of the base of the tail, 13–16 subdigital lamellae on 1st toe and 17–19 on 4th toe, no Y-shaped mark on head, non-banded dorsal pattern on a dark chocolate brown to black background, and a dark brown iris. Description of holotype. Adult male (Figures 1–3). SVL 118.0 mm. Head long (HeadL/SVL 0.28), relatively broad (HeadW/HeadL 0.74), somewhat depressed (HeadD/HeadL 0.40), strongly distinct from the neck. Lores and interorbital region slightly inflated. Snout moderate (SnOrb/HeadL ratio 0.43), less than twice eye diameter (OrbD/SnOrb ratio 0.66); scales on snout and forehead small to moderate in size, granular and slightly domed to flattened; scales on snout much larger than those on interorbital region. Eye relatively large (OrbD/HeadL ratio 0.28). Pupil vertical with crenulated margins. Supraciliaries short. Ear opening oval, vertically oriented, moderate (EarL/HeadL ratio 0.08); eye to ear distance slightly shorter than diameter of orbit (OrbEar/OrbD ratio 0.91). Rostral approximately twice wider (5.6 mm) than deep (2.7 mm). No rostral groove present; two much enlarged supranasals partly separated by a single, small, internasal; rostral in contact with supralabial I and supranasals. Nostrils oval, each surrounded by supranasal, first supralabial, two dorsally located postnasals and a large nasal, which enters the nostril forming a recessed valvular flange in its posterior third. Mental triangular, deeper (3.7 mm) than wide (2.4 mm) and not much deeper than infralabials; one pair of greatly enlarged postmentals meeting behind the mental. Each postmental bordered anteriorly by first infralabial, medially by mental, laterally by the first scale in a row of enlarged scales bordering the infralabials, and posteriorly by two chin shields (larger than the granular gular scales), the medial one in shared contact with both left and right postmentals. Infralabials bordered by a row of enlarged scales, decreasing in size posteriorly. Enlarged supralabials to midpoint of orbit 10 (left and right); supralabials to angle of jaws 13 (left)-12 (right); enlarged infralabials 11 (left and right). Interorbital scale rows across narrowest point of frontal 12, between supraciliaries 28. Supraorbital scales heterogeneous in size, largest at medial edge of orbit in midorbital position. Body robust, trunk relatively long (TrunkL/SVL ratio 0.42), dorsoventrally depressed in cross-section, with distinct ventrolateral folds without denticulate margins. Dorsal scales heterogeneous, granular, rectangular to oval, and flattened. Regularly arranged, small (about 6 times size of granules), conical, posteriorly directed tubercles extending from posterior margin of orbit to tail; tubercles smaller on parietal region than elsewhere; tubercles in 18 rows at midbody. Ventral scales smaller than dorsal tubercles, subimbricate, becoming granular and much smaller in gular region. Midbody scale rows across belly between ventrolateral folds 30. Eleven pore-bearing precloacal scales, in a continuous row. Scales on palm and sole smooth, flat, rounded. Scales on dorsal aspects of hind limbs heterogeneous - granular, intermixed with larger tubercles, some conical, others flattened. Scales on dorsal surface of forelimb proximal to elbow subimbricate, weakly heterogeneous, those distal to elbow granular, intermixed with larger tubercles. Fore- and hind limbs moderately long, stout; forearm and tibia moderately long (FAL/SVL ratio 0.15; TibL/SVL 0.17). Digits relatively short; digit I, both manus and pes, clawless, all remaining digits strongly clawed; distal portions of digits strongly curved, arising from distal portion of expanded subdigital pad. Scansors beneath each toe undivided; scansors from proximalmost at least twice diameter of palmar scales to distalmost: 13-14-16-15-14 (left manus), 12-13-16-16-13 (right manus), 14-15-17-17-15 (left pes), 13-14-16-17-15 (right pes). Relative length of digits of manus: III>IV=II>I>V; of pes: IV>V>III>II>I. Tail depressed, slightly longer than head and body (TailL/SVL ratio 1.10); dorsal surface of tail covered with small, square to oval, juxtaposed to weakly subimbricate granules forming regular transverse rows. Median pair of enlarged subcaudal plates extending about 2/3 across the width of tail. Posterior portion of dorsum of each tail segment with a single transverse row of 6 enlarged, rounded tubercles. A single post-cloacal tubercle on each side of tail base. Coloration in life. Dorsal surface of head, dorsum, and dorsal surfaces of members and tail uniformly dark chocolate brown, with very contrasting, white spots (covering one tubercle and sometimes also some adjacent small scales) not arranged in transverse rows. Background color of dorsal surfaces of fingers and toes brown but slightly lighter than dorsum background color, with some white tubercles. In the anterior half of the tail, white dorsal tubercles are more or less arranged in five regularly spaced transverse rows, followed in the second half of the tail by five transverse white bands, the last one near the tail tip. No nuchal collar. No Y-shaped mark on head. Iris dark brown. Infralabials chocolate brown. Throat whitish, with irregularly distributed yellowish scales. Venter and lower surfaces of members and tail whitish, with a few irregular chocolate brown spots, and a few yellowish scales. Palms gray. Finger and toe lamellae white. Variation. Main morphometric and meristic characters of the type series are provided in Table 1. Morphological characters of the paratypes agree in most respects with the holotype. The female lacks precloacal pores. All three paratypes have a partly regenerated tail; AUP-02009 even shows a bifurcated regenerated tail (Figure 4). Distribution and natural history. All types and observed individuals were found within Khao Chan Cave, and the species is known only from its type-locality (Figure 7). The cave, in which the species is abundant, has been radically transformed into a Buddhist temple (Figure 8), and is totally surrounded by cultivated fields. The ground of the cave has been tiled; many cracks and holes have been filled with cement. The holotype was found while it was foraging on a plastic barrel (Figure 1). Nothing is known about the diet or the reproduction of the new species. Five other gecko species were found to share the cave with the new species: Dixonius siamensis Boulenger, Gehyra cf. angusticaudata (Taylor), Gekko gecko (Linnaeus), Hemidactylus frenatus Duméril & Bibron and H. platyurus (Schneider). We also found at least 20 shed skins of Lycodon davisonii (Blanford) (Colubridae) within the cave. Etymology. The specific epithet is a name in apposition, invariable, based on the combination of the Thai words “pradap” (decorated) and “dao” (star). The starry effect is due to the fact that, unlike in other members of the Gecko subgenus such as G. (G.) siamensis or G. (G.) smithii, the white dorsal tubercles are not aligned in transverse rows, but irregularly distributed. It is reinforced by the contrast between the white tubercles and the unusually dark background color of the new species. We suggest the following common names: ตุ๊กแกประดับดาว (Took-kay pradap dao; Thai); Starry Tokay Gecko (English), and Gecko tokay étoilé (French). Comparison to other species. The subgenus Gekko currently includes seven species: G. (G.) albofasciolatus (Günther); G. (G.) gecko (represented by its nominal subspecies and by G. (G.) gecko azhari Mertens); G. (G.) nutaphandi Bauer, Sumontha & Pauwels; G. (G.) reevesii (Gray); G. (G.) siamensis Grossmann & Ulber; G. (G.) smithii Gray; and G. (G.) verreauxi Tytler. The main diagnostic characters of these species are compared in Table 2. Gekko (G.) pradapdao sp. nov. differs from the Bornean G. (G.) albofasciolatus by its smaller SVL (127.1 vs. 165.1 mm), its much higher DTR number (16–18 vs. 10), its higher VentR (30–34 vs. 26), lower PrePo number (11–13 vs. 16), its dark brown (vs. green) iris and the absence (vs. presence) of a Y-shaped mark on the head. It can be differentiated from the Bangladeshi Gekko (G.) gecko azhari by its smaller SVL (127.1 vs. 155 mm), its higher InterCilS number (24–28 vs. 20–23), higher DTR number (16–18 vs. 15), lower number of postcloacal tubercles (one vs. 2 or 3), lower SubDLT1 (13–16 vs. 17 or 18), lower SubDLT4 (17–19 vs. 22–24), and the absence (vs. presence) of a Y-shaped mark on the head. It differs from the widespread Gekko (G.) gecko gecko by its smaller SVL (127.1 vs. 161 mm), higher DTR number (16–18 vs. 11–13), different iris color (dark brown vs. pale golden, copper, or brown to olive), absence (vs. usual presence) of a Y-shaped mark on head, and a distinct dorsum background color (dark chocolate brown vs. ultramarine gray to bluish- or brownish-gray). It can be separated from the western Thai Gekko (G.) nutaphandi by its much lower MSR (89–91 vs. 115 or 116), its higher DTR number (16–18 vs. 14), lower PrePo number (11–13 vs. 17–22), higher SubDLT1 (13–16 vs. 12), and higher SubDLT4 (17–19 vs. 15), dark brown (vs. brick red) iris, and dark chocolate brown (vs. grayish brown to chestnut brown) dorsum background color; moreover in Gekko (G.) nutaphandi the white dorsal tubercles are arranged in transverse rows, while they are irregularly distributed in Gekko (G.) pradapdao sp. nov. The new species can be differentiated from the Sino- Vietnamese Gekko (G.) reevesii by its much smaller SVL (127.1 vs. 173 mm), its dark brown (vs. pale golden) iris, absence (vs. presence) of a Y-shaped mark on the head, and its dark chocolate brown dorsum (vs. grayish-brown to grayish-green) with irregularly arranged white tubercles (vs. disposed in transverse rows). From the central Thai Gekko (G.) siamensis it can be distinguished by its generally lower SL number (12–14 vs. 13–21), much lower MSR (89–91 vs. 121–132), its single postcloacal tubercle (vs. two), dark brown (vs. green) iris, and its dark chocolate brown (vs. gray-brown to dark green) dorsum background color with irregularly arranged white tubercles (vs. disposed in transverse rows). It can be differentiated from Gekko (G.) smithii by its much smaller SVL (127.1 vs. 191 mm), lower InterCilS number (24–28 vs. 31–43), higher DTR number (16–18 vs. 8–13), dark brown (vs. green) iris, absence (vs. presence) of a Y-shaped mark on head, and by its dark chocolate brown (vs. yellow-green to dark green) dorsum background color with irregularly arranged white tubercles (vs. disposed in transverse rows). It can be separated from Gekko (G.) verreauxi, endemic to the Andaman Islands, by its smaller SVL (127.1 vs. 155 mm), the contact between its nostril and rostral (vs. no contact), much higher DTR number (16–18 vs. 11), smaller SubDLT4 (17–19 vs. 20–22), dark brown (vs. green) iris, and its dark chocolate brown (vs. gray-brown) dorsum background color.

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. 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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. 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(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"]}