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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. 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