18 research outputs found
FIGURE 12 in Night stalkers from above: A monograph of Toxicodryas tree snakes (Squamata Colubridae) with descriptions of two new cryptic species from Central Africa
No full textFIGURE 12. Photos of Toxicodryas adamanteus sp. nov. in life. (A and B) adult male holotype (UTEP 22204, SVL = 812 mm) from Npenda, Democratic Republic of the Congo (DRC). Photo: E. Greenbaum, (C) adult female (UTEP 22205) from Salonga National Park, DRC. Photo: E. Greenbaum, (D) adult female (UTEP 22201) from Kihungwe, DRC, with a vespid wasp. Photo: E. Greenbaum, (E) ventral view of adult male (UTEP 22200) from Mayindombe, DRC. Photo: E. Greenbaum, (F) unsexed individual from Banalia, DRC. Photo: K. Mebert, (G) juvenile female (UTEP 22198) from Punia Gorilla Reserve, DRC. Photo: G.M. Mitamba.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, Rödel, 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 page 29, DOI: 10.11646/zootaxa.4965.1.1, http://zenodo.org/record/472302
Toxicodryas vexator Greenbaum & Allen & Vaughan & Pauwels & Wallach & Kusamba & Muninga & Aris- Tote & Mali & Badjedjea & Penner & Rödel & Rivera & Sterkhova & Johnson & Tapondjou & Brown 2021, sp. nov.
No full textToxicodryas 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 <1 meter in 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’ out when he detected food nearby. The bite occurred just after I opened the front [of] the cage (a tall arboreal habitat); he ‘poured’ out and delivered what was clearly a feeding strike with firm attachment on the ulnar zone, dorsal aspect [of the right] wrist. Attachment lasted approximately 30–40 seconds with attempted jaw advancement; I was able to detach him by gently [maneuvering] his head. Pain was akin to several punctures with a sewing needle; immediate bleeding was proportional to the wounds and transient (<3 min). I developed some very slight edema also proportional to the extent of physical trauma, but did have some persistent erythema (lasting approximately 36 hr), and some slight pruritus that also lasted a couple of days. The second bite was from an approximately 45 cm male and was [a] quick bite and release delivered to the lateral aspect of the [right] index finger, consistent with a brief defensive bite. I only experienced some sharp punctures, very brief, slight bleeding (lasting barely a minute) and a few minor lacerations from a few of the teeth inflicting a ‘dragging’ effect; there were no other effects. It should be noted that although most of the minor effects of the first bite (the second was too brief to assess) could be assigned to local effects of low volume venom inoculation, some contribution of local hypersensitivity is certainly possible. I was sensitised in the 1970s during the beginning of my venom research.” Etymology. The specific epithet is a noun in apposition, invariable, from the Latin noun vexator, meaning harasser or stalker, in reference to the fact that this snake stalks prey when they are sleeping, and to its aggressiveness when disturbed.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, Rödel, 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/472302
Toxicodryas adamanteus Greenbaum & Allen & Vaughan & Pauwels & Wallach & Kusamba & Muninga & Aris- Tote & Mali & Badjedjea & Penner & Rödel & Rivera & Sterkhova & Johnson & Tapondjou & Brown 2021, sp. nov.
No full text<i>Toxicodryas adamanteus</i> sp. nov. <p>(Table 2, Figs. 8, 12–13)</p> <p> 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 <i>Toxicodryas pulverulenta</i> 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).</p> <p> <b>Diagnosis.</b> A species of <i>Toxicodryas</i> restricted to West, Central and East Africa, east of the Niger Delta, defined by the following combination of characters: maximum SVL <1 meter (vs. maximum SVL> 1 meter in <i>T</i>. <i>blandingii</i> and <i>T</i>. <i>vexator</i> <b>sp. nov.</b>), DSRN 18–23 (vs. 23–25 in <i>T</i>. <i>blandingii</i> and 23–29 in <i>T</i>. <i>vexator</i> <b>sp. nov.</b>), DSRM 18–21 (vs. 21–25 in <i>T</i>. <i>blandingii</i> and <i>T</i>. <i>vexator</i> <b>sp. nov.</b>); cloacal plate undivided (vs. usually divided in <i>T</i>. <i>blandingii</i>, and divided or undivided in <i>T</i>. <i>vexator</i> <b>sp. nov.</b>); 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 <i>T</i>. <i>blandingii</i> and <i>T</i>. <i>vexator</i> <b>sp. nov.</b>); 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 <i>T</i>. <i>blandingii</i> and <i>T</i>. <i>vexator</i> <b>sp. nov.</b>).</p> <p> <b>Holotype.</b> 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.</p> <p> <b>Paratopotype.</b> UTEP 22203 (field no. ELI 2212) adult female with same collection details as the holotype.</p> <p> <b>Paratype.</b> 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.</p> <p> <b>Description of the holotype.</b> 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.</p> <p> <b>Coloration (in life) of the holotype.</b> 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.</p> <p> <b>Coloration (in preservative) of the holotype.</b> 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.</p> <p> <b>Variation.</b> Morphometric variation of <i>Toxicodryas adamanteus</i> <b>sp. nov.</b> 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., <i>T</i>. <i>adamanteus</i> <b>sp. nov.</b>) had more subcaudals than snakes from West Africa (i.e., <i>T</i>. <i>pulverulenta</i>). 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.</p> <p> 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 <i>T</i>. <i>pulverulenta</i>.</p> <p> 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 <i>et al</i>. (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 <i>et al</i>. (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.</p> <p> <b>Diet.</b> 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 <i>Agama</i> sp. lizard (<i>sensu</i> Leaché <i>et al</i>. 2017). Spawls & Branch (2020) listed the diet as arboreal lizards and rodents.</p> <p> <b>Parasites.</b> Pitman (1938) noted a juvenile from Uganda with numerous ticks.</p> <p> <b>Behavior.</b> 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.</p> <p> <b>Reproduction.</b> 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.</p> <p> <b>Habitat.</b> 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 (<i>Platycerium angolense</i>) 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 <i>(</i> 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), <i>T. adamanteus</i> <b>sp. nov.</b> was among the most rarely encountered species.</p> <p> <b>Geographic distribution and habitat.</b> Based on molecular data from Allen <i>et al</i>. (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.</p> <p> <b>Venom.</b> Based on a specimen from Cameroon, Taub (1967) described the histological morphology of the Duvernoy’s gland. Venom composition and effects are unknown (Weinstein <i>et al</i>. 2011).</p> <p> <b>Etymology.</b> The specific epithet <i>adamanteus</i> is a Latin adjective referring to the diamond-shaped marks on the flanks and dorsum of this species.</p>Published as part of <i>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, Rödel, 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)</i> on pages 27-32, DOI: 10.11646/zootaxa.4965.1.1, <a href="http://zenodo.org/record/4723024">http://zenodo.org/record/4723024</a>
Toxicodryas pulverulenta
No full textToxicodryas 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 Ceríaco 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. Thomé, not Edina, Grand Bassa County, Liberia as stated by Loveridge (1958:269).” The latter citation is in error (Loveridge 1957). Ceríaco 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 São Tomé over the last century failed to document this species, Ceríaco 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 (Ceríaco 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 Ceríaco 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. Günther (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 Leaché et al. 2017). Böhme (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, Böhme (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. Rödel & Mahsberg (2000) found snakes in Ivory Coast from thick undergrowth in a swamp bordered by fields and secondary forest, and swampy primary forest. Branch & Rödel (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, Rödel, 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/472302
Systematics of the Thirteen-scaled Green Snake <i>Philothamnus carinatus</i> (Squamata: Colubridae), with the description of a cryptic new species from Central and East Africa
No full textRecent molecular phylogenies of African green snakes suggested the geographically widespread species Philothamnus carinatus includes at least two distinct lineages. We utilised an integrative taxonomic approach with morphological and genetic data to reconcile the taxonomic status of these cryptic lineages, including the recently described taxon P. brunneus from West Africa. We sequenced three mitochondrial (16S, cyt b and ND4) and two nuclear (c-mos and RAG1) genes from several Central African populations of P. carinatus and combined our data with other closely related species to infer a maximum likelihood phylogenetic tree with IQ-TREE. Our results are consistent with previous studies that showed P. cf. carinatus populations from Democratic Republic of the Congo (DRC) represent a cryptic lineage that is distinct from P. carinatus sensu stricto in Cameroon, Central African Republic, Equatorial Guinea (including Bioko Island), Gabon, eastern Nigeria, Republic of the Congo, and extreme western DRC. In our preferred tree, P. brunneus (limited to 16S molecular data) was recovered as a relatively long branch in a moderately supported clade with P. carinatus sensu stricto, whereas P. cf. carinatus populations from northern Angola, most of DRC, and East Africa (Burundi, Kenya, Rwanda, Tanzania and Uganda) were described as a new species. A possible hybrid population between south-eastern Cameroon and north-western DRC is consistent with an increasing body of evidence suggesting the Ubangi River might represent a hybrid zone area.</p
Hemidactylus gramineus Ching & Tay & Brown & Mohareb & Sethi & Annamalai 2023, sp. nov.
No full textHemidactylus gramineus sp. nov. Figures 1, 2; Table 1 urn:lsid:zoobank.org:act: 8C27856D-9BAD-4880-930F-2503794734C3 Hemidactylus cf. nzingae: Ceríaco et al. (2020a: 14) Hemidactylus sp.: Ceríaco et al. (2020a: 19) Holotype.— UTEP 22208 (field number ELI 1645; Figs. 1, 2), adult male collected near bank of Lumene River, Bombo-Lumene Reserve (-4.42027 °, 16.04687 °, 548 m), Kinshasa Province, DRC, collected by Chifundera Kusamba and Wandege M. Muninga on 7 June 2013. Paratype.— UTEP 22209 (field number ELI 1674), adult male collected in Bombo-Lumene Reserve (-4.41843 °, 16.04420 °, 555 m), Kinshasa Province, Democratic Republic of the Congo, collected by Chifundera Kusamba, Wandege M. Muninga, Mwenebatu M. Aristote, and Eli Greenbaum on 7 June 2013. Diagnosis.—A small-sized Hemidactylus, maximum SVL 39.5 mm, with a short, blunt snout (Fig. 1). Dorsal pholidosis heterogeneous, with 12–14 irregularly arranged longitudinal rows of subtrihedral, striated, strongly keeled tubercles at midbody. Two well-developed pairs of postmentals, the inner pair longer than the outer pair, about half the size of the mental, and in contact behind the mental. Ventrolateral folds distinct, 23– 25 scale rows across midventer. Five divided scansors beneath first digit of both manus and pes, seven beneath fourth digit of manus, seven beneath the fourth digit of pes. Males with a continuous series of eight precloacal pores. Body dorsum grayish brown, bordered by irregular, squiggly, dark brown markings that are interrupted by pairs of light gray spots, bordered by a cream/tan band that passes from the tip of the snout through the eye to the upper portion of the flanks and onto the tail, with numerous whitish tubercles on the flanks. Description of Holotype.—The holotype is an adult male in good condition (Fig. 1). The body shape is somewhat dorsoventrally flattened. Measurements and meristic characters of the holotype are presented in Table 1. Head large (HL/ SVL 0.30 mm), wide (HW/HL 0.60 mm), not depressed (HH/HL 0.38 mm), distinct from neck. Canthus rostralis moderately distinct. Snout short (SE/HL 0.43 mm), longer than eye diameter; scales on snout, canthus rostralis, forehead and between eyes heterogeneous, juxtaposed, and weakly pointed; scales on snout, canthus rostralis and forehead twice the size of those on the occipital and interorbital regions. Eye small (ED/HL 0.18 mm), supraciliaries large, pointed, with those at the anterior end of orbit slightly larger. Ear opening oval (greatest diameter 0.7 mm); eyeto-ear distance greater than diameter of eye. Rostral wider than deep; three internasals, with the two outer ones enlarged and separated from each other by the smaller one in the middle; one supranasal on each side that is smaller than internasal, one pair of still smaller postnasals; rostral in contact with nostril, supralabial I, supranasal and internasal; nostrils small (> 1 mm), oval, each surrounded by supranasal, rostral, supralabial I and postnasal; one or two rows of scales separate orbit from supralabials. Mental triangular; two postmentals, smaller than the mental and in extensive contact with each other behind mental; outer pair about one third the size of the inner pair, separated from each other by inner pair and four gular scales. Inner postmental bordered by mental, infralabial I, outer postmental and two gular scales; outer postmental bordered by infralabials I and II, inner postmental, three to four small gular scales of which the outer one is enlarged and continue as a single row of enlarged scales below infralabials. Infralabials bordered by a double row of enlarged scales. Supralabials and infralabials to angle of jaw 10 (right and left) and 9 (right and left), respectively. Body relatively stout, moderately elongate (TRL / SVL 0.42 mm), ventrolateral folds without denticulate scales. Dorsal pholidosis heterogeneous, comprising conical, granular, striated scales intermixed with enlarged, fairly regularly arranged, longitudinal rows of 14 subtrihedral, strongly keeled, striated tubercles at midbody, extending from occipital region to tail, each enlarged tubercle roughly six to eight times longer than adjacent granules, surrounded by rosette of 10–12 small granules, 2 or 3 granules between two adjacent enlarged tubercles; enlarged tubercles on nape and shoulder small and pointed, those on occipital region heterogeneous. Ventral scales larger than dorsal, smooth, imbricate; midbody scale rows across belly 25 (between ventrolateral folds); gular region with still smaller, subimbricate scales, those on lateral aspect of neck granular, anterior gular scales slightly larger than the rest. Eight precloacal pores. Scales on the palm and sole smooth, imbricate, rounded; scales on dorsal aspect of upper arm larger than granules on dorsum, subimbricate and striated, dorsal aspect of forearm with smaller, striated, conical and granular scales, intermixed with a few enlarged conical tubercles; those on dorsal part of thigh and shank conical, granular, striated, intermixed with enlarged, striated, subtrihedral tubercles, which are numerous on shank compared with anterior aspect of thigh; posterior aspect of thigh lacks enlarged tubercles. Forelimbs and hind limbs relatively short, stout; forearm short (FL/ SVL 0.14 mm); tibia short (CL/ SVL 0.16 mm); digits moderately short, strongly clawed; all digits of manus and digits I–IV of pes indistinctly webbed; terminal phalanx of all digits curved, arising angularly from distal portion of expanded lamellar pad, half or more than half as long as associated toepad; scansors beneath each toe in straight transverse series, divided except for distal and two basal scansors on digit I and one or two distal and basal scansors in all other digits that are single; scansors from proximal-most part of the toe at least twice diameter of palmar scales to distal-most single scansor: 5-6-6-7-7 (right manus), 5-8-7-7-6 (right pes). Relative length of digits: III = IV> V> II> I (right manus); IV> III> V> II> I (right pes). The tail bears spinose tubercles dorsally, and subimbricate subcaudal scales, about one fourth of the width of the tail. Coloration (in Life and PreserƲatiƲe).—Vertebral region of dorsum grayish brown, bordered by irregular, squiggly, dark brown markings that are interrupted by pairs of light gray spots; the dark brown markings transition to chevrons on the tail. Crown of head with scattered, poorly defined brown markings that resemble irregular chevrons. A cream/tan band passes from the tip of the snout through the eye to the upper portion of the flanks and onto the tail, with numerous whitish tubercles on the flanks. The lower portion of the flanks has a grayish brown band that is bordered by numerous light gray or whitish tubercles. The lowest margin of the flanks is light brown with irregular brown markings, and transitions to the venter. Forelimbs and hind limbs grayish brown with irregular dark brown crossbars. Venter cream; palms and soles cream. In preservative the specimens look more brownish with the pattern not so clearly delimited. Variation.—Variation in scalation and body measurements of the paratype of Hemidactylus gramineus sp. nov. is reported in Table 1. The paratype agrees almost entirely with the holotype, although its coloration is less contrasting than that of the holotype. Coloration of a tentatively assigned third specimen (based on a photo of a live specimen) from Mayongongo village [-4.19, 14.96, 368 m a.s.l.], Pool Department, Republic of Congo (Fig. 4), is nearly identical to that of the holotype (Fig. 2). Comparison aeith West and Central African Congeners.—The newly described species can be morphologically distinguished from all of its regional congeners, including those occurring in the DRC and in the areas neighboring the western DRC: Gabon, Republic of the Congo, and Angola. Hemidactylus gramineus sp. nov. may be readily distinguished from H. ituriensis and Hemidactylus coalescens Wagner, Leaché and Fujita, 2014, by being much smaller (maximum [max] SVL 39.5 vs. 78 mm in H. coalescens and 89 mm in H. ituriensis) and in not having the typical dorsal crossbands that characterize these two species (and the H. fasciatus group as a whole). It may be distinguished from H. angulatus Hallowell, 1854 by having only 8 precloacal pores (vs. 20–46 uninterrupted precloacal-femoral pores in males), by having a lower number of tubercle rows on the dorsum (12–14 vs. 14–25), and by having small subcaudal scales (vs. transversely enlarged). Hemidactylus gramineus sp. nov. is readily distinguished from H. kamdemtohami Bauer and Pauwels, 2002, and H. richardsonii (Gray, 1845) by the lack of basal digital webbing and from H. matschiei (Tornier, 1902) by having spiny tubercles on the dorsum and small subcaudal scales. It may be distinguished from H. steindachneri by lacking a longitudinal row of keeled tubercles on the ventrolateral border of the flanks, and from H. hecqui by not having the nostril in contact with the first supralabial, by having a relatively larger head depth (vs. flat head in H. hecqui), lower number of tubercle rows on the dorsum (12–14 vs. 16), by having fewer rows of ventral scales (23–25 vs. 26), and a higher number of precloacal-femoral pores in males (8 vs. 6). Hemidactylus gramineus sp. nov. can be distinguished from H. longicephalus by having a lower number of tubercle rows on the dorsum (12–14 vs. 17–18), fewer granular scales between the dorsal tubercles (1–3 vs. 3–6), and fewer rows of ventral scales (23–25 vs. 30–33). It can be distinguished from H. paiƲae by its smaller size (max SVL 39.5 vs. 68.4 mm), a lower number of granular scales between the dorsal tubercles (1–3 vs. 4–9), and a lower number of rows of ventral scales (23–25 vs. 28–34). Hemidactylus gramineus sp. nov. may be distinguished from H. benguellensis by a much lower number of precloacal femoral pores in males (8 vs. 23–33) and fewer granular scales between the dorsal tubercles (1–3 vs. 4–9). The new species can be distinguished from H. bayonii and H. Ʋernayi by fewer granular scales between the dorsal tubercles (1–3 vs. 3–4), and fewer rows of ventral scales (23–25 vs. 28–32 in H. bayonii and 28–31 in H. Ʋernayi). Hemidactylus gramineus sp. nov. is easily distinguishable from H. mabouia by its smaller size (max SVL 39.5 vs. 67.4 mm), lower number of precloacal femoral pores in males (8 vs. 28–39), fewer granular scales between the dorsal tubercles (1–3 vs. 5–10), and lower number of rows of ventral scales (23–25 vs. 34–37). Hemidactylus gramineus sp. nov. can be distinguished from H. muriceus sensu stricto by a higher number of tubercle rows on the dorsum (12–14 vs. 7–12) and fewer granular scales between the dorsal tubercles (1–3 vs. 5–10). The new species can be distinguished from H. pseudomuriceus by having fewer precloacal femoral pores in males (8 vs. 14–17) and fewer granular scales between the dorsal tubercles (1–3 vs. 5–10). The newly described species can be distinguished from H. echinus by its lack of two rows of scattered spiny tubercles on the ventral side of the tail (a character that differentiates H. echinus from all other known Hemidactylus), and by being smaller (max SVL 39.5 vs. 68.0 mm). Hemidactylus gramineus sp. nov. differs from H. ansorgii by a lower number of precloacal femoral pores (8 vs. 10– 11) and a stockier body (H. ansorgii has a slender body and narrow head with a short interorbital distance). Hemidactylus ansorgii also has a row of pointed tubercles separating the venter and the flanks, which are completely devoid of any tubercles (see figures in Tornier, 1902; Perret, 1975), whereas in H. gramineus sp. nov. this ventrolateral row of tubercles is lacking, and its tubercles extend across the flanks and dorsum. Hemidactylus gramineus sp. nov. can be distinguished from H. nzingae by having a lower number of tubercle rows on the dorsum (12–14 vs. 16–21) and by having irregular, undulate, dark brown markings that are interrupted by pairs of dorsal light gray spots (vs. well-marked series of dark-brown continuous ‘‘W-shaped’’ transverse markings from occiput to sacrum in H. nzingae); the new species can be distinguished from H. hannasabinae by having a lower number of tubercle rows on the dorsum (12–14 vs. 16–18), by having strongly keeled, striated tubercles at midbody (vs. smoothly keeled in H. hannasabinae), and by being smaller (max SVL 39.5 vs. 47.4 mm). Distribution.—The new species is confirmed from Bombo-Lumene Reserve in Kinshasa Province, southwestern DRC (Fig. 3). A photo of a third specimen from Mayongongo village [-4.19, 14.96, 368 m a.s.l.], Pool Department, Republic of the Congo (Fig. 4) presents a remarkable morphological resemblance with the newly described species. The specimen is deposited in the laboratory of the Institut National des Sciences Exactes et Naturelles of the Republic of Congo in Brazzaville. We tentatively assign this specimen to the new species, but it was not possible to include it in our morphological or molecular datasets, and further research on this specimen and additional material are needed to confirm the presence of the species in the Republic of the Congo. Habitat and Natural History Notes.—The holotype was collected in the early evening while on the ground near the bank of the Lumene River. The paratype was collected in the early evening while it was crawling through grass in sandy soil near a small pond. Bombo-Lumene Reserve is located ~ 130–150 km east of Kinshasa, in a hilly area known locally as the Bateke Plateau (name derived from the local Bateke tribal people), at an elevation of ~ 400–960 m a.s.l. (Kayumba et al., 2015). It was originally created as a Game Reserve in 1968, but was elevated in 1976 to Natural Reserve International Union for Conservation of Nature (IUCN) category VI because of its ecological importance (Kayumba et al., 2015). The rainy season lasts for 8 mo from mid-September to mid-May, whereas the 4-mo dry season occurs from mid-May to mid-September. The rainfalls reach an annual range from 1,600 to 1,629.7 mm, and the average annual temperature is 24.4 ° C (Kayumba et al., 2015). Bombo-Lumene Reserve is a forest–savanna mosaic (i.e., western Congolian forest–savanna mosaic sensu Burgess et al. [2004]), with nearly all forest occurring as gallery forest adjacent to the Bombo or Lumene Rivers (Greenbaum, pers. obs.). We encountered the new species during the dry season that typically lasts for 4 mo (Milau et al., 2016), and we observed large brush fires during our (CK and EG) brief visit to the reserve. Other lizards encountered at the reserve included Hemidactylus mabouia (in or near human habitations), Trachylepis maculilabris Gray, 1845; Gerrhosaurus cf. nigrolineatus Hallowell, 1857; and Panaspis cabindae Bocage, 1866 (Medina et al., 2016; Allen et al., 2019). The specimen from Republic of Congo was collected in ‘‘savannah’’ near a village (K. Jackson, pers. com. 14 January 2009). Etymology.—The specific epithet ‘‘ gramineus ’’ refers to the grassy type of habitat from which the species was collected in the DRC and the Republic of the Congo (Fig. 5) and is applied here as an adjective in the genitive singular. We propose the French common name of ‘‘Gecko de l’herbe’’ and the English common name of ‘‘grass-dwelling tropical gecko.’’Published as part of Ching, Ann Hui, Tay, Tricia, Brown, Bryan, Mohareb, Amir M, Sethi, Aisha & Annamalai, Aniyizhai, 2023, Dermatologic conditions of adult refugees following resettlement in the United States, 2015 to 2018., pp. 105-111 in Journal of Herpetology 7 (7) on pages 105-111, DOI: 10.1670/20-094, http://zenodo.org/record/787712
Toxicodryas blandingii
No full textToxicodryas blandingii (Hallowell, “1844” 1845) (Table 1, Figs. 5–8) Dipsas Blandingii: Hallowell (“1844” 1845:170); type locality: “ Liberia, West Africa. ” Triglyphodon fuscum: Duméril, Bibron & Duméril (1854:1101); type locality: “ Grand-Bassam, sur la Côte d’Ivoire (Guinée)” [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 Duméril (1859:212); no type locality provided. Triglyphodon fuscum var. obscurum: Duméril (1861:211); type locality: “ Côte d’Or ” [Ghana]. Dipsas regalis: Jan (1871:3, Livraison 38, pl. vi, fig. 2) in Jan & Sordelli (1870 –1881); type locality: “ Côte d’Or ” [Ghana]. Dipsas globiceps var. tumboensis: Müller (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 Duméril 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. variété? (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. Müller (1885:687) seemed to suggest that Jan illustrated his specimen (collected by Dr. Mähly from “Goldküste” [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 <1 meter in 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. Müller (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 Müller (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 & Mané (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 & Mané (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 Müller (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, “barré” [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 veloutés” [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 two color morphs (Lawson 1993; Rasmussen 1997a), described Boiga blandingi occidentalis for Cameroonian specimens that were uniform black or bluish black, whereas B. blandingi subfulva was named for Cameroonian specimens that were yellowish brown with faint whitish or dark brown diamondshaped blotches. Lawson (1993) attributed these color morphs to sexual dimorphism, but Hughes (2000) suggested exceptions are possible. Trape & Mané (2006:168) added, although there are a few exceptions, the differences in coloration are clearly associated with the sex and age of the specimens. Greenbaum & Carr (2005:15) documented the color in life of an adult female from Guinea as “dorsum and flanks were pinkish tan with 39 brown blotches outlined in a creamy tan border along the flanks. The dorsum of the head is brown; the labials are tan with brown outlines, and the venter is white.” Pauwels et al. (2020) documented an adult specimen from Gabon that was uniformly beige in life. Based on photos of a subadult of unknown sex from Mbiliki, Cameroon (Fig. 5G) and a juvenile of unknown sex from Gamba, Gabon (Fig. 5H), the base of the tongue is bluish black, and the forked tip is silvery gray. Hemipenis. Doucet (1963:299) illustrated (Fig. 8) and described the hemipenis of a specimen from Ivory Coast as non-bifid, short and massive [i.e., broad], and flattened. Proximal third covered with spines except at the level of a “mamelon” [nipple] near the root. The distal two-thirds are dimpled, and the apex is flattened. Hemipenes of our examined specimens of T. blandingii had a simple, subcylindrical shape, simple sulcus spermaticus, and spinose ornamentation with a rough apical structure. Diet. In his description of Dipsas globiceps var. tumboensis Müller (1885) noted his specimen from Guinea had a bird’s egg in its stomach. Mocquard (1896) mentioned this species eats lizards in Guinea. Sternfeld (1909) suggested that small birds are the main food of this species in Cameroon, and this contention was repeated by Hughes (2000). Villiers (1950a) noted both species of Toxicodryas in West Africa feed mainly on birds. Villiers (1950b) remarked that one of the listed specimens (either from Ivory Coast or Liberia) had a bird in its stomach. Monard (1951) noted a snake from Cameroon had an Orange Weaver (Ploceus aurantius) in its stomach. Villiers (1951) documented a bird in the stomach of a snake from Benin, and an Agama sp. (sensu Leaché et al. 2017) in the stomach of a snake from Togo. Cansdale (1955:31) suggested that “it is reputed to be an egg-eater in the Gold Coast [Ghana] and has the same Twi name as the true Egg-eating Snake; but I... cannot confirm it from the literature.” Cansdale (1965) repeated the latter remark and added that they will eat eggs in captivity. Dekeyser (1955) cited Villiers (1955), who supposedly noted a large number of bats in a West African T. blandingii, and the identity of these bats was later mistakenly attributed to Lavia frons by Wickler & Uhrig (1969) and Happold & Happold (2013), perhaps because this bat species was mentioned earlier in the same paragraph by Dekeyser (1955). However, Villiers (1955) is a study about parasites in African vertebrates, and there is no mention of Toxicodryas or any bat species. It is likely that Dekeyser (1955) was referring to Villiers (1956), who documented a snake from Guinea that was found in the cracks of a cave’s ceiling, and its stomach contained several bats in the genus Hipposideros. Hughes (2000) reported complaints by local people in Ghana that this species will raid eggs from domesticated poultry, but it is possible that the true culprit was Naja guineensis (sensu Wüster et al. 2018), which is also long and has a tendency to be melanistic in adults. Pitman (1958:84) quoted T.S. Jones from Sierra Leone who shot three snakes as they raided Villa
Phylogeny and biogeography of the African burrowing snake subfamily Aparallactinae (Squamata: Lamprophiidae)
No full textInternational audienc
Evolutionary history of burrowing asps (Lamprophiidae: Atractaspidinae) with emphasis on fang evolution and prey selection
No full textAtractaspidines are poorly studied, fossorial snakes that are found throughout Africa and western Asia, including the Middle East. We employed concatenated gene-tree analyses and divergence dating approaches to investigate evolutionary relationships and biogeographic patterns of atractaspidines with a multi-locus data set consisting of three mitochondrial (16S, cyt b, and ND4) and two nuclear genes (c-mos and RAG1). We sampled 91 individuals from both atractaspidine genera (Atractaspis and Homoroselaps). Additionally, we used ancestral-state reconstructions to investigate fang and diet evolution within Atractaspidinae and its sister lineage (Aparallactinae). Our results indicated that current classification of atractaspidines underestimates diversity within the group. Diversification occurred predominantly between the Miocene and Pliocene. Ancestral-state reconstructions suggest that snake dentition in these taxa might be highly plastic within relatively short periods of time to facilitate adaptations to dynamic foraging and life-history strategies.</div
Ancestral-state reconstructions with ML optimization on the ML trees from the concatenated analyses shown in Fig 2.
No full text(A) fang morphology, (B) dietary preference. Aparallactus 1 = A. niger; Aparallactus 2 = A. modestus; Aparallactus 3 = A. capensis, A. cf. capensis, A. guentheri, A. jacksonii, A. lunulatus, and A. werneri; Amblyodipsas 1 = A. concolor; Amblyodipsas 2 = A. dimidiata, A. polylepis, and A. unicolor; Amblyodipsas 3 = A. ventrimaculata; Amblyodipsas 4 = A. microphthalma.</p
