Rhampholeon bombayi Hughes & Behangana & Lukwago & Menegon & Dehling & Wagner & Tilbury & South & Kusamba & Greenbaum 2024, sp. nov.

Rhampholeon bombayi sp. nov. Hughes, Dehling, Menegon, Kusamba, and Greenbaum

Bombay’s pygmy chameleon urn:lsid:zoobank.org:act: DA3436FD-9EE8-4ED8-BC25-A1BC771C1CE8

Synonymy.

Rhampholeon boulengeri — de Witte 1965 (partim), Fischer & Hinkel 1993, Hinkel 1996, Tilbury & Tolley 2015 (partim), Spawls et al. 2018 (partim), Tilbury 2018 (partim)

Rhampholeon sp. 3 — Hughes et al. 2018

Etymology. The specific epithet honors Sidi Mubarak Bombay, an explorer of the waYao tribe who guided, led, and interpreted for the expeditions of both Speke and Burton to discover the source of the Nile, helped Stanley find Livingstone, and with Cameron, became the first known African to cross the African continent from east to west (Millard 2022). Bombay was sold into slavery as a child and ended his life as the greatest African explorer of all time.

Holotype. UTEP 21701 (field no. ELI 602), adult female, DRC, South Kivu Province, near Kalundu, 03.15552° S, 28.42108° E, 1482 m elevation, 21 December 2010, collected by E. Greenbaum, C. Kusamba, M.M. Aristote, and W.M. Muninga (Fig. 12A).

Paratypes. Two adult females, UTEP 21702, 22722 (field nos. ELI 617–618), DRC, South Kivu Province, Kalundu / Mwana River, 03.15786° S, 28.4273° E, 1455 m elevation, 22 December 2010, collected by E. Greenbaum, C. Kusamba, M.M. Aristote, and W.M. Muninga. One adult male, UTEP 22721 (field no. ELI 598), DRC, South Kivu Province, vicinity of Kalundu, 03.15387° S, 28.42487° E, 1525 m elevation, 21 December 2010, collected by E. Greenbaum, C. Kusamba, M.M. Aristote, and W.M. Muninga. Four adult females, UTEP 22674, 21703, 22675– 22676 (CRSN HERP 2920, 2984–2986), DRC, South Kivu Province, Mwana Kisanga, 03.15146° S, 28.44403° E, 1529 m elevation, November 2014, collected by C. Kusamba. One adult male and three adult females, UTEP 22677–22680 (CRSN HERP 2987–2988, 2995–2996), DRC, South Kivu Province, Mwana Kisanga, 03.15678° S, 28.43448° E, 1450 m elevation, November 2014, collected by C. Kusamba. One adult female, UTEP 22681 (CRSN HERP 2999), DRC, South Kivu Province, Hill Nkala Summit, 03.16036° S, 28.43740° E, 1555 m elevation, November 2014, collected by C. Kusamba.

Referred specimens. UTEP 22689, 22692, 22693–22696 (field nos. JMD 2014-53, 2014-101, 2014-103 – 106), RWANDA, Western Province, Nyungwe National Park, Kamiranzovu Swamp (02.46573° S, 29.15917° E, 2213 m elevation) (6 specimens). UTEP 22690–22691 (field nos. JMD 2014-55 – 56), RWANDA, Western Province, Nyungwe National Park, Kamiranzovu Swamp (02.47350° S, 29.16647° E, 2330 m elevation) (2 specimens) (Fig. 12 D-F). MTSN 7075–7076, RWANDA, Western Province, Nyungwe National Park (02.56753° S, 29.23079° E, 2003 m elevation) (2 specimens). MTSN 7123, RWANDA, Western Province, Cyamudongo Forest (02.54529° S, 28.98507° E, 2033 m elevation) (1 specimen). UTEP 22697 (field no. JMD 2014-107), RWANDA, Western Province, Cyamudongo Forest (02.53851° S, 28.99328° E, 1857 m elevation) (1 specimen). UTEP 21706, 22724– 22726 (field nos. ELI 739–741, 796), DRC, South Kivu Province, Itombwe Plateau, vicinity of Tumungu (03.53545° S, 28.67411° E, 1835 m elevation) (4 specimens) (Fig. 12C). UTEP 21704–21705 (field nos. EBG 1286, 1346), DRC, South Kivu Province, vicinity of Irangi (*collection notes indicate that these specimens were collected at a nearby location by a local person [see Hughes et al. 2018]) (2 specimens) (Fig. 12B). UTEP 21708 (field no. EBG 1189), DRC, South Kivu Province, Tshibati (02.22640° S, 28.77940° E, 2030 m elevation) (1 specimen). MTSN 1640–1641 (field nos. D.C. Moyer 1640–1641), DRC, South Kivu Province, Kabobo Mountains, Kizamba River (05.38097° S, 29.19563° E, 1904 m elevation) (2 specimens). UTEP 22682 (field no. MUSE 10154), DRC, South Kivu Province, Itombwe Plateau, Atuyaumbu (03.56424° S, 28.25707° E, 1582 m elevation) (1 specimen). UTEP 21707, 22723 (field nos. MUSE 10146–10147), DRC, South Kivu Province, Itombwe Plateau, Mabwe (03.60583° S, 28.34308° E, 1582 m elevation) (2 specimens).

Diagnosis. Rhampholeon bombayi sp. nov. is in the subgenus Rhinodigitum because of its distinctly bicuspid claws, prominent rostral process, smooth plantar surfaces, and phylogenetic placement, thus easily distinguishing it from the six species in the other two subgenera (i.e., Rhampholeon and Bicuspis): R. gorongosae, R. marshalli, R. spectrum, R. spinosus, R. temporalis, and R. viridis. Rhampholeon bombayi sp. nov. can be distinguished from all other Rhampholeon species by the following combination of traits: (1) lack of prominent mite pockets in the inguinal region distinguishes it from R. beraduccii, R. platyceps, R. chapmanorum, R. maspictus, R. tilburyi, R. bruessoworum, and R. nebulauctor; (2) presence of prominent mite pockets in the axillary region distinguishes it from R. nchisiensis and R. acuminatus; (3) distinct supra-orbital and canthal crests distinguishes it from R. hattinghi; (4) geographic restriction to the Albertine Rift distinguishes it from R. uluguruensis, R. moyeri, R. colemani, R. sabini, R. rubeho, R. nicolai, R. waynelotteri, and R. princeeai; (5) shorter tail length in males, genetic divergence, and non-overlapping elevational range at parapatric sites distinguishes it from R. boulengeri; (6) larger mean body size, longer snout length, and larger eye diameter in females, and longer inter-limb length in both sexes distinguishes it from R. monteslunae sp. nov.; (7) slightly shorter head, snout, and mouth lengths, smaller eye diameter, shorter inter-limb and hind limb lengths in females distinguishes it from R. nalubaale sp. nov.; (8) genetic divergence and non-overlapping elevational range at sites of co-occurrence distinguishes it from R. plumptrei sp. nov. and R. msitugrabensis sp. nov.

Genetic differentiation and variation. A summary of pairwise sequence divergence for three DNA markers (16S, ND2, and RAG-1) among individuals of R. bombayi sp. nov. and other Rhampholeon species is presented in Supplementary Material 1.

Description of holotype (UTEP 21701). Adult female, SVL 54.1 mm and TL 11.0 mm. Body shape leaf-like. Casque flattened, with short head. Neck indistinct from head. Supra-orbital crests distinct with cluster of tubercles connected by a ridge with 13 tubercles across casque and 20 tubercles from peak-to-peak of crests. Rostral process 1.97 mm, composed of elongated tubercles. Temporal crest discrete with several enlarged tubercles extending posteriorly from mid-eye. Nares open in a posterior orientation. Canthal ridge consists of raised tubercles, one raised higher than others near snout. Ninety-eight upper and 98 lower labial tubercles present along tip of snout to rictus of mouth. Body covered in nearly homogenous, flattened tubercles. Several larger conical tubercles present on dorsal flanks around midbody. Crenulated dorsal crest, more prominent from mid-body to nape. Several enlarged conical tubercles present on limbs. Claws markedly bicuspid.

Coloration of holotype (in life). A photograph of the holotype is presented in Figure 9A. Background color light brown to gray with a darker brown hue. Limbs, tail, and top of head a darker brown color than body. Gular region white, and this color extends from tip of chin to ventral area and towards cloaca. One prominent diagonal dark brown line extends from near dorsal crest posteriorly to lateral flank, with a less prominent line posterior to that, which together resemble veins on a leaf. Several small yellow tubercles present on upper lateral flanks. Largest body tubercle near neck dark brown to black.

Variation. A summary of descriptive morphometrics for R. bombayi sp. nov. is presented in Table 3, comparative boxplots in Figure 4, and measurements of the type specimens in Table 7. Photographs displaying color variation in life are presented in Figure 12. Morphological proportions are generally consistent with those of the holotype. Males have smaller body sizes (M: mean 41.5 mm, range 31.0– 47.6 mm, n = 14; F: mean 47.9 mm, range 40.8–59.1 mm, n = 22) and longer tails than females (M: mean 12.9 mm, range 9.3–18.9 mm, n = 14; F: mean 11.4 mm, range 10.0– 13.4 mm, n = 22). Body coloration is consistently brown to tan and/or gray with lighter orange, yellow, or red hues. One, sometimes two or three, dark brown to red lines on the lateral flanks extend diagonally from the dorsal crest toward the hind limbs, resembling veins of a leaf. Legs and tail coloration can be markedly darker brown than the body. Small tubercles, especially on the eyelids, dorsal crest, and legs can often be yellowish.

Reproduction. Seven females were gravid (UTEP 21703, UTEP 22674, UTEP 22679, UTEP 22680, UTEP 22689, UTEP 22692, and UTEP 22694). Gravid females had a mean SVL of 49.1 mm (44.4–55.1 mm) and a mean TL of 11.7 mm (10.0– 13.4 mm). Most of these females were collected in November (n = 4). Two females had clutch sizes of 4 eggs each, and one female had a clutch size of 3 eggs. The smallest specimen examined (UTEP 21708) was collected 19 August 2007 with SVL 18.4 mm and TL 4.2 mm from Tshibati, South Kivu Province, DRC.

Distribution, natural history, and conservation. Rhampholeon bombayi sp. nov. is found in montane forests at an elevation range of 1450–2330 m. Specimens were collected from within forests at heights of approximately 1 m above the ground. Specimens have been collected from Kahuzi-Biega National Park, Kabobo Natural Reserve, and Itombwe Natural Reserve, DRC, and Nyungwe Forest National Park, Rwanda. See Greenbaum & Kusamba (2012) for detailed information on the conservation of Itombwe’s herpetofauna, and Greenbaum (2017) for a description of the overall threats facing the herpetofauna of eastern DRC. Behavior and activity patterns are unknown, but likely similar to that described for R. boulengeri (sensu lato) (Spawls et al. 2018; Tilbury 2018). Other lizard species collected from montane forest in the Itombwe Plateau included Congolacerta vauereselli, Holaspis cf. guentheri, Leptosiaphos blochmanni, L. graueri, Trachylepis varia, Trioceros johnstoni, and T. schoutedeni.