Common name

Hughes, Daniel F., Kusamba, Chifundera, Behangana, Mathias & Greenbaum, Eli, 2017, Integrative taxonomy of the Central African forest chameleon, Kinyongia adolfifriderici (Sauria: Chamaeleonidae), reveals underestimated species diversity in the Albertine Rift, Zoological Journal of the Linnean Society 181, pp. 400-438 : 413-417

Common name : Rugege Highlands forest chameleon.

Holotype: UTEP 21485 (field no. ELI 1156 ), adult female, BURUNDI, Bubanza Province, near Kibira National Park, Mpishi village , 03°4′ 11.064″S 29°29′4.02″E, 1660 m elevation, 20 December 2011, collected by E. Greenbaum, C. Kusamba, M.M. Aristote, and W.M. Muninga ( Fig. 6A).

Paratopotypes: Same collection details as holotype, one adult male, UTEP 21481 (field no. ELI 1155) ( Fig. 6B), and another adult male, UTEP 21482 (field no. ELI 1238), collected on 23 December 2011 ( Fig. 6C).

Paratypes: One adult female, UTEP 21483 (field no. ELI 1220 ), BURUNDI, Bubanza Province, Kibira National Park, Mpishi village , 03°3′42.372″S 29°29′36.348″E, 1986 m elevation, 22 December 2011, collected by same collectors of holotype ; one adult female, UTEP 21484 (field no. ELI 1256 ), BURUNDI, Kayanza Province, Kibira National Park, near Rwegura village , 02°56′20.292″S 29°29′54.78″E, 2130 m elevation, 25 December 2011, collected by E. Greenbaum, M.M. Aristote, and W.M. Muninga ( Fig. 6D) .

Diagnosis: Kinyongia rugegensis sp. nov. can be distinguished from all other Kinyongia species by the following combination of traits: (1) lack of rostro-nasal ornamentation in both sexes; (2) moderate body size (mean SVL = 55.9 mm); (3) anterior dorsal keel with 8–10 conical tubercles; (4) a slightly elevated casque that tapers posteriorly to a prominent apex; (5) absence of a gular and ventral crest; (6) 16–18 upper and 15–17 lower labials; (7) generally flat shape of the upper casque; (8) tail length longer than SVL in both sexes; (9) indistinct parietal crest with slightly raised tubercles; (10) background body coloration in adult females generally green to yellow-green with darker pigmented regions on the flanks and tail; background body coloration in adult males generally brown with tan and yellow speckling on the flanks; (11) interstitial skin between the tubercles of the body generally black for both sexes; (12) a light brown stripe passes through the middle of the eye and extends from the canthal ridge to the temporal crest; (13) top of the head is typically a darker green/brown colour than elsewhere; (14) the gular region is distinctly lighter in colour, with a combination of green, white and tan.

Differential diagnosis: A medium-sized forest chameleon that is distinguished from most congeners by the absence of a rostral process in both sexes [ K. asheorum , K. boehmei ( Lutzmann & Nečas, 2002) , K. carpenteri , K. fischeri , K. magomberae Menegon et al. (2009) , K. matschiei ( Werner, 1895) , K. msuyae Menegon et al. (2015) , K. multituberculata ( Nieden, 1913) , K. oxyrhina ( Klaver & Böhme, 1988) , K. tavetana ( Steindachner, 1891) , K. tenuis ( Matschie, 1892) , K. uluguruensis ( Loveridge, 1957) , K. uthmoelleri ( Müller, 1938) , K. vanheygeni Nečas, 2009 , K. vosseleri ( Nieden, 1913) and K. xenorhina ]. The new species can be distinguished from K. adolfifriderici by its larger cranial crest gap and head length, larger body size (52.8–58.7 mm vs. 47.9–54.9 mm), and more upper (16–18 vs. 10–14) and lower labials (14–17 vs. 12–15). The new species can be distinguished from K. tolleyae sp. nov. by the lack of two distinctly bulging and rounded portions of the upper casque, slightly larger head width, larger fleshy papillae medial to rotulae on hemipenis and more upper labials (16–18 vs. 13–17). The new species can be distinguished from K. itombwensis sp. nov. by its larger fore- and hind limbs, slightly larger cranial crest gap and head length, and more conical tubercles on the dorsal crest (8–10 vs. 6–7). The new species can be distinguished from K. mulyai and K. excubitor by the presence of a dorsal crest with 8–10 conical tubercles and marked mitochondrial sequence divergence. The new species can be distinguished from K. gyrolepis by its smaller mean body size (55.9 vs. 67.3 mm) and current distribution in moist Afromontane rainforest.

Genetic differentiation and variation: A summary of pairwise sequence divergence for each molecular marker (16S, ND2 and RAG 1) among individuals of K. rugegensis sp. nov. and other species of Kinyongia endemic to the AR are presented in Table S2. For the ND2 locus, p -distances among K. rugegensis sp. nov. samples ranged from 0.1 to 0.7%.

Description of holotype: Adult female, SVL 56.6 mm and TL 67.3 mm. Four oviductal eggs present (see Reproduction in the following text). Casque low, slightly raised above nape. Distinct, elevated apex on posterior casque. Neck distinct from head. Parietal crest largely indistinct with few enlarged and flattened tubercles in an inconsistent pattern. Supra-orbital ridges mostly smooth. Temporal crest comprises three enlarged tubercles extending posteriorly from mid-eye and ascending along posterior ridge of casque to its apex. Nares open laterally and in a posterior orientation. Canthal ridge consists of four slightly raised tubercles, one raised higher than others near snout. Sixteen upper and 14 lower labials are present along tip of snout to posterior margin of orbit. No gular or ventral crests present. Nine small conical tubercles present on anterior portion of dorsal crest, absent near mid-body. Tail and lateral flanks smooth. Body covered in nearly homogenous, flattened tubercles. Some larger polygonal tubercles present on dorsal flanks. Patches of small tubercles in rosette patterns on ventral flanks. Some enlarged flattened tubercles present on outer portions of limbs. Claws typical of Kinyongia species.

Coloration of holotype (in ethanol): Photographs of the body and head detail of the holotype (in preservative) are presented in Fig. 10. The background coloration is greyish blue with some darker blotches on the flanks and tail. Patches of lighter blues and greens are present near the anterior portions of the body, and the sides of the head and tail. Light yellow (almost white) patches occur near axillary and inguinal regions and a few places on the lateral body flanks. The soles of the feet are yellowish-white.

Coloration of holotype (in life): A photograph of the holotype (in life) is presented in Fig. 6A. The top of the head is covered in brown and dark green tubercles with black interstitium. Beginning below the temporal crest, the head is lighter green in colour and covered in yellowish-green tubercles with powder-blue interstitium. At mid-eye, there is a dark brown lateral stripe that connects the coloration on the canthal ridge to the temporal crest. Near the tip of the snout is a pronounced yellow coloration that fades posteriorly.The background coloration of the body is yellowish-green with black interstitium. The powder-blue coloration of the head interstitium extends posteriorly on the ventral flanks and gradually changes to black by mid-body, then blue reappears on the posterior third of body. Tubercles on the venter, near axillary and inguinal regions, and hidden parts of the limbs, are off-white with flecks of green. The dorsal crest is adorned with darker green tubercles than elsewhere on the body and this coloration extends onto the tail. The posterior third of the tail is darker brown and the greenish coloration of the tail in general is less bright compared to the body. Differential distribution of interstitial coloration (light blue or black) on the body form broad vertical dark brown bands.

Hemipenis: Hemipenal drawings and description are based on specimen UTEP 21481. Line drawings depicting the general hemipenis morphology of K. rugegensis sp. nov. are presented in sulcal and lateral views ( Fig. 7A). Hemipenes are calvate and the pedicel is less than one-fifth of the hemipenis length. The truncus is covered with calyces ranging in size from smaller on the asulcal apex to larger ones near the asulcal pedicel. Distal calyces are smaller and more hexagonal in shape. The sulcal lips and sulcus spermaticus are smooth and devoid of ornamentation. The flesh on the sulcus is highly envaginated (folded), forming numerous sulcal ridges. Sulcal lips diverge towards the apex and continue as a ridge that encircles the apex. The apex is bilobed and each lobe possesses a large, sharply denticulated rotulae. A sizeable protuberant fleshy papilla is positioned medially from each rotulae.

Variation: Descriptive morphometrics of K. rugegensis sp. nov. are presented in Table 4, and a summary of mean measurements in Table 3. Chameleon photographs displaying colour variation in life are presented in Fig. 6. Morphological proportions in paratopotypes and paratypes are generally consistent with those in the holotype. Males have longer tails than females [M: 85.3 ± 6.9 (80.4–90.2 mm, n = 2); F: 66.7 ± 0.9 (65.6–67.3 mm, n = 3)] (P <0.01), but similar body sizes [M: 56.9 ± 2.6 (55.0– 58.7 mm, n = 2); F: 55.4 ± 2.3 (52.8–56.8 mm, n = 3)] (P> 0.05). Males have overall yellowish-brown background coloration, in contrast to the lighter green colour of females. When agitated, the tip of the snout, eye skin and various regions on the flanks can be brightly coloured with yellow. One female ( UTEP 21484) in an aggressive posture and coloration with an open mouth, showed a dark patch laterally at mid-body, white gular and ventral regions and bright yellow areas on the head ( Fig. 6D).

Reproduction: The holotype ( UTEP 21485) with SVL 56.6 mm and TL 67.3 mm collected on 23 December 2011 was gravid. This individual contained four oviductal (shelled) eggs with mean dimensions (in mm), length 12.75 ± 0.21 (range: 12.49–12.93) and width 6.49 ± 0.25 (range: 6.31–6.84). Exact measurements of eggs were as follows: 12.65 L × 6.84 W; 12.49 L × 6.32 W; 12.91 L × 6.49 W; 12.93 L × 6.31 W. This individual had moderate fat bodies. Another female ( UTEP 21484 ) with SVL 52.8 mm and TL 55.6 mm collected on 25 December 2011 was also gravid. This individual contained four enlarged, yolked ovarian follicles with mean dimensions (in mm), length 7.09 ± 0.19 (range: 6.83–7.25) and width 5.48 ± 0.26 (range: 5.2–5.83). Exact follicular measurements were as follows: 7.25 L × 5.43 W; 7.19 L × 5.2 W; 6.83 L × 5.83 W; 7.09 L × 5.44 W. This individual possessed extensive fat bodies. Conversely, a female ( UTEP 21483 ) with SVL 58.8 mm and TL 67.1 mm collected on 22 December 2011 was not gravid, as demonstrated by the largest ovarian follicles measuring <3 mm in diameter and lacking evidence of yolk. This individual had minor fat bodies .

All males had darkly pigmented testes (i.e. black coloration), which is characteristic of all chameleon species examined to date ( Tolley & Herrel, 2013). All collected males were sexually mature. One male ( UTEP 21481 ) with SVL 55.0 mm and TL 80.4 mm collected on 20 December 2011 had enlarged testes. The right testis of this individual measured 6.89 mm in length and 5.04 mm in width. Another male ( UTEP 21482 ) with SVL 58.7 mm and TL 90.2 mm collected on 23 December 2011 also had enlarged testes. The right testis of this individual measured 6.45 mm in length and 4.51 mm in width. Fat bodies were minor for both of these individuals .

Diet: All five specimens examined for gut contents had remains of arthropod prey items that could be identified to order. The stomach of one female ( UTEP 21484) contained Hemiptera, Lepidoptera , Hymenoptera and Coleoptera. A second female ( UTEP 21485) stomach contained Diptera, Hemiptera, Araneae and Acari. The stomach of a third female ( UTEP 21483) contained Araneae, Orthoptera and Hemiptera. A male ( UTEP 21481) stomach contained Diptera and Hemiptera. Another male ( UTEP 21482) stomach contained Diptera, Hemiptera, Araneae and Psocoptera.

Distribution and natural history: Kinyongia rugegensis sp. nov. is found in moist Afrotemperate montane and sub-montane forests at an elevation range from 1660 to 2130 m. Most specimens were collected from forest edges near and inside Kibira National Park. This montane forest extends from southern Rwanda (Nyungwe Forest National Park) to northern Burundi (Kibira National Park). We speculate that this new species is present throughout the Rugege Highlands

*Enlarged ovarian follicles present in body cavity of specimen.

in areas of suitable forest habitat. For example, Hinkel (1993) recorded Bradypodion adolfi -friderici (= Kinyongia adolfifriderici ) from both Nyungwe and Cyamudongo forests (= Nyungwe Forest National Park) in Rwanda, and these records potentially represent this new species. Two specimens ( UTEP 21482 and UTEP 21483) were collected inside a banana tree plantation just outside the national park. One specimen ( UTEP 21484) was collected from natural roadside vegetation, and was found c. 2.5 m above ground in a small tree. Two of the three females were gravid, and both males were sexually mature. No juveniles were detected during the search period (c. 3.5 weeks). Behaviour and activity patterns are essentially unknown, but likely similar to that of K. adolfifriderici ( Tilbury, 2010) . Other lizard species collected near the type locality included typical AR lizard fauna, including Adolfus africanus , Chamaeleo dilepis , Congolacerta vauereselli , Hemidactylus mabouia , Lygodactylus cf. gutturalis , Rhampholeon boulengeri , Trioceros ellioti , T. johnstoni , Trachylepis striata and T. maculilabris .

Conservation: Nyungwe Forest National Park is the largest protected area in Rwanda, and Kibira National Park is the largest protected area in Burundi. These contiguous parks together form one of the largest montane forest blocks in eastern Africa ( Barakabuye et al., 2007). However, despite this high level of connectivity, similarity of threats and biodiversity importance, these forests have been managed in near isolation to the neighbouring protected areas ( Barakabuye et al., 2007). The highlands of these countries are renowned for their nutrient-rich soils. As a result, the regions are burdened with extremely dense human populations that greatly threaten the biological integrity of the remaining forests with severe agricultural pressures. Moreover, a longstanding history of armed conflict in the region has left a legacy of irreparable anthropogenic damage in these fragile ecosystems ( Kanyamibwa, 1998).

Etymology: The specific epithet is derived from Rugege Highlands, the greater mountainous region where the species was collected, with the Latin suffix – ensis denoting a place or locality. Although the holotype was collected from Kibira National Park in northern Burundi, this Afromontane forest is contiguous with Nyungwe Forest National Park in Rwanda via the Rugege Highlands. This new species likely occurs in suitable forested habitat across this mountain range. The view that these neighbouring protected areas are independent is outdated, and unfortunately, park management in bordering countries has sustained this position for some time ( Barakabuye et al., 2007). Thus, we felt that the taxonomy should reflect the natural connectivity of the region and chose a broader name accordingly.