Lygodactylus kibera, Lobón-Rovira & Bauer & Pinto & Trape & Conradie & Kusamba & Júlio & Cael & Stanley & Hughes & Behangana & Masudi & Pauwels & Greenbaum, 2024

Lobón-Rovira, Javier, Bauer, Aaron M., Pinto, Pedro Vaz, Trape, Jean-Francois, Conradie, Werner, Kusamba, Chifundera, Júlio, Timóteo, Cael, Garin, Stanley, Edward L., Hughes, Daniel F., Behangana, Mathias, Masudi, Franck M., Pauwels, Olivier S. G. & Greenbaum, Eli, 2024, Integrative revision of the Lygodactylus gutturalis (Bocage, 1873) complex unveils extensive cryptic diversity and traces its evolutionary history, Zoological Journal of the Linnean Society (Zool. J. Linn. Soc.) 201 (2), pp. 447-492 : 474-476

LYGODACTYLUS KIBERA SP.NOV.

( Figs 17–18 , Table 3 ; Supporting Information, Fig. S6 ; Table S7)

Zoobank registration: https://zoobank.org/ 26F76FD8-043A-4226-BDD5-E971235D474C

Lygodactylus picturatus gutturalis : Schmidt (1919) [part]; de Witte (1953) [part]; Pasteur, 1965(1964) [part]; Wermuth (1965) [part].

Lygodactylus gutturalis : Röll (2005) [part]; Spawls et al. (2018) [part].

Lygodactylus kibera sp. nov. belongs to a distinctive clade (B1), from the Albertine Rift of Burundi and eastern DRC, which clusters as sister to another candidate new species from the northern Albertine Rift in the Lendu Plateau of DRC and several highlands of Uganda, but differs from it by c. 6.07% in 16S uncorrected p-distance (Table 2), and a lack of nuclear haplotype sharing in RAG1 ( Fig. 2B–C ).

Holotype: UTEP 22566 ( ELI 1145 ), a male with a ventral incision, collected in a village near montane forest at Mpishi , near Kibira National Park, Bubanza Province, Burundi, S03.06974, E29.48445, 1660 m a.s.l. on 20 December 2011 by locals and brought to Eli Greenbaum.

Paratypes (11 specimens): • Burundi (10 specimens) : UTEP 22567–69 ( ELI 1146–48 ), females, and UTEP 22570 ( ELI 1149 ), male with the same collection data as the holotype ; UTEP 22571–73 ( ELI 1195–97 ), males, and UTEP 22574–75 ( ELI 1199–98 ), a male and a female, respectively, collected in a banana field at Mpishi, near Kibira National Park , Bubanza Province, S03.06749, E29.48560, 1705 m a.s.l. on 21 December 2011 by Wandege M. Muninga and Eli Greenbaum ; UTEP 22576 ( ELI 1071 ), female, collected at Bujumbura City , Bujumbura Mairie Province, S03.38236, E29.36419, 811 m a.s.l. on 16 December 2011 by Wandege M. Muninga and Eli Greenbaum . • DRC (one specimen) : UTEP 22586 ( EBG 1556 ), male, collected in a gallery forest at N’Komo River, road. Bukavu-Uvira , South Kivu Province, S02.71471, E28.94641, 1260 m a.s.l. on 15 June 2008 by Maurice Luhumyo, Chifundera Kusamba, Mwenebatu M. Aristote, Wandege M. Muninga, John Akuku, Felix Akuku, Asukulu M’Mema, and Eli Greenbaum .

Diagnosis: A large Lygodactylus [maximum SVL 37.7 mm (mean 34.7 ± 2.5 mm)], that shares a similar distinctive gular chevron ornamentation with the L. gutturalis subgroup. It has 7–9 supralabials and 5–7 infralabials. Dorsal pholidosis with granular scales that become flattened, larger, and imbricate on original tails. Large triangular mental followed by usually three (occasionally two) symmetrical postmental scales (Supporting Information, Fig. S6 ). Nostril never in contact with rostral. Ventral pholidosis with large, flattened, imbricate scales. Five to six terminal scansors on the tail tip. Digits elongated with five terminal scansors on the fourth toe (Supporting Information, Table S7).

This species may be easily differentiated from the L. angularis group by the characteristic ∩ - shaped gular pattern as L. gutturalis (see L. gutturalis diagnosis). It can be differentiated from L. paurospilus , found in the same region, on the basis of three ∩ -shaped thick chevrons reaching the chest vs. two V -shaped broken gular chevrons in paurospilus ; and by having a reduced, almost vestigial postorbitofrontal bone vs. well-developed postorbitofrontal in L. paurospilus . It can also be differentiated from other members of the L. picturatus group based on dorsal coloration and gular pattern (see L. gutturalis account).

Lygodactylus kibera sp. nov. can be differentiated from other species within the L. gutturalis subgroup by subtle morphometric and meristic features. This species is best regarded as cryptic, but we provide some characters that are diagnostically useful. Lygodactylus kibera sp. nov. differs from L. dysmicus by its larger size (maximum SVL 37.7 mm vs. 27.6 mm); gular patterning always with three ∩ - shaped thick chevrons reaching the chest (vs. two thinner ∩ -shaped chevrons, that never extend beyond the posterior part of the lower jaw); fewer precloacal pores (7–8 vs. 9); usually three symmetrical postmental scales vs. two; nostril never in contact with rostral scale (vs. nostril contacting rostral); and lower number of ventral scales across the body (16–18 vs. 21 in L. dysmicus ). It can be distinguished from L. gutturalis s.s. by its slightly larger size [maximum SVL 37.7 mm (mean 34.7 ± 2.5 mm) vs. 36.2 mm (mean 33.3 ± 1.8 mm)]; snout proportionally narrower ( IN / HW 0.20–0.27 vs. 0.29–0.34) with usually one large internasal scale vs. two smaller internasal scales (this character has shown to be variable in both species). It also differs from L. gutturalis and L. depressus based on gular pattern always having three thick ∩ -shaped chevrons reaching the chest [vs. two or three (in L. gutturalis ), or one or two (in L. depressus ) thinner ∩ -shaped chevrons that never extend beyond the posterior part of the lower jaw]; proportionally more elongated head (HL/ SVL 0.27–0.30 vs. 0.24–0.26) with larger eyes ( OD /HL 0.24–0.27 vs. 0.20–0.23) than L. depressus ( Fig. 4 ; Table 3 ). The new species occurs in mid-elevation moist forest, agricultural fields, and human habitations, vs. lowland dry sub-Saharan savannah ( L. gutturalis ) and lowland rainforest of the Congo Basin ( L. dysmicus ; see Fig. 6 ). For a distinction with other species described below, see their respective diagnoses.

Etymology: The name ‘ kibera ’ derives from the word ‘kibira’ or ‘kibera’ in Kinubi—a Sudanese Arabic-based creole language spoken in some regions of Burundi, Kenya, and Uganda —that means ‘forest’, the main habitat type associated with the species.

Description of the holotype ( Fig. 17 ): Measurements and meristic characters of the holotype are presented in Supporting Information, Table S7. Adult male, with a snout–vent length ( SVL) of 36.9 mm and a slightly larger original tail length (TL = 40.6 mm). Body slender, nape moderately distinct. Head as broad as body, and moderate head length ( HW /HL 0.70). Canthus rostralis not prominent. Eye diameter 2.5 mm, with circular pupil. Ear to eye distance slightly larger than orbit diameter (3.7 mm). Snout rounded and slightly pointed. Frontal granular scales larger than occipital scales. Dorsal scales granular from rostral to tail. Rostral undivided, in contact with 1st supralabial, prenasals, and one large internasal scale. Eight to nine supralabials and six infralabials. Prenasal scale in contact with 1st supralabial. Nostril circular, bordered by 1st supralabial, prenasal, one supranasal, and one postnasal. Four rows of scales between supralabials and the orbit. Mental large, triangular, and rounded posteriorly, with two large rounded postmental scales separated by one small rounded postmental scale. Five post-postmental scales. Gular scales granular, rounded, and slightly smaller than ventral scales. Ventral scales large, imbricate, with 17 scales rows across the venter. Body relatively robust and slightly elongated ( TRL / SVL 0.43). Tail with 51 enlarged transverse scales and six pairs of terminal scansors on the tip. Seven precloacal pores. Fore- and hind limbs moderately short, stout; forearm medium-sized ( FL / SVL 0.14); tibia short (CL/ SVL 0.16). Digits elongated and unwebbed with 5–6 terminal scansors. Thumb rudimentary with a small claw. Relative length of digits: I <II = V <III <IV (manus); I <II < V <III <IV (pes).

Coloration: In life ( Fig. 18 ), dorsal coloration brownish grey with light cream-beige lines from nape to tail on each side of the dorsum interspersed by five or six lighter cream dots surrounded by black flanks. Black line from nostril to the anterior insertion of the forelimb. Gular region with white coloration and three black ∩ -shaped chevrons. First and second chevrons in contact. Venter uniform orange from second chevrons to anterior portion of the tail and extending onto the hind limbs. Ventral surface of tail tip and digits whitish. In preservative (holotype; Fig. 17 ) dorsum dark brown and venter with uniform light cream coloration.

Variation: The meristic characters of the head and body of this species are variable (see Supporting Information, Table S7). Coloration of this species seems to be consistent, with the gular coloration slightly lighter in females. First and second gular chevrons may be in contact or not (Supporting Information, Fig. S6 ). Habitat and distribution ( Figs 6 , 18 ): Lygodactylus kibera sp. nov. is known from the Albertine Rift adjacent to Lake Tanganyika in Burundi and DRC. Specimens from Mpishi, Burundi, were collected near montane forest in banana fields. The specimen from Bujumbura, Burundi, was collected inside a hotel in the middle of the city. The specimen from N’Komo River, DRC, was found on a tree near a small river between secondary gallery forest and grassland.

Natural history: An arboreal species with diurnal habits frequently found on the trunk or branches of trees, and also around anthropogenically altered areas such as buildings and plantations.

Figures

Figure 2. A, Multilocus species tree and Bayesian chronogram of divergences among Afro-American Lygodactylus clade (Clade C of Gippner et al. 2021), based on an alignment of 13 concatenated markers (including eight nuclear-encoded genes, four protein-coding mitochondrial genes, and the 16S mitochondrial rRNA gene, see Gippner et al.2021). Branches with posterior probability> 0.9 are denoted by asterisks (*) at relevant nodes. Blue bars depict 95% High Probability Density (HPD) intervals of estimated divergence dates.B, Median-joining nuclear allele network showing the relationships between the L. picturatus-fischeri groups, inferred from the RAG1 nuclear gene (262 bp including all available RAG1 sequences of the L. picturatus-fischeri groups).C, Median-joining nuclear allele network of the RAG1 nuclear gene (980 bp) showing the relationships within the L. gutturalis subgroup, excluding L. gutturalis s.s. In all networks, circle frequency of alleles and small lines represent mutational steps.

Figure 4. A, PCA plots of the first principal component (PC1) vs. the second principal component (PC2). Species are represented by different colours included in the legend. B, Boxplots (top whisker–maximum value; lower whisker–minimum value; dark horizontal line–median) of the body size (SVL) and linear measurements analysed in this study.Values in all the variables with the exception of SVL, are represented using the Least-squares means interaction plot (function Lsmip, package lsmeans). For abbreviations and further details see Material and methods section.

Figure 6. Geographical distribution of the L.gutturalis complex in Africa, on a hybrid map of the major vegetation divisions (Sayre et al. 2013) and 1 arc-second elevation map across tropical Africa (NASA 2000). Different colours depict records of different species within the L. gutturalis subgroup; see inset for explanations of symbols.White circles denote records of L. cf. gutturalis that cannot be assigned to any of the nominate taxa. Black stars represent the type localities of each nominate taxon within the L. picturatus subgroup (1– L. picturatus, 1S– L. picturatus sudanensis, 1U– L. picturatus ukerewensis, 2– L. inexpectatus, 3– L. keniensis, 4– L. kimhowelli, 5– L. manni, 6– L. mombasicus, 7– L. scorteccii, 8– L. tsavoensis, 9– L. wojnowskii, 10– L. williamsi).

Figure 17. Holotype of Lygodactylus kibera sp. nov. (UTEP 22566) from Mpishi, near Kibira National Park, Bubanza Province, Burundi. A, Dorsal and ventral view of body. B, Detailed view of head in dorsal, ventral, and lateral views (from top to bottom). Details of cloacal region (C), (D) tail tip, and (E) left manus.

Figure 18. Specimens of Lygodactylus kibera sp. nov. in life or freshly euthanized from (A) Mpishi (holotype: UTEP 22566), near Kibira National Park, Bubanza Province, Burundi, (B) N'Komo River (UTEP 22586), South Kivu Province, DRC, and (C) Bujumbura city, Bujumbura Mairie Province (UTEP 22576), Burundi.Photo of habitat at (D) Mpishi, Bubanza Province, Burundi. Photographs by E.G. (A–C) and C.K. (D).