Lygodactylus leopardinus, 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 : 481-483

LYGODACTYLUS LEOPARDINUS SP.NOV.

( Figs 22–23 , Table 3 ; Supporting Information, Table S9)

Zoobank registration: https://zoobank.org/ 9010CCA9-953F-4091-9D1E-40BBFD0B9FF8

Lygodactylus leopardinus sp. nov. is the rainforest member within subgroup C, and the most morphologically distinguishable species within the L. gutturalis subgroup. It also differs by a minimum of c. 10.10% for 16S uncorrected p-distance (Table 2) from related taxa, and it lacks nuclear haplotype sharing in RAG1 ( Fig. 2B–C ).

Holotype: UTEP 22577 ( ELI 2330 ), male with broken tail and ventral incision, collected at Balolombo Village , Busira River, Équateur Province, DRC, S00.25939, E19.63575, 305 m a.s.l. on 18 July 2013 by Chifundera Kusamba, Wandege M. Muninga, Mwenebatu M. Aristote, and Eli Greenbaum.

Paratype: UTEP 22596 ( ELI 3584 ) , female, collected from Katopa, ICCN Camp , Maniema Province, DRC, S02.74769, E25.10323, 450 m a.s.l. on 2 July 2015 by Eli Greenbaum .

Additional material: RMCA 1981.065 . R.0004, male with original tail, collected at River Lotende (canopy), Mabali, Équateur Province, DRC, between 17 and 18 August 1955 by Raymond Laurent ; RMCA R.2947, female, collected at Basongo , Kasai Province, DRC, on 27 July 1921 by Henri Schouteden ; RMCA R.16752, male, collected at Yokamba (Boende), Tshuapa Province, DRC, in 1953 by J. Stevenart .

Diagnosis: This species represents a moderately sized Lygodactylus [maximum SVL 34.7 mm (mean 33.6 ± 1.3 mm)], with males having the most distinctive gular patterning of the L. gutturalis group (described below, Fig. 3 ). Females without gular pattern. Eight supralabials and 6–7 infralabials. Males with seven precloacal pores.

This species can be differentiated from all taxa of the angularis group taxa by the same osteological differences described above (e.g. postorbitofrontal reduced almost vestigial vs. well developed in the angularis group) and a unique leopard-like pattern on the dorsum. Also, it can be easily differentiated from L. angularis and L. baptistai based on the gular patterning (broken pattern vs. V-shaped pattern in L. angularis and the unique pattern of L. baptistai described above; see Fig. 5 ). The species can be confused with L. heeneni and L. paurospilus based on the broken gular ornamentation; however, it can be differentiated from them as follows: unique leopard-like dorsal pattern in L. leopardinus sp. nov. (vs. light brown dorsal background with a light cream vertebral line of blotches in L. heeneni ( Fig. 7 ), similar to L. paurospilus (based on preserved specimens; Fig. 8 ; Supporting Information, Fig. S4 ); broken pattern that converges anteriorly (vs. a broken pattern that converges posteriorly (V -shaped) in L. heeneni and L. paurospilus ); fewer precloacal pores than L. heeneni (7 vs. 9–10); and osteological features described above.

Lygodactylus leopardinus sp. nov. is the most distinctive species within the gutturalis subgroup and the only species within the picturatus group with a broken gular pattern. It can be easily distinguished from other closely related species based on its unique gular pattern and the leopard-like dorsal pattern, which is not present in related taxa. Females lack gular patterning (vs. weakly present in other members of the gutturalis subgroup). It can also be differentiated based on minor meristic differences as follows: greater number of scales between the eyes (30–36 vs. 17–24 in L. gutturalis , 25 in L. dysmicus , 19–28 in L. kibera sp. nov., 19–22 in L. karamoja sp. nov., and 19 in L. mirabundus sp. nov.); and lower number of ventral scales at midbody (14–15 vs. 15–20 in L. gutturalis , 21 in L. dysmicus , 16–19 in L. depressus , 16–18 in L. kibera sp. nov., 16–18 in L. karamoja sp. nov., and 21 in L. mirabundus sp. nov.).

Etymology: The name ‘ leopardinus ’ is an adjective referring to the leopard-like dorsal pattern present in males of this species.

Description of the holotype ( Fig. 22 ): Measurements and meristic characters of the holotype are presented in Supporting Information, Table S9. Table 3 . Adult male, with a snout–vent length (SVL) of 32.4 mm and a broken tail. Body slender and nape slightly distinct. Head as broad as body, and moderate head length (HW/HL 0.63). Canthus rostralis not prominent. Eye diameter 2.0 mm with circular pupil. Ear to eye distance almost twice the orbit diameter (3.4 mm). Snout rounded and moderately pointed. Frontal granular scales larger than occipital scales, with 36 small granular scales between the eyes. Dorsal scales granular from rostral to tail. Rostral undivided, in slight contact with nostril. Three small symmetrical internasals. Eight supralabials and seven infralabials. Prenasal not in contact with the 1st supralabial. Nostril circular, in narrow contact with the rostral and bordered by 1st supralabial, prenasal, one supranasal, and one postnasal. Four rows of scales between supralabials and the orbit. Mental large, triangular, and in contact posteriorly with two rounded and symmetrical postmental scales. Five smaller post-postmental scales. Gular scales granular and smaller than ventral scales. Ventrals imbricate, rounded, and larger than dorsal scales, in 14 scale rows across the venter. Body relatively robust and slightly elongated (TRL/SVL 0.41). Seven precloacal pores. Fore- and hind limbs moderately short and stout; forearm intermediate (FL/SVL 0.15); tibia short (CL/SVL 0.17). Digits elongated, unwebbed, with 5–6 terminal scansors. Thumb rudimentary with a small terminal claw. Relative length of digits: I <II = V <III <IV (manus); I <II <V <III <IV (pes).

Coloration: In life (holotype; Fig. 23A–B ), dorsal background colour light grey to brown, with a black reticulate dorsal pattern from snout to anterior insertion of the hind limbs; light cream blotches surrounded by black dots from eye to anterior insertion of the hind limb on each side of dorsum; tail brownish dorsally with light cream ocelli; throat white with two ∩ - shaped broken dark chevrons; venter uniformly white. In preservative (holotype; Fig. 22 ), dorsum uniformly dark grey from snout to tail. Venter with silvery appearance, becoming brownish on tail and fore- and hind limbs; black reticulated dorsal pattern visible, with two white paravertebral lines (one on each side) from eyes to midbody; digits dark brown ventrally.

Variation: In contrast to the holotype, the female paratype has a dorsal coloration in life that is entirely black with an orangish tail and dorsolateral dots in paravertebral lines that dissipate posteriorly, without a gular pattern ( Fig. 23C–D ). The lack of gular pattern is corroborated in preservative, in the paratype, and in another female specimen ( RMCA R.2947) from Basongo , DRC.

Habitat and distribution ( Figs 6 , 23 ): Lygodactylus leopardinus sp. nov. is known from Congo Basin primary rainforest at Balolombo, Katopa (Lomani River, Congo River affluent), and Basongo. This pattern suggests a large distribution in the Congo Basin like its sister taxon, L. depressus , which is found in sympatry at Lake Tumba and Katopa. However, it seems to be less abundant.

Natural history: A diurnal and arboreal rainforest species. The holotype was collected early in the morning on a tree about 1 m above the ground.

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 3. Comparative figure including dorsal (or dorsolateral) and ventral views of preserved holotypes (with exception of Lygodactylus gutturalis that is represented by a paratype) of all species within the L.gutturalis subgroup used in the current study.

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 5. Gular and skull osteology comparisons between holotype material or representative material of all members of the L. gutturalis subgroup and the L. angularis group mentioned in this work.From top to bottom, ventral view gular section; dorsal, lateral, and ventral view of the skull; lateral, dorsal, medial, and ventral view of jaw; and number of tooth loci. PM = premaxilla, MX(R) = right maxilla, MX(L) = left maxilla, DN(R) = right dentary of the jaw.

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 7. Holotype of L. heeneni [RMCA (= MRAC) R.8477] from Kapiri, Lualaba Province, DRC. A, Ventral and dorsal view of body. B, Details of head in dorsal, ventral, and lateral views (from top to bottom). Photos by J.L.R.

Figure 8. Paratype of L. paurospilus [RMCA (=MRAC) R.27409] from Haute Lubitshako, South Kivu Province, DRC. A, Ventral and dorsal views of body. B, Details of head in ventral, dorsal, and lateral views (from top to bottom). C, Details of cloacal region. D, Details of right manus. Photos by J.L.R.

Figure 22. Holotype of L. leopardinus sp. nov. (UTEP 22577) from Balolombo Village, Busira River, Équateur Province, DRC.A, Dorsal and ventral view of body. B, Details of right manus and (C) pes. D, Detailed view of head in ventral, dorsal, and lateral views (from top to bottom). Photos by J.L.R.

Figure 23. A–B, Holotype (UTEP 22577) and (C–D) paratype (UTEP 22596) of L. leopardinus sp. nov. in life and photo of habitat from (E) type locality at Balolombo Village, Busira River, Équateur Province, DRC. F–G, Holotype (UTEP 22587) and (H–I) paratype (UTEP 22584) of L. gamblei sp. nov. in life and photo of habitat from (J) type locality at Manono, Tanganyika Province, DRC. Photographs by E.G.