Lygodactylus gamblei, 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 : 483-485

LYGODACTYLUS GAMBLEI SP.NOV.

( Figs 23–24 , Table 3 ; Supporting Information, Table S10)

Zoobank registration: https://zoobank.org/ 411D833D-2017- 4930-BD11-B783AAD6A9B7

Lygodactylus picturatus gutturalis : de Witte (1953) [part].

Lygodactylus gutturalis : Pasteur (1964) [part]; Broadley and Cotterill (2004) [part]; Röll (2005) [part].

Lygodactylus gamblei sp. nov. is a miombo-savannah form within subgroup C, described above.It also differs by a minimum of c. 9.96% for the 16S mitochondrial gene (Table 2) from its sister taxon L. leopardinus sp. nov. ( Fig. 2A ) and lacks nuclear haplotype sharing in RAG1 ( Fig. 2B–C ).

Holotype: UTEP 22587 ( ELI 340 ), male with regenerated tail and ventral incision, collected at Manono , Tanganyika Province, DRC, S07.29363, E27.39472, 634 m a.s.l. on 19 January 2010 by locals and brought to Eli Greenbaum.

Paratypes (nine specimens): UTEP 22584 ( ELI 293 ), female with original tail and ventral incision, collected at Mulongo, Haut-Lomami Province , DRC, S07.65509, E27.34027, 875 m a.s.l. on 18 January 2010 by Chifundera Kusamba, Wandege M. Muninga, Mwenebatu M. Aristote, and Eli Greenbaum ; RBINS 2721 (formerly under RBINS 6157 ), adult male, collected at Rivière Kande, affluent de la rive gauche de la Lupiala et sousaffluent de la rive droite de la Lufira, Upemba National Park, Haut-Katanga Province , DRC, 700–730 m a.s.l., on 4–8 October 1947 ; RBINS 2722 (formerly under RBINS 6160 ), adult male collected at Rivière Lukawe, affluent de la rive droite de la Lufira, Upemba NP, Haut-Katanga Province , DRC, 700 m a.s.l. on 28 October 1947 ; RBINS 2723 and RBINS 6124 (formerly under RBINS 6124 ), male and female, respectively, collected at Munoi, bifurcation de la rivière Lupiala, affluent de la rive droite de la Lufira, Upemba NP, Haut-Katanga Province , DRC, 890 m a.s.l. on 3 June 1948 ; RBINS 2725–27 (formerly in a series of nine specimens under RBINS 6122 ), two males and one female, respectively, collected at Munoi, bifurcation de la rivière Lupiala, affluent de la rive droite de la Lufira, Upemba NP, Haut-Katanga Province , DRC, 890 m a.s.l. on 28–31 May 1947 ; RBINS 2728 (formerly under RBINS 6156 ), collected at Kaswabilenga, région du cours inférieur de la Lupiala, affluent de la rive droite de la Lufira, Upemba NP, Haut-Katanga Province , DRC, 700 m a.s.l. on 31 January 1949 . All paratype material from RBINS was collected by G.-F. de Witte during his expeditions in the former Katanga region between 1946 and 1949 .

Additional material: RMCA R.8933, male, collected at Kapanga , Haut-Katanga Province, DRC, in September 1932 by G.F. Overlaet ; RBINS 6121 and RBINS 20749 (formerly in RBINS 6121 series) , male and female, respectively, collected at Kambi , affluent de la Grande Kafwe ( Masombwe ), Upemba NP, Haut-Katanga Province, DRC, on 25–27 July 1945 by G.-F. de Witte ; RBINS 6139 (male) and RBINS 20748 (female) (formerly in RBINS 6139 series) , collected at Mabwe , Upemba NP, Haut-Lomami Province, DRC, in August 1945 by G.-F. de Witte .

Diagnosis: A large Lygodactylus [ SVL to 39.8 mm (mean 36.1 ± 2.1 mm)] with 6–9 supralabials and 6–7 infralabials. As with other Lygodactylus within the L. gutturalis group, this species can be easily differentiated from the L. angularis group species by the gular ornamentation, dorsal pattern, and lack of terminal scansors on the tail tip, and from other members of the L. picturatus group based on dorsal coloration and gular pattern (see L. gutturalis account).

Lygodactylus gamblei sp. nov. can be differentiated from other members within the L. gutturalis subgroup by the following combination of characters: dorsal coloration lacking dorsolateral series of cream ocelli on flanks (present in L. gutturalis , L. karamoja sp. nov., L. kibera sp. nov., and L. dysmicus ), with two parallel dorsolateral lines from head to forelimbs, the lower line extending on to the anterior side of the forelimb, reaching the cubital fossa ( Fig. 25B ); males usually with three ∩ - shaped chevrons (vs. one or two in L. depressus ; a broken chevron in L. leopardinus sp. nov.; and the exceptional inverted Y -shaped pattern in L. mirabundus sp. nov.) and thicker gular lines than L. gutturalis , L. dysmicus , and L. karamoja sp. nov.. Lygodactylus gamblei sp. nov. always has three postmental scales, with the lateral pair separated by a posterior extension of the mental ( Fig. 25A ). It can be differentiated from L. karamoja sp. nov., L. kibera sp. nov., and L. leopardinus sp. nov. by having a proportionally larger tibia (CL/ SVL ≥ 0.19 vs. CL/ SVL ≤ 0.17). As in its sister taxon L. leopardinus sp. nov., females lack gular patterning (present in all the other members of the group, Fig. 25A ). Lygodactylus gamblei sp. nov. can be differentiated from L. mirabundus sp. nov. by having a lower number of precloacal pores (7–9 vs. 10) and by subtle morphometric differences ( Table 3 ).

Etymology: We name this new species after the American evolutionary biologist and herpetologist Tony Gamble of Marquette University, in recognition of his substantial contributions to the evolutionary biology of geckos. The name is a patronym formed in the genitive case.

Description of the holotype ( Fig. 24 ): Measurements and meristic characters of the holotype are presented in Supporting Information, Table S10. Adult male, with a snout–vent length ( SVL) of 35.0 mm and a regenerated tail of 31.0 mm.Body slender and nape slightly distinct. Head slightly broader than body, and moderate head length ( HW /HL 0.66). Canthus rostralis not prominent. Eye diameter 2.51 mm with circular pupil. Ear to eye distance slightly larger than the orbit diameter (3.26 mm). Snout rounded and moderately pointed. Granular scale of frontal larger than occipital scales, with 25 small interorbital granular scales. Dorsal scales granular from rostral to tail. Rostral undivided, in narrow contact with nostril. Two small symmetrical internasals. Seven supralabials and six infralabials. Prenasal contacts the 1st supralabial. Nostril circular and surrounded by 1st supralabial, prenasal, one supranasal, and one postnasal. Four or five rows of scales between supralabials and the orbit. Mental large, triangular, and posteriorly in contact with three rounded symmetrical postmental scales. Mental scales extending between the lateral postmentals reaching ~50% of the medial surface suture of the postmentals. Five smaller post-postmental scales. Gular scales granular, rounded, and smaller than ventral scales. Ventrals imbricate, denticulate, and larger than dorsal scales, with 19 scale rows across the venter. Regenerated tail with 30 enlarged subcaudal scales, some fragmented. Body relatively robust and slightly elongated ( TRL / SVL 0.42). Eight precloacal pores. Fore- and hind limbs moderately short and stout; forearm of medium length ( FL / SVL 0.16); tibia moderately long (CL/ SVL 0.19). Digits elongated and 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 ( Fig. 23A–D, F–I ), dorsal colour light brown to grey, with dark brown mottled pattern in the female; male with three rows of interrupted, but conspicuous black stripes on lateral surface from head to neck, converging at the anterior insertion of the forearm, giving the impression of a single large blotch; male and female with a black-brown line from snout to ear aperture; venter uniform orange-yellowish from posterior part of gular to cloaca, and cream to white elsewhere; gular immaculate white in female, and white in male with three thick and black ∩ - shaped chevrons. In preservative (holotype; Fig. 24 ): similar coloration as in life, slightly darker, with gular pattern more brownish.

Variation: All paratypes largely agree with the holotype, and their measurements are summarized in Supporting Information, Table S 10. The specimen RBINS 6138 has a bold dorsal pattern from head to midbody. It is worth noting that all the female specimens examined from de Witte’s (1953) expedition lack a gular pattern, in contradiction to de Witte’s (1953) statement. However, we noted that some young male specimens were catalogued as females, which could have misled de Witte to this conclusion.

Habitat and distribution ( Figs 6 , 23 ): Lygodactylus gamblei sp. nov. is known from Manono, Upemba National Park, and other nearby areas in south-eastern DRC. The species is associated with Miombo Woodland forest and gallery forest between 500 and ~ 1500 m a.s.l. This species is sympatric with L. heeneni in this area.

Natural history: A diurnal and arboreal species. The holotype (UTEP 22587) was collected active during the daytime with an air temperature of 45 °C and one of the paratypes (UTEP 22584) was found scampering around a tree close to the ground in an open area.

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 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 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.

Figure 24. Holotype of L. gamblei sp. nov. (UTEP 22587) from Manono, Tanganyika Province, DRC.A, Dorsal and ventral view of body.B, Details of the cloacal region.C, Detailed view of head in ventral, dorsal, and lateral views (from top to bottom). Photos by J.L.R.

Figure 25. A, Dorsal and ventral views of head, showing gular ornamentation and pholidosis variation in L. gamblei sp. nov. B, Specimens of L. gamblei sp. nov. in lateral view, showing its characteristic scapular pattern. Red points denote males and blue points denote females. Photos by J.L.R.