Hemidactylus cinganji, Lobón-Rovira & Conradie & Iglesias & Ernst & Veríssimo & Baptista & Pinto, 2021

Lobon-Rovira, Javier, Conradie, Werner, Iglesias, David Buckley, Ernst, Raffael, Verissimo, Luis, Baptista, Ninda & Pinto, Pedro Vaz, 2021, Between sand, rocks and branches: an integrative taxonomic revision of Angolan Hemidactylus Goldfuss, 1820, with description of four new species, Vertebrate Zoology 71, pp. 465-501 : 465

Hemidactylus cinganji sp. nov.

Figs 12 , 13

Hemidactylus benguellensis : Bocage (1893:115; 1895:12); Monard (1937:52)

Hemidactylus benguellensis : Marques et al. 2018; Ceríaco et al. 2020 [part]

The original description of H. benguellensis by Bocage make reference to “… species with 16 to 18 longitudinal series, 9 upper labials and 9 lower and 26 pre-cloacal and femoral pores, 13 on each side in a continuous series ". These characters were later corroborated by Monard (1937), with one male from Ebanga differing only from Bocage material on the number of preclocal-femoral pores (28 vs. 26).

In February 2020, a survey was conducted at Ebanga, Benguela Province, about 40 kilometers south of Cahata, the original type locality ( Bocage 1893), and from where some of the earliest material ascribed to this species had been collected ( Monard 1937). During this expedition an adult male morphologically assigned to H. benguellensis was collected on a granite rock boulder. A few days after, a second male was collected on a rock face overhanging a stream, near Passe, Benguela Province, ~80 km west from Ebanga and the type locality (Table S2, Fig. 12 ).

Interestingly, our phylogenetic analysis revealed a genetically separate lineage well-differentiated from the material assigned to H. benguellensis by Ceríaco et al. (2020a) (Fig. 2 ), with uncorrected p-distance 11.64% (Table 1 ). However, the available historical material collected between 1925 and 1953, from Ebanga, Hanha and Entre Rios ( Ceríaco et al. 2020a), was morphologically included as part of the taxon genetically assigned to H. benguellensis by Ceríaco et al. (2020a), without any molecular support. It should be noted that, not only it is unknown to which phylogenetic clade should the material from Hanha and Entre Rios be assigned to, but the problem is further compounded by some geographic inconsistencies. The specimens from Hanha, obtained during the VAE were most likely collected at Hanha Estate (also known as Hanha do Norte) and located on the semi-arid coast north of Lobito (see Branch et al. 2017 for a detailed discussion on misinterpretations surrounding the toponym Hanha), and not on the geographic location given by Ceríaco et al. (2020a), about 130 km to the southeast and on the base of the escarpment at 917 m a.s.l. On the other hand, the locality of Entre Rios (meaning "between rivers") can also be confusing, as several historical toponyms in Angola bear that name, and various publications have placed Entre Rios in different locations, ranging from 1,200 to 1,500 m a.s.l. in Benguela and Huambo provinces (e.g. Conradie et al. 2013; Marques et al. 2018; Ceríaco et al. 2020a). On his first Angolan expedition, Hellmich established his main base at the farm of Entre Rios, owned by a German citizen named Alfons Burger ( Hellmich 1957), which is located further south than previously acknowledged, at -13.30385° 14.70761°, 1076 m a.s.l, Benguela Province. Two specimens were here collected by Hellmich in 1953 and a third specimen was added by Herr Burger in the following year, all initially identified as H. mabouia , and only recently reassigned to H. benguellensis ( Ceríaco et al. 2020a). Nevertheless, the lack of phylogenetic context, the morphological variability recognized for specimens belonging to the neotypic clade, small sample size of the topotypic material, and the fact that Hanha and Entre Rios are intermediately located in relation to specimens genetically ascribed to different clades, recommends caution before assigning those specimens.

Based on field observations, the material assigned to H. benguellensis by Ceríaco et al. (2020a) is a very variable species present in different biomes and biogeographic units and occupying different niches, often showing arboreal habits, frequently found on the trunk or branches of trees, but sometimes in rocks, and also around human infrastructures such as man-made walls. In contrast, the new linage revealed here from the topotypic region, was only found associated with large granitic boulders in mountain habitats, even though it is a very limited sample.

Furthermore, the material recently assigned to H. benguellensis by Ceríaco et al. (2020a) presents some incongruence with our new topotypic material and the original H. benguellensis description. The historical material collected from Cahata and Ebanga ( Bocage 1893; Monard 1937), makes reference to large sized specimens (SVL 55 mm and 42 mm, respectively), with 9 supralabials and 26-28 precloacal-femoral pores. Remarkably, this description agrees fully with the new specimens collected in Ebanga and Passe while underscoring differences from the recently reassigned H. benguellensis , which consistently presents 10-11 supralabials (Table S6). Moreover, the neotype specimen that was ascribed to H. benguellensis by Ceríaco et al. (2020a) is an adult male with 40.8 SVL and 33 precloacal-femoral pores. Therefore, based on a morphological evidence and incongruence with the original description, clear well-differentiated genetic distance and morphological differences between the two clades were detected, adding to the large distance from the original type locality (>250 km). Notably, the new specimens found near the original topotypic agrees perfectly with the original H. benguellensis described by Bocage (1893) and later reported by Monrad (1937) from Ebanga. However, due to the recent redescription and new type locality assigned by Ceríaco et al. (2020a), and in order to maintain taxonomic stability and to avoid more confusion within this group, we consider the new material recovered from near the original type locality of H. benguellensis as a new species.

Holotype.

ANGOLA • 1 ♂; Benguela Prov., Ebanga; -12.77082°, 14.77102°; 1916 m a.s.l; 22 Feb. 2020; Pedro Vaz Pinto and Javier Lobón-Rovira; FKH0439.

Paratype.

ANGOLA • 1 ♂; Benguela Prov., Passe; -12.66774°, 14.01607°; 828 m a.s.l; 13 Feb. 2020; Pedro Vaz Pinto and Javier Lobón-Rovira; FKH0413.

Diagnosis.

A large sized Hemidactylus , with maximum SVL of 55 mm (mean) and maximum width of 10.5 mm (Fig. 13 ). Nine supralabials and 9-11 infralabials. Dorsal pholidosis with 13-18 rows of moderate strongly keeled tubercle scales and ventral pholidosis with 33-41 smooth and rounded scale rows on midbody. Hemidactylus cinganji sp. nov. present a moderate, triangular mental scale, two large postmentals followed by two medium post-postmentals. Tail with six strongly keeled dorsal tubercles rows dorsally and subcaudal scales medium sized, interspersed by 11 horizontal whorls of keeled scales on the original portion of the tail. Males with 26-28 continuous precloacal-femoral pores and 1-4 postcloacal tail spurs. Six divided scansors beneath first digit of both manus and pes, seven or eight beneath fourth digit of manus, eight or nine beneath the fourth digit of pes. Dorsum presents dark orange-beige coloration across the body with darker coloration on the dorsal part of the head that continues along the medial part of the dorsum until the sacrum.

Comparative diagnosis.

Hemidactylus cinganji sp. nov. can be distinguished from other non-Angolan western and central Africa congeners based on the same characteristics of H. benguellensis ( Ceríaco et al. 2020a). Hemidactylus cinganji sp. nov. can be distinguished from H. mabouia by the presence of smaller subcaudal scales (vs. large, elongated scales). It differs from the H. bayonii -group by its large size, largely keeled dorsal tubercle scales vs slightly keeled and larger number of precloacal-femoral pores (26-28 vs. 4-9 in H. bayonii , 4-6 in H. vernayi , 7-8 in H. nzingae and 8 in H. gramineus ). It differs from H. longicephalus -group by larger number of precloacal-femoral pores (26-28 vs. 6-11 in H. longicephalus and 6-8 in H. paivae ). It differs from H. benguellensis by lower number of supralabials (9 vs. 10-11 in H. benguellensis ).

Holotype description (Fig. 13 ).

Measurements and meristic characters of the holotype are presented in Table S6. Adult male with a snout-vent-length (SVL) of 45.05 mm and a tail length (TL) slightly smaller than SVL 42.07 mm. Body slender, nape distinct. Head slightly narrower than the body and moderate head length (HW/HL 0.72). Canthus rostralis not prominent. Eye diameter (3.05 mm), with vertical pupil and crenulated margin. Supraciliar scales small and rounded. Ear height (1.47 mm). Ear to eye distance slightly larger than orbit diameter (4.17 mm). Snout rounded and pointed. Frontal scales granular and larger than occipital scales. Occipital scales granular interspersed with large number of smooth and keeled tubercle scales from eyes to nape. Rostral undivided, in contact with 1st supralabial, nostril, supranasal and one internasal scales. 9 supralabial and 8-9 infralabials. First supralabial and rostral in direct contact with the nostril. Nostril circular rounded by rostral, 1st supralabial, supranasal, and two postnasals. One row of scales between supralabials and the orbit. Mental large, triangular and rounded posteriorly, with two larges rectangular postmental scales in broad contact posteriorly to the mental. 2 post-postmental scales, in contact with post-postmental slightly smaller than postmental scales and 1st and 2nd infralabials. Gular scales slightly smaller than ventral scales and granular.

Body relatively robust and slightly elongated (TRL/SVL 0.40). Ventral scales widely larger than dorsal scales, with 32 scales across the belly. The dorsal pholidosis present heterogenous conical, granular scales interspersed by 15 strongly keeled dorsal tubercle rows of at midbody. Dorsal tubercle rows are separated by 3 granular scales. Tubercle scales reach the posterior part of the eye region where tubercle scales remain keeled. Tail with six strongly keeled dorsal tubercles rows dorsally and subcaudal scales medium sized, interspersed by 11 horizontal whorls of keeled scales on the original portion of the tail. Regenerated portion of the tail presents homogeneous scales all around the surface without tubercle scales. 27 precloacal-femoral pores enlarged and 2 well-developed postcloacal spurs on each side.

Fore- and hindlimbs relatively short, stout; forearm short (FL/SVL 0.15); tibia short (CL/SVL 0.16). Short digits strongly clawed. All digits of manus and pes indistinctly webbed. Scansors beneath each toe composed by 1st and terminal scansor undivided with the exception of 4th and 5th toes with 3 and 2 undivided terminal scansor, respectively. Scansors beneath each finger equally divided, with exception of 1st and last terminal scansor undivided. Number of scansors: 5-5-6-7-6 (right manus), 6-8-8-9-7 (right pes). Relative length of digits: V < IV=III > II > I (right manus); V < IV < III > II > I (right pes).

Variation.

Variation in scalation and body measurements of the holotype and paratype of H. cinganji sp. nov. are reported in Table S6.

Coloration.

In life (holotype FKH0439; Fig. 12A ): this species displays a dark orange-beige coloration across the body with darker coloration on the dorsal part of the head that continues along the medial part of the dorsum until the sacrum; in the dorsal section it presents two lighter cream dorsolateral bands from the nares to the tail; the ventral part of the head shows a cream coloration with dispersed black speckles that increase in the anterior part of the infralabials; ventrum is uniform light beige or cream; supralabials are darker than infralabials; fore- and hindlimbs with irregular dark brownish coloration; tail with similar color and slightly banded; iris silvered with a black narrow pupil and brownish-golden reticulation. In preservative (Holotype; Fig. 13 ): dorsum with dark brownish coloration and two lighter dorsolateral bands from the narine till the sacrum; ventrum is light uniform beige. Variation: no variation was observed.

Etymology.

The name " Hemidactylus cinganji " is a widespread traditional word used in Angolan local languages that represents an ancestral spiritual entity that reincarnates assuming different physical forms in different places and occasions. This name is suitable as the new species corresponds to a taxon that was first described under a different name, then became lost and now resurfaces after its original name had been hijacked by a surrogate sister-species. The species epithet is used as a neuter singular noun in opposition to the generic name.

Distribution and conservation (Fig. 12B ).

The full distribution range of the species remains unknown, even though so far it has only been confirmed to occur in the interior of Benguela Province, from Cahata to Passe and Ebanga, above 800 m a.s.l. Due to poor sampling and few material confirmed to belong to this species, we cannot calculate the EOO and thus we regard the conservation status of the species as Data Deficient. Additional data will need to be collected to address its conservation status.

Natural history and habitat (Fig. 12C ).

Hemidactylus cinganji sp. nov exhibited a rock-dwelling behavior associated to large granitic boulders in mountainous regions from medium to high altitudes. The confirmed records fall in Benguela’s transition region between the Scarp and Transitional Zone and the Angolan Highlands. Both specimens were collected at night while foraging in large vertical granitic walls, one on a mountain cliff and the other on a boulder overhanging a forest stream. However, due to the poor sampling of this species, its natural history and real habitat requirements remain uncertain.

Figures

Figure 12. A - From top, dorsolateral view of Hemidactylus cinganji sp. nov. (FKH 0435), H. benguellensis (FKH 0413) and H. carivoensis sp. nov. (MNCN 50543); B - records of H. benguellensis - group (purple color represents records of H. cinganji sp. nov.; blue H. benguellensis; dark green H. carivoensis sp. nov.; circles represent genetically assigned records, while triangles represent historical or non-genetically assigned records by Ceriaco et al. 2020 a; color stars depict type or neotype localities; black star depicts original type locality of H. benguellensis assigned by Bocage (1893); C - habitat of H. cinganji sp. nov. at Ebanga; D - habitat of H. carivoensis sp. nov between Dombe-Echimina. E - habitat of H. benguellensis at Tchivinguiro. Photos Javier Lobon-Rovira (A-D) and William R. Branch (E).

Figure 13. Holotype of Hemidactylus cinganji sp. nov. (FKH 0439) from Ebanga, Benguela Province, Angola. A - Ventral and dorsal view of body. B - Detailed of head in lateral, dorsal and ventral views (from top to bottom). Photos by Pedro Vaz Pinto (A) and Rogerio Ferreira (B).

Figure 2 A. Bayesian Inference (BI) phylogenetic tree based on ND 2 + RAG 1 concatenated data, with Maximum Likelihood (ML) support overlaid. Numbers above the key nodes indicates BI posterior probability (≥ 0.90 were considered supported), while below the key nodes indicate ML bootstrap values (≥ 75 % were considered supported). Major clades are marked with a thick vertical line.