Cyrtodactylus kampingpoiensis, Quah & Grismer & Sinovas & Chourn & Chhin & Hun & Cobos & Geissler & Ching & Murdoch & Thi & Gregory & Nguyen & Hernandez & Kaatz & Grismer, 2025

Quah, Evan S. H., Grismer, L. Lee, Sinovas, Pablo, Chourn, Phyroum, Chhin, Sophea, Hun, Seiha, Cobos, Anthony, Geissler, Peter, Ching, Christian, Murdoch, Matthew L., Thi, Sothearen, Gregory, Jeren J., Nguyen, Eddie, Hernandez, Alexis P., Kaatz, Amanda & Grismer, Jesse L., 2025, Geographically structured genetic and morphological variation in a new species of Cyrtodactylus (Squamata, Gekkonidae) from a karstic archipelago in western Cambodia, ZooKeys 1240, pp. 73-115 : 73-115

Cyrtodactylus kampingpoiensis sp. nov.

Figs 5 , 6 , 7

Type material.

Holotype • Adult male ( LSUHC 15206 ) collected from Phnom Kamping Poi , Banan District, Battambang Province, Cambodia at 13°5.795'N, 102°55.798'E, at 114 m and nearby areas on 22 March 2024 by Pablo Sinovas, Seiha Hun, Phyroum Chourn, Matthew L. Murdoch, L. Lee Grismer, Evan S. H. Quah, Sothearen Thi, Christian Ching, and Anthony Cobos . Paratypes • Two adult males ( LSUHC 15203 and 15211) and five adult females ( LSUHC 15205 , 15207 , 15209–10 , and 15212) bear the same collection data as the holotype. The type series was collected from 1530–2200 hrs .

Additional specimens examined.

• Six hatchlings ( LSUHC 15196 –200, and 151202) bear the same collection data as the type series. The specimens were too small to recover reliable morphometric and meristic data but were included in the phylogenetic analyses.

Diagnosis.

Cyrtodactylus kampingpoiensis sp. nov. can be separated from all other species of the intermedius group by the combination of having a maximum SVL of 79.6 mm (female); 9–11 supralabials; nine or 10 infralabials; 30–37 paravertebral tubercles; 19–21 rows of longitudinally arranged tubercles; 38–46 longitudinal rows of ventrals; 5–7 expanded subdigital lamellae on the fourth toe; 11–13 unmodified subdigital lamellae on the fourth toe; 18–20 total subdigital lamellae on the fourth toe; 26–34 total number of enlarged femorals; no femoral pores; 5–9 enlarged precloacals; 7–9 precloacal pores in males (n = 3); three or four rows of large post-precloacal scales; 0–3 postcloacal tubercles; four dark body bands; 11–13 dark caudal bands (n = 7); 10–12 light caudal bands (n = 7); body tubercles not greatly reduced and moderately keeled; caudal tubercles extend beyond base of tail; subcaudals transversely expanded but not extending up onto side of tail; enlarged femorals and enlarged precloacals not continuous; enlarged proximal femorals equal (rarely subequal, one of seven) in size to distal femorals; no interdigital pockets; dorsal pattern not faded; no distinct dark markings on the top of head; lightened centers in dark body bands variable; no dark markings in light interspaces between body bands; dark body bands equal in width to light interspaces; light interspaces not reduced to a narrow thin white band; dark body bands bordered by prominent white tubercles; dark caudal bands slightly wider than light caudal bands; light caudal bands bearing dark markings in adults; posterior margin of nuchal loop not smoothly rounded (Table 4 ; Fig. 7 ; Suppl. materials 3, 4).

Description of holotype.

(Fig. 5 ; Table 4 ). Adult male SVL 78.9 mm; head moderate in length (HL / SVL 0.29), width (HW / HL 0.67), somewhat flattened (HD / HL 0.39), distinct from neck, triangular in dorsal profile; lores weakly concave anteriorly, weakly inflated posteriorly; prefrontal concave; canthus rostralis rounded; snout elongate (SN / HL 0.38), slightly convex, rounded in dorsal profile; eye large (ED / HL 0.23); ear opening elliptical, obliquely oriented, moderate in size; eye to ear distance slightly greater than diameter of eye; rostral rectangular, partially divided, bordered posteriorly by large left and right supranasals and a small internasal, bordered laterally by first supralabials; external nares bordered anteriorly by rostral, dorsally by large supranasal, posteriorly by two moderately sized postnasals, bordered ventrally by first supralabial; nine (R, L) rectangular supralabials tapering abruptly to below posterior margin of eye, first – sixth supralabials largest; nine (R, L) infralabials tapering smoothly to slightly past the termination of enlarged supralabials to corner of mouth; scales of rostrum and lores flat, much larger than granular scales on top of head and occiput; scales of occiput intermixed with small, rounded, tubercles; superciliaries elongate, largest dorsally; mental triangular, bordered laterally by first infralabials and posteriorly by large left and right trapezoidal postmentals contacting medially for ~ 40 % of their length posterior to mental; one row of enlarged, sublabials extending posteriorly to seventh infralabials (R, L); gular and throat scales small, granular, grading posteriorly into slightly larger, flatter, smooth, imbricate, pectoral and ventral scales.

Body relatively long (AG / SVL 0.45) with well-defined ventrolateral folds; dorsal scales small, granular, interspersed with moderately sized, keeled, semi-regularly arranged tubercles extending from occiput to beyond base of tail; ~ 20 longitudinal rows of tubercles at midbody; ~ 32 paravertebral tubercles; 43 flat, imbricate, ventral scales much larger than dorsal scales; seven enlarged pore-bearing precloacal scales, 4 R, 3 L separated by two poreless scales; no deep precloacal groove or depression; and three rows of large post-precloacal scales on midline.

Forelimbs moderate in length and stature (FL / SVL 0.15); granular scales of forelimbs larger than those on body, small rounded tubercles on dorsal surface of forearms; palmar scales rounded, juxtaposed; digits well-developed, inflected at basal interphalangeal joints, slightly narrower distal to inflections; subdigital lamellae transversely expanded, those proximal to joint inflections much wider than the nearly unmodified lamellae distal to inflections; claws well-developed, sheathed by a dorsal and ventral scale; hind limbs robust, wider and longer than forelimbs (TBL / SVL 0.19), covered dorsally by granular scales interspersed with moderately sized tubercles, larger and flatter scales anteriorly; ventral scales of thighs flat, imbricate, larger than dorsals; subtibial scales, flat, imbricate, slightly smaller than those of thigh; one row of 16 (R) 18 (L) slightly enlarged femoral scales terminating distally before knee, not continuous with enlarged precloacal scales, and poreless; proximal femorals nearly same size as distal femorals, all femorals forming an abrupt union with smaller, granular, posteroventral scales of thigh; plantar scales flat, juxtaposed; digits well-developed, inflected at basal interphalangeal joints; claws well-developed, sheathed by a dorsal and ventral scale at base; and five (R, L) wide subdigital lamellae on fourth toe proximal to joint inflection, 13 (R, L) narrower lamellae distal to joint inflection, 18 total subdigital lamellae.

Tail long (TL / SVL 1.04), partially regenerated, original portion 9.6 mm, regenerated portion 72.4 mm, 7.3 mm wide at base, tapering to a point; dorsal caudal scales of original portion of tail small, generally square, juxtaposed; median row of subcaudals significantly larger than dorsal caudals, transversely expanded, not extending dorsally onto lateral side of tail; body tubercles extending slightly beyond base of tail; hemipenal swellings at base of tail, three large postcloacal tubercles on both sides; and postcloacal scales flat, imbricate.

Color and pattern.

(Figs 5 – 7 ). Ground color of top of head, limbs, and dorsum straw to light-brown; top of head immaculate; prominent dark-brown nuchal loop edged in light-yellow bearing a somewhat flat posterior border extends between postorbital regions across the nape; no dark-brown band on nape; four immaculate dark-brown, slightly wavy-edged, dorsal body bands edged with bright-white tubercles have slightly lightened centers and are equal in width to straw-colored interspaces, extend from shoulders to groin, terminating ventrally slightly above the ventrolateral folds; first dorsal band extends anteriorly across shoulders; light-colored dorsal interspaces immaculate; forelimbs and hind limbs generally immaculate; one dark-brown caudal band on original portion of tail; regenerated portion of tail straw-colored overlain with faint, dark, irregularly shaped markings; iris gold with thin black reticulations; venter beige with faint, dark shadowing on lateral edges of belly, limbs, and throat; and subcaudal region essentially unicolor light-brown.

Etymology.

The species name kampingpoiensis is in reference to the type locality at Phnom Kamping Poi, Banan District, Battambang Province, Cambodia (Fig. 1 ).

Distribution.

The type series of Cyrtodactylus kampingpoiensis sp. nov. is known only from the type locality at Phnom Kamping Poi, Banan District, Battambang Province, Cambodia (Fig. 1 ).

Variation.

(Figs 6 , 7 ). The paratypes remarkedly approach the holotype in general coloration and pattern. The most notable variation pertains to the dorsal banding of the juvenile LSUHC 15203 whose third and fourth bands are broken on the midline and offset (Fig. 6 ). The overall pattern of LSUHC 15212 is far less bold than that of the holotype and the medial portion of its first band is posteriorly protracted. The juveniles have darker bands lacking lightened centers and are edged with yellow tubercles and there are no dark markings in the caudal interspaces. This is all accentuated in the hatchlings whose coloration is often slightly faded in the posterior ~ 25 % of the tail. The precloacal pore series in the holotype is separated medially by two poreless scales whereas the pore-bearing precloacal scales on the paratypes are continuous. Differences in meristics, morphometrics, and categorical characters are detailed in Table 4 and Suppl. material 3.

Geographic variation.

(Tables 3 , 5 ). Potential diagnostic differences in maximum SVL, meristics, and categorical characters between Cyrtodactylus kampingpoiensis sp. nov. and all other species in the intermedius group – except for species in the Battambang clade (used here in reference to all four populations) – are detailed in Suppl. material 4. Cyrtodactylus kampingpoiensis sp. nov. has an uncorrected pairwise sequence divergence from the intermedius group species ranging from 3.5–23.6 % (Suppl. material 2). Cyrtodactylus kampingpoiensis sp. nov. bears a 1.1–2.2 % uncorrected pairwise sequence divergence from the other three populations. In meristics, it differs further from the P. Banan population in having a statistically significant fewer number of enlarged precloacals (PS; 5–9 vs 9–12), unmodified subdigital lamellae (U 4 TL; 11–13 vs 13–16) and total number of subdigital lamellae (T 4 TL; 18–20 vs 20–23) (Table 5 ). It differs further in morphometrics in having a statistically significant longer ear-eye distances (EE), longer snout (SN), and a shorter foreleg (TBL). The PERMANOVA analyses indicate that Cyrtodactylus kampingpoiensis sp. nov. differs significantly in morphospace from the P. Banan population in the MFA (Figs 2 B , 4 C ) and the meristic PCA (Fig. 4 C ). Categorically, C. kampingpoiensis sp. nov. differs from the P. Banan population in that its slightly enlarged femorals and enlarged precloacals are usually discontinuous versus being continuous (four of five specimens; Suppl. materials 3, 4); the dark body bands are equal in width to the light interspaces as opposed to being wider; and hatchings and small juveniles have only a slightly faded tail tip as opposed to a nearly immaculate white tail tip (Fig. 6 D vs Fig. 9 E ).

In meristics, Cyrtodactylus kampingpoiensis sp. nov. differs significantly from the P. Sampeu population in having more rows of longitudinal tubercles ( LRT: 19–21 vs 15–17). It differs significantly from the P. Khpoh population and the P. Sampeu population in having a greater number of paravertebral tubercles ( PVT 30–37 vs 28–30 and 28–31, respectively) (Tables 3 – 5 ). It differs significantly morphometrically from the P. Sampeu population in having a significantly shorter ear-eye distance (EE) and a wider and longer head (HL and HW, respectively) and from the P. Sampeu and P. Khpoh populations in having a longer snout (SN) and longer hind limbs (TBL) (Table 3 ). Categorically it differs from the P. Sampeu population in having discontinuous versus continuous enlarged femorals and precloacals ( FS-PS) and there are no dark markings in the light interspaces between the dark body bands (BB-intr) whereas faint markings occur in the P. Khpoh population (Figs 7 , 9 , 10 ; Suppl. material 3).

Natural history.

All specimens of the type series were collected on P. Kamping Poi from 1030–2000 hrs. Phnom Kamping Poi is a long rectangularly shaped karstic hill ca 6.5 km in length and 1.6 wide and its widest point near its southeastern margin (Fig. 1 ). Phnom Kamping Poi reaches ca 242 m in elevation and is covered with drought-deciduous karst vegetation (Fig. 8 ). The hillsides are covered with karstic boulders of varying size which constitutes the prime microhabitat for C. kampingpoiensis sp. nov. although some specimens were found in a wide, open cave near the entrance. All age classes from hatchlings ( LSUHC 15198; 39 mm SVL) to adults ( LSUHC 15206, 15209, 15212; 82 mm SVL) were found and LSUHC 15207 ( SVL 77.9 mm) was gravid with two eggs. Although geckos were found on all substrates, they were most commonly found on karst in all planes of orientation with fewer specimens being found on the ground and the karst vegetation – often at the base of trees. One specimen ( LSUHC 15204 [in the FFI collection]) was found deep within a cave on top of a large (2.5 m diameter) log. Other species found on P. Kamping Poi were the lizards Eutropis longicaudata (Hallowell) , Dixonius siamensis (Boulenger) , Gekko gecko (Linnaeus) , Gehyra cf. lacerata (Taylor) , Gehyra mutilata (Wiegmann) Hemidactylus frenatus Duméril & Bibron , and Subdoloseps bowringii (Günther) , and the snakes Lycodon capucinus Boie and Indotyphlops cf. braminus (Daudin) .

All specimens from P. Banan (Figs 9 , 10 ) were collected near Rum Say Sok from 1930–2000 hrs. Phnom Banan is an irregularly shaped elongate karstic hill ca 6.6 km in length and 1.6 km wide at its widest point near its the eastern margin (Fig. 1 ). It reaches ca 203 m in elevation and is covered with drought-deciduous karst vegetation (Fig. 11 ). All age classes from hatchlings ( LSUHC 15177 [in FFI collection]; 35.0 mm SVL) to adults ( LSUHC 15168 [in FFI collection]; SVL 82.0 m) were found, indicating that mid-March is within the reproductive season for this population. Although geckos were occasionally found on the ground (most hatchings and juveniles), they were most commonly found on the karst surfaces in all planes of orientation with fewer specimens being found on karst vegetation (only at the base of trees less than 0.5 m above the ground or on fallen logs). Other species found on P. Banan were the frogs Polypedates cf. leucomystax (Gravenhorst) ; Glyphoglossus guttulatus (Blyth) ; Kaloula pulchra Gray ; Microhyla mukhlesuri Hasan, Islam, Kuramoto, Kurabayashi & Sumida ; the lizards Gehyra mutilata (Wiegmann) ; Subdoloseps bowringii (Günther) ; and Eutropis macularia (Blyth) ; and the snakes Oligodon cf. kampucheaensis Neang, Grismer & Daltry ; and a Chrysopelea ornata (Shaw) found dead on a nearby road.

All specimens were from P. Sampeu (Figs 12 , 13 ) from were collected from 1930–2050 hrs. Phnom Sampeu is a small oval karstic hill ca 1.8 km in length and 0.6 km across the center (Fig. 1 ). It reaches 102 m in elevation and is covered with drought-deciduous karst vegetation. The hillsides are covered with karstic boulders of varying size as well as shear vertical faces where specimens were most commonly found (Fig. 14 ). Geckos, however, were also found on the ground and on karst vegetation, usually at the base of trees. All age classes from hatchlings ( LSUHC 15126; 35 mm SVL) to adults ( LSUHC 151116; 80 mm SVL) were observed, indicating mid-March is within the reproductive season of the P. Sampeu population. Specimen LSUHC 15113 had eaten a large Huntsman Spider. Other species found on P. Sampeu were the frogs Duttaphrynus cf. melanostictus (Schneider) , Kaloula pulchra Gray , and Polypedates cf. leucomystax (Gravenhorst) , the lizards Dixonius siamensis (Boulenger) , and the snakes Lycodon capucinus Boie and Trimeresurus cf. albolabris Gray.

Phnom Khpoh is overall a somewhat circularly shaped karstic hill with deeply incised margins 8.6 km to the west of P. Sampeu. It is 5.0 km by 4.5 km in size with three major peaks – the western and eastern peaks reaching ca 250 m in elevation and a southern peak reaching ca 242 m in elevation (Fig. 1 ). Like the other hills, P. Khpoh is covered with drought-deciduous karst vegetation, varying karst boulder microhabitats, and caves of varying length and depth (Fig. 15 ). The three specimens collected were found on karst and vegetation from 1900–2030 hrs at the opening of a small cave. A fourth specimen was found low on a cave wall ~ 4 m from the opening (Figs 13 , 15 ). All age classes from hatchlings ( LSUHC 15232 [in the FFI collection; 36 mm SVL]) to adults ( LSUHC 15230; 80.0 mm SVL) were observed. LSUHC 15229 ( SVL 78.5 mm) is gravid. Other species found on P. Khpoh were the frogs Duttaphrynus cf. melanostictus (Schneider) , Kaloula pulchra Gray , and Polypedates cf. leucomystax (Gravenhorst) , the lizards Dixonius siamensis (Boulenger) and Hemiphyllodactylus khpoh Grismer, Sinovas, Quah, Thi, Chourn, Chhin, Hun, Cobos, Geissler, Ching & Murdoch , and the snakes Lycodon capucinus Boie and Trimeresurus cf. albolabris Gray.

Figures

Figure 1. Location of the karstic hills surveyed in Battambang Province and distribution of the new species Cyrtodactylus kampingpoiensis sp. nov., from Phnom Banan, and the other populations from Phnom Kamping Poi, Phnom Sampeu, Phnom Khpoh.

Figure 2. A Maximum Likelihood consensus tree of the Cyrtodactylus intermedius group B enlarged section of the Maximum Likelihood consensus tree highlighting the Battambang clade C MFA plots and percent character contribution bar graphs based on the nearly total evidence data set of the Battambang clade.

Figure 4. A, B PCA and DAPC plots, respectively, of the meristic data of the Battambang clade C summary statistics from the PERMANOVA analysis. Shaded cells denote populations whose morphospatial positions are statistically different.

Figure 5. Adult male holotype of Cyrtodactylus kampingpoiensis sp. nov. LSUHC 15206 A dorsal view B ventral view of gular region, throat, forelimbs, and feet C ventral view D ventral view of tail, pelvic region, hind limbs, and feet.

Figure 6. Cyrtodactylus kampingpoiensis sp. nov. A adult male holotype LSUHC 15206 B gravid adult female LSUHC 15207 C adult female LSUHC 15205 D juvenile LSUHC 15176 E juvenile male LSUHC 15203.

Figure 7. Type series of Cyrtodactylus kampingpoiensis sp. nov.

Figure 8. Habitat at Phnom Kamping Poi A, B optimal microhabitat of boulders of varying size within the karst vegetation C cave microhabitat inside Damrei Cave D typical deciduous karst vegetation.

Figure 9. The Phnom Banan population A adult male LSUHC 15173 B adult female LSUHC 15168 in FFI collection C adult female LSUHC 15170 D juvenile LSUHC 15176 E hatchling LSUHC 15175.

Figure 10. Specimens from the Phnom Banan population.

Figure 11. Habitat at Phnom Banan A hillside microhabtiat B Cliff face microhabitat C deciduous karst vegetation following summer rains. Photographs by Sothearen Thi.

Figure 12. A, C The Phnom Sampeu population A adult male LSUHC 15113 B juvenile LSUHC 15119 in FFI collection C hatchling LSUHC 15123 D – F the Phnom Khpoh population D gravid adult female LSUHC 15229 E juvenile female LSUHC 15231 F hatchling LSUHC 15233 in FFI collection.

Figure 13. Comparison of specimens from the Phnom Khpoh and Phnom Sampeu populations.

Figure 14. Habitat at Phnom Sampeu A vertical face microhabitat of a low karstic ridge B boulder microhabitat within open karst vegetation C vertical microhabitat D typical deciduous karst vegetation where LSUHC 15118 was found on the base of the tree in the foreground.

Figure 15. Habitat at Phnom Khpoh A typical boulder microhabitat within karst vegetation B open rocky microhabitat where specimens are rarely found C vertical opening into a cave microhabitat D cave microhabitat shared with Python bivittatus.