Two new cave-dwelling species of Bent-toed Geckos from Saraburi and Loei provinces, Thailand (Squamata: Gekkonidae: Cyrtodactylus) Author Sumontha, Montri 0000-0003-4829-7731 montri.sumontha@gmail.com Author Panitvong, Nonn 0000-0002-3979-0762 npanitvong@gmail.com Author Kunya, Kirati 0000-0002-7219-6239 Nakhonratchasima Zoo, 111 M. 1, Ratchasima-Pak Tongchai Rd., Chaimongkol, Muang, Nakhon Ratchasima 30000, Thailand. Author Donbundit, Nattasuda 0000-0002-3074-9288 Department of Biology, Faculty of Science, Khon Kaen University, Muang, Khon Kaen 40002, Thailand. Author Suthanthangjai, Winai 0009-0000-6617-5557 iansuthan@gmail.com Author Suthanthangjai, Maneerat 0009-0000-6617-5557 iansuthan@gmail.com Author Phanamphon, Eakarit Author Pauwels, Olivier S. G. 0000-0002-3265-5496 Royal Belgian Institute of Natural Sciences, Rue Vautier 29, B- 1000 Brussels, Belgium. & Corresponding author text Zootaxa 2024 2024-09-19 5512 2 272 294 http://dx.doi.org/10.11646/zootaxa.5512.2.9 journal article 10.11646/zootaxa.5512.2.9 e2ed3424-4a4a-40e0-82a2-eb4e80c81a9d 1175-5326 13848855 485907D7-0C23-4A44-8B77-FFE223E36408 Cyrtodactylus wiboonatthapoli sp. nov. ( Figures 7–10 ) Cyrtodactylus papilionoides –– Ellis & Pauwels 2012: 20 [ partim ] Cyrtodactylus papilionoides ––Chan-ard et al . 2015: 54 [ partim ] Cyrtodactylus papilionoides –– Cota et al . 2018 [ partim ] Cyrtodactylus papilionoides –– Jantarit & Ellis 2023: 174 Holotype . CUMZ-R-2603 (field no. MS 741 ), adult male caught on 11 July 2014 at the entrance of a cave on Phu Pha Sana ( 17°10’33.8”N , 101°57’52.5”E ), a limestone hill in Erawan District , Loei Province , northeastern Thailand , by K. Kunya and M. Sumontha. Paratype (1). CUMZ-R-2604 (field no. MS 743 ) , adult female; same locality and collectors as holotype . Diagnosis. Cyrtodactylus wiboonatthapoli sp. nov. can be distinguished from all other congeneric species by the combination of its maximal known SVL of 91.6 mm , 22 interorbital scales, 38 interciliary scales, 9 or 10 supralabials, 9 or 10 infralabials, 29–31 paravertebral tubercles of which 20–22 between limb insertions, 24 longitudinal rows of dorsal tubercles at midbody, 30–32 ventral scale rows at midbody, poorly defined, non-denticulate ventrolateral folds, a series of 19 enlarged femoroprecloacal scales including 2 pores on the proximalmost scales on each side separated by two small medial scales in male, no precloacal groove or depression, three postcloacal spurs on each side of the base of the tail in male, no interdigital webbing, 17–21 total subdigital lamellae on 4th toe, no enlarged median row of subcaudal plates, a dorsal pattern including an uninterrupted blackish brown nuchal collar and three blackish brown butterfly-shaped blotches on dorsum, 11 blackish brown rings on original tail, and a greysih golden iris. Description of the holotype . Adult male ( Figures 7 and 8 ). SVL 88.7 mm . TailL 107.5 mm , tail original and complete. Head relatively long (HeadL/SVL ratio 0.31), wide (HeadW/HeadL ratio 0.68), not markedly depressed (HeadD/HeadL ratio 0.39), distinct from slender neck. Loreal region inflated, canthus rostralis not prominent. Snout elongate (SnOrb/HeadL ratio 0.39), rounded, longer than orbit diameter (OrbD/SnOrb ratio 0.78); scales on snout small, rounded to oval, granular, mostly homogeneous, larger than those on crown, interorbital and occipital regions. Eye relatively large (OrbD/HeadL 0.30); pupil vertical with crenelated margins; supraciliaries short, anterior ones bearing small conical spines. Ear opening vertically elliptical, small (EarL/HeadL 0.02); orbit to ear distance subequal to orbit diameter (OrbEar/OrbD 0.95). Rostral wider (4.0 mm) than deep ( 2.8 mm ), rostral crease less than half of rostral height. Two enlarged supranasals separated from one another by a smaller scale. Rostral in contact with first supralabials, nostrils, supranasals and internasal. Nostrils oval, more or less laterally directed, each surrounded by supranasal, rostral, first supralabial and three postnasals. Three or four rows of small scales separate orbit from supralabials. Mental triangular, wider ( 3.1 mm ) than deep ( 2.6 mm ). A single pair of greatly enlarged postmentals in broad contact behind mental, each bordered anteromedially by mental, anterolaterally by first infralabial, posterolaterally by an enlarged lateral chinshield; both postmentals collectively bordered posteriorly by six gular scales. Supralabials to mid-orbital position 8/8, enlarged supralabials to angle of jaws 9/10. Infralabials 10/9. Interorbital scale rows across narrowest point of frontal bone 22. Body slender, relatively short (AG/SVL0.42; TrunkL 53.9 mm )with poorly defined, non-denticulate ventrolateral folds. Dorsal scales relatively homogeneous, domed; dorsal tubercles about five times size of adjacent dorsal scales, extending from nape onto tail, smaller tubercles on postocular region, crown, occiput and nape; most tubercles bearing a strong keel, less marked on lower flank tubercles; tubercles on posterior trunk and sacral region most prominent; tubercles in 24 rows at midbody, separated from one another by three or four dorsal granules. Nine to 11 granular dorsal scales surround each tubercle. Ventral scales larger than dorsals, smooth, oval and subimbricate, largest on posterior abdomen and in precloacal region. Midbody scale rows across belly between ventrolateral folds 30. Gular region with homogeneous, smooth, juxtaposed granular scales. No precloacal groove or depression. On each side a continuous series of 19 FemPreEnSc, the two proximalmost bearing each a pore; the two series separated by a diastema of two small poreless medial scales. Postcloacal spurs each bearing three enlarged, conical scales. Scales on palm and sole rounded to oval or hexagonal, domed. Scalation on dorsal surface of hind and forelimbs similar to body dorsum with enlarged tubercles interspersed among smaller scales. Fore and hind limbs long, slender (FaL/SVL 0.19, TibL/SVL 0.20). Digits long, slender, inflected at interphalangeal joints, all bearing robust, slightly recurved claws. Basal subdigital lamellae broad, oval to rectangular, without scansorial surfaces (4-5-5-5-4 right manus, 3-5-6-6-6 right pes); narrow lamellae distal to digital inflection and not including ventral claw sheath: 6-10- 11-9-10 (right manus), 6-10-10-14-14 (right pes); no interdigital webbing. Relative lengths of digits: III>IV>V>II>I (manus), IV>V>III>II>I (pes). Tail original and complete, gently tapering to pointed tip, distinctly longer than SVL (TailL/SVL ratio 1.21). No median row of enlarged subcaudals. FIGURE 7. Live male holotype of Cyrtodactylus wiboonatthapoli sp. nov . in situ at the type locality. A. General dorsal view. B. Ventral view. C. Right profile. Photos by M. Sumontha. In life, dorsal ground color of head dark brown; dorsum and dorsal surfaces of limbs light brown. Dark reticulations on head. A blackish brown continuous nuchal loop, pursuing a brown preorbital stripe, connects orbits and extends posteriorly to the level of the shoulders. Three blackish brown butterfly-shaped blotches on dorsum, distinctly longer than their light interspace, the anterior one beginning above the shoulder; numerous brown and blackish brown irregular marks on the light interspaces ( Figures 7 and 8 ). Irregular brown bars on upper surface of limbs, hands and feet. Background color of tail light brown as dorsum, becoming slightly lighter posteriorly, with 11 blackish brown rings, encircling the tail; irregular brown marks on the light interspaces between the dark rings. Iris greyish golden. Supralabials and infralabials white, barred with blackish brown. Throat and venter light grey; undersides of fore- and hind limbs dark grey. Variation. Main morphometric and meristic characters of the paratype are provided in Table 1 . Eleven or 12 granular dorsal scales surround each tubercle in the paratype . The female paratype also has a complete, original tail, with a TailL/SVL ratio of 1.13, thus proportionally a bit shorter than in the male. An adult and a subadult from the type-locality ( Figure 9 ), not collected, show a dorsal pattern identical to the one shown by the types. Distribution and natural history. The types and several other individuals were all encountered near the entrance of the cave on Phu Pha Sana. Within the cave, Cyrtodactylus wiboonatthapoli sp. nov. was found in syntopy with Cyrtodactylus cf. interdigitalis ( Figure 12 ; possibly C. ngati , which was confirmed from Phu Luang by Grismer et al. 2022b ), Gehyra cf. fehlmanni (Taylor) and Gekko gecko (Linnaeus) . Just outside the cave it was encountered in direct proximity to the squamates Calotes versicolor (Daudin) ( Agamidae ), Dixonius cf. siamensis (Boulenger) , Hemidactylus frenatus and H. platyurus ( Gekkonidae ), Lycodon capucinus Boie and L. davisonii (Blanford) ( Colubridae ), Rhabdophis siamensis (Mell) ( Natricidae ), Trimeresurus macrops Kramer ( Viperidae ) and the anurans Fejervarya limnocharis (Gravenhorst) ( Ranidae ) and Polypedates megacephalus Hallowell ( Rhacophoridae ). We also encountered Cyrtodactylus wiboonatthapoli sp. nov. in Erawan Cave ( 17°20’52.8”N , 102°01’05.9”E , altitude about 445 m above sea level; Figure 10 ), on the border between Na Wang District in Nong Bua Lamphu Province and Erawan District of Loei Province , at about 20 km N-NE of the type-locality. Ellis & Pauwels (2012) presented photographs of Cyrtodactylus wiboonatthapoli sp. nov. (under the name C. papilionoides ) taken in situ in Tham Yai Wat Pha Sawan ( 17°07’05.9”N , 101°56’01.3”E , altitude 420 m above sea level), Ban Pha Sawan, Pha Khao District, Loei Province . This latter cave is located about seven km SW of the type-locality. FIGURE 8. Preserved type series of Cyrtodactylus wiboonatthapoli sp. nov. From above: adult male holotype CUMZ-R-2603 and adult female paratype CUMZ-R-2604. Photo. by M. Sumontha. Being found both inside and near limestone caves, Cyrtodactylus wiboonatthapoli sp. nov. is a trogloxene species ( sensu Jantarit & Ellis 2023 ) with a karst ecotype ( sensu Grismer et al . 2020 , 2021). Etymology. The specific epithet honors the late Suthep Wiboonatthapol (1953–2023), former science teacher at Khamsaenwittayasan School in Nong Bua Lamphu Province . We suggest the following common names: ตุ๊กกายเอราวั ( Took-kai Erawan ; Thai), Erawan Bent-toed Gecko (English) , and Cyrtodactyle d’Erawan (French). Comparison to other species. Its lack of widened subcaudal plates distinguishes Cyrtodactylus wiboonatthapoli sp. nov. from C. amphipetraeus (which also shows, among others differences, four to seven dark bands on dorsum), C. auribalteatus (whose adults show regular round dark spots on the light interspaces on dorsum), C. bansocensis Luu, Nguyen, Le, Bonkowski & Ziegler (regular dorsal bands with straight transversal borders), C. calamei Luu, Bonkowski, Nguyen, Le, Schneider, Ngo & Ziegler (four dark bands between limb insertions), C. chanhomeae (regular dorsal bands with straight transversal borders), C. chauquangensis Hoang, Orlov, Ananjeva, Johns, Hoang & Dau (discontinuous nuchal collar), C. darevskii Nazarov, Poyarkov, Orlov, Nguyen, Milto, Martynov, Konstantinov & Chulisov (four or five thin dark bands between limb insertions), C. doisuthep Kunya, Panmongkol, Pauwels, Sumontha, Meewasana, Bunkhwamdi & Dangsri (six or seven thin dark bands between limb insertions), C. dumnuii Bauer, Kunya, Sumontha, Niyomwan, Pauwels, Chanhome & Kunya (four of five dark bands between limb insertions), C. hinnamnoensis Luu, Bonkowski, Nguyen, Le, Schneider, Ngo & Ziegler (four to six dark bands between limb insertions), C. houaphanensis Schneider, Luu, Sitthivong, Teynié, Le, Nguyen & Ziegler (five irregular dark bands between limb insertions, discontinuous nuchal collar), C. interdigitalis (short nuchal collar), C. intermedius (four regular dark bands between limb insertions), C. inthanon Kunya,Sumontha,Panitvong,Dongkumfu, Sirisamphan & Pauwels (three to five bands between limb insertions), C. jaegeri Luu, Calame, Bonkowski, Nguyen & Ziegler (or four regular dorsal bands with straight transversal borders between limb insertions), C. jarujini (dorsal pattern made of numerous irregular blotches), C. khammouanensis Nazarov, Poyarkov, Orlov, Nguyen, Milto, Martynov, Konstantinov & Chulisov (three or four regular dorsal bands with straight transversal borders between limb insertions), C. khelangensis Pauwels, Sumontha, Panitvong & Varaguttanonda (four dark bands between nuchal loop and hind limbs insertion), C. kunyai (four or five regular dark bands between nuchal collar and hind limbs insertion), C. lomyenensis Ngo & Pauwels (three regular dorsal bands with straight transversal borders between limb insertions), C. meesookae sp. nov. (banded pattern), C. muangfuangensis (four dark bands between limb insertions); C. multiporus Nazarov, Poyarkov, Orlov, Nguyen, Milto, Martynov, Konstantinov & Chulisov (dorsal pattern made of numerous blotches), C. pageli Schneider, Nguyen, Schmitz, Kingsada, Auer & Ziegler (five dark bands between limb insertions), C. phukhaensis Chomdej, Pradit, Pawangkhanant, Kuensaen, Phupanbai, Naiduangchan, Piboon, Nganvongpanit, Yuan, Zhang, Che, Sucharitakul & Suwannapoom (four to six dark bands between limb insertions), C. rufford Luu, Calame, Nguyen, Le, Bonkowski & Ziegler (three or four regular dorsal bands with straight transversal borders between limb insertions), C. saiyok (three to five irregular, thin dark bands between limb insertions), C. soudthichaki Luu, Calame, Nguyen, Bonkowski & Ziegler (five dark bands between limb insertions), C. spelaeus Nazarov, Poyarkov, Orlov, Nguyen, Milto, Martynov, Konstantinov & Chulisov (four or five pairs of dark blotches between limb insertions), C. teyniei David, Nguyen, Schneider & Ziegler (dorsal pattern made of numerous irregular blotches), C. thathomensis Nazarov, Pauwels, Konstantinov, Chulisov, Orlov & Poyarkov (four dark bands between limb insertions), C. tigroides (regular dorsal bands with straight transversal borders) and C. uthaiensis (regular banded pattern). Its four PrePo in a discontinuous series and its lack of FemPo in males separate it from C. amphipetraeus (seven to nine PrePo and 10–12 FemPo in males), C. auribalteatus (six PrePo and eight to ten FemPo in males), C. bansocensis (34 FemPrePo in males), C. calamei (35–39 FemPrePo in males), C. chanhomeae (32–34 FemPrePo in males), C. chauquangensis (six or seven PrePo in males), C. darevskii (38–44 FemPrePo in males), C. denticulatus (13 PrePo and 20 FemPo in males), C. doisuthep (six or seven FemPi and six PrePo in males), C. dumnuii (five or six PrePo and 12 FemPo in males), C. hinnamnoensis (36–44 FemPrePo in males), C. houaphanensis (six PrePo in males), C. interdigitalis (14 PrePo and 17 FemPo in males), C. intermedius (eight to ten PrePo in males), C. inthanon (five PrePo and 8–12 FemPo in males), C. jaegeri (44 FemPrePo in males), C. jarujini (52–54 FemPrePo in males), C. khammouanensis (40–44 FemPrePo in males), C. khelangensis (two to six PrePo and six or seven FemPi or FemPo in males), C. kunyai (3 PrePo and 10–12 FemPo in males), C. lomyenensis (39–40 FemPrePo in males), C. meesookae sp. nov. (6 PrePo in males, no enlarged femorals), C. monilatus (no pores in males; dorsal pattern made of numerous roundish blotches), C. muangfuangensis (six PrePo and 15 FemPo in males), C. multiporus (58–60 FemPrePo in males), C. ngati (13 PrePo and 14 FemPo in males; five dark bands between limb insertions), C. phukhaensis (seven PrePo and 10–12 FemPo in males), C. rufford (42 or 43 FemPrePo in males), C. saiyok (five PrePo in males), C. soudthichaki (29 FemPrePo in males), C. spelaeus (eight or nine PrePo in males), C. teyniei (14 PrePo in males), C. thathomensis (10 PrePo and 36–38 FemPo in males), C. thongphaphumensis (15 PrePo and 12–16 FemPo in males), C. tigroides (eight or nine PrePo and 10–14 FemPo in males), C. uthaiensis (14 PrePo and 12 FemPo in males) and C. wayakonei Nguyen, Kingsada, Rösler, Auer & Ziegler (six to eight PrePo in both sexes). Cyrtodactylus wiboonatthapoli sp. nov. is separated from C. vilaphongi Schneider, Nguyen, Le, Nophaseud, Bonkowski & Ziegler , whose males are still unknown, by its blotched ( versus banded) pattern, its higher DTR (24 versus 15 or 16) and its enlarged femorals ( versus not enlarged). The only species within 400 km radius which Cyrtodactylus wiboonatthapoli sp. nov. resembles are C. angularis and C. papilionoides . It differs from the former by its first dorsal blotch beginning above the shoulder ( versus behind the shoulder), its blackish brown ( versus brown) dorsal blotches distinctly longer than the lighter interspaces ( versus smaller blotches whose length is subequal to the lighter interspaces), and its four ( versus five or six) PrePo. It is distinguished from C. papilionoides by its continuous ( versus interrupted or at least near-interrupted) nuchal collar, much higher DTR number (24 versus 12–14) DTR, and four PrePo in a discontinuous series ( versus four to six in a continuous series). The differences in color and pattern appear clearly when comparing Figures 7–10 with photographs of C. angularis and C. papilionoides in Ulber & Grossmann (1991) and Chan-ard et al . (1999) and Figures 13 and 14 . It should be noted that these two species were long confused because Taylor (1963) illustrated Cyrtodactylus angularis with a photograph showing two C. papilionoides (see Ulber & Grossmann 1991 ). The type locality of Cyrtodactylus wiboonatthapoli sp. nov. lies about 250 km N-NE of those of C. angularis and C. papilionoides . While Cyrtodactylus wiboonatthapoli sp. nov. is a cave and limestone dweller, C. papilionoides is a ground dweller, often found in plantations ( Ulber & Grossmann 1991 ; Chan-ard et al . 1999). FIGURE 9. Live adult (A; tail tip regenerated) and subadult (B; tail original and complete) individuals of Cyrtodactylus wiboonatthapoli sp. nov. at the type-locality. Individuals not collected. Photos by M. Sumontha. Although the ground-dwelling Cyrtodactylus papilionoides was described from Pak Chong District in Nakhon Ratchasima Province , Chan-ard et al . (2015) mentioned that it only occurred in the more northern provinces of Chaiyaphum , Loei and Phetchabun , without any mention of Nakhon Ratchasima Province , and they described its habitat as limestone caves in dry evergreen forest. The type-locality of Cyrtodactylus wiboonatthapoli sp. nov. in Loei Province is relatively close to the borders with Chaiyaphum and Phetchabun Provinces; it is hence possible that at least the Loei record is actually attributable to the new cave-dwelling species described here. The drawing that Chan-ard et al. (2015) provided however represents a typical Cyrtodactylus papilionoides and is probably based on a published photograph of this species.