Research Article
Research Article
Common but ignored: a new species of Cyrtodactylus (Chordata, Reptilia, Squamata, Gekkonidae) from lowland Sumatra Barat, Indonesia
expand article infoFitra Arya Dwi Nugraha, Yuni Ahda, Djong Hon Tjong§, Nia Kurniawan|, Awal Riyanto, Muhammad Alif Fauzi|, Si-Min Lin#
‡ Universitas Negeri Padang, Padang, Indonesia
§ Universitas Andalas, Padang, Indonesia
| Universitas Brawijaya, Kota Malang, Indonesia
¶ Herpetology Group Research, Biosystematic and Evolution Research Center - National Research and Innovation Agency (BRIN)-("Museum Zoologicum Bogoriense"), Jakarta, Indonesia
# National Taiwan Normal Universit, Taipei, Taiwan
Open Access


The lowland region of Sumatra Barat has received little attention in previous biodiversity studies. Past studies have mainly focused on highland habitat and conservation areas. However, many populations of Cyrtodactylus in the lowland habitats of Sumatra Barat were not correctly identified. A phylogenetic tree based on the NADH dehydrogenase subunit 2 (ND2) gene showed that the lowland Sumatran population is the sister group of the Malaysian lowland species, C. semenanjungensis, together nesting within the agamensis group. The genetic divergence within the Sumatra Barat population is 0–4.2% and 18.3–20% to C. semenanjungensis. Further examination of morphological characters revealed that they differed from the sister clade and other Sumatran Cyrtodactylus members by a unique combination of characters such as absence of tubercle on brachium, presence of tubercle on ventrolateral fold, 32–41 paravertebral tubercles, 38–46 ventral scales, enlarged femoral scales, presence of precloacofemoral pores and 22–23 subdigital lamellae under fourth toe. Based on the morphological and molecular evidence, the lowland Sumatran population is herein described as a new species, increasing the number of species in Sumatra to seven. More comprehensive and intensive sampling efforts would most likely yield further discoveries in the group of Sumatran Cyrtodactylus in the near future.

Key words

Bent-toed gecko, diversity, morphology, ND2 gene, phylogeny, Sumatra, taxonomy


Species diversity and new discoveries in the Bent-toed gecko genus Cyrtodactylus Gray, 1827 is remarkable in recent years. In only two decades from 2000, the number of new species descriptions has dramatically increased this taxon from only 77 species to nearly 300. This steep upward trajectory has indeed indicated that the true diversity of Cyrtodactylus is highly underestimated. Among Southeast Asian countries, Indonesia is ranked fifth as the largest contributor (considering type localities) to the number of Bent-toed gecko discoveries, behind Myanmar, Vietnam, Malaysia and Thailand (Grismer et al. 2021). Despite having a larger area than Peninsula Malaysia, the number of species discovered in Sumatra island was significantly lower (6 vs 26), presumably suggesting a low sampling or research effort undertaken on the island. But, perhaps the most influencing factor contributing to its diversity is the evolution of habitat preferences where some habitat transitions have occurred. The changes in habitat preference may in part be responsible for its diversity (Grismer et al. 2020).

Currently, there are six species of Cyrtodactylus distributed across the mainland of Sumatra (Teynie et al. 2010; Uetz et al. 2022). Four species were described based on Sumatran type series: C. agamensis (Bleeker, 1860); C. psarops Harvey, O’Connell, Barraza, Riyanto, Kurniawan & Smith, 2015; C. semicinctus Harvey, O’Connell, Barraza, Riyanto, Kurniawan & Smith, 2015; and C. lateralis (Werner, 1896). Meanwhile, C. consobrinus (Peters, 1871) dan C. quadrivirgatus Taylor, 1962 were described based on non-Sumatran type series.

Sampling efforts for herpetofauna specifically targeting the Sumatra Barat region have not been comprehensively performed until the last decade. Teynie et al. (2010) carried out sampling in Sumatra Barat Province and Jambi Province but the majority was in the surroundings of Lake Maninjau. Although they found a significant number of reptiles and amphibian and reported a new species of Bufonidae (Amphibia: Anura), no Cyrtodactylus were collected. Milto and Bezman-Moseyko (2021) also surveyed the surroundings of Lake Maninjau, reporting two new records Limnonectes deinodon Dehling, 2014, Dicroglossidae and Sphenomorphus scotophilus (Boulenger, 1900) and one new species Cnemaspis calderana Milto & Bezman-Moseyko, 2021 for Sumatra and provided extended descriptions of Cyrtodactylus agamensis. Nugraha et al. (2022) surveyed lowland habitat generally close to human settlement such as paddy fields, peat swamp near the coastline, streams and other plantations. They reported Cyrtodactylus sp. from a rocky stream and did not provide a description. Unfortunately, there was no sampling effort that focused on collecting lowland Cyrtodactylus from Sumatra.

Several potentially new species from Sumatra have been reported by O’Connell et al. (2019). At least four potentially new species among the C. psarops group await formal description. We excluded Cyrtodactylus marmoratus Gray, 1831 from Sumatra as this species only occurs in Java (O’Connell et al. 2019) and morphometric assessment of specimens labelled as C. marmoratus showed that they did not belong to this species (Fauzi et al. 2022).

Between 2020 and 2022, we surveyed lowland forest in four locations of Sumatra Barat targetting Cyrtodactylus. We found several individuals of Cyrtodactylus and examined those specimens using both morphological and molecular approaches. Based on our examination on those individuals, we found they differed from other Cyrtodactylus in Sumatra and they should be treated as a new species.

Materials and methods

Field survey and specimen preservation

A total of 16 individuals from lowland of Sumatra Barat Province were collected by hand during fieldwork in 2020–2022 (Fig. 1). All Individuals were collected during the night from 19.00 to around 24.00. The locality of samples were recorded using a Garmin GPS 65 s. The collected specimens were euthanized by immersion in 250 mg/L benzocaine solution. The fresh tissues (liver) were preserved in 96% ethanol and stored at -20 °C for genetic analysis, and the specimens were fixed using 10% formalin and then transferred to 70% alcohol for long term preservation. Photographs were taken before and immediately after euthanization using a mirrorless camera Sony Alpha 6000 with Laowa 65 mm f/2.8 macro lens. In order to document the original coloration, we defered documentation until the next day if the individuals showed stress coloration. All photos were deposited at the Department of Biology, Universitas Negeri Padang (UNP) along with the preserved specimens.

Figure 1. 

Localities of samples used in this study. The star indicates the type locality. The black-filled circle indicates the locations of the paratypes.

Morphological analysis

Color characters were assessed from digital images of living individuals prior to preservation. The sex was determined by confirming the presence of the hemiphenal structure by injecting formaline into the postcloacal region. The measurements were taken with a dial caliper to the nearest 0.05 mm under a Nikon SMZ1270 stereo microscope. We followed Grismer and Leong (2005) for examination of morphological and meristic characters including:

SVL Snout-vent length, measured from tip of snout to vent;

AX Axial length, measured from posterior margin of forelimb insertion to anterior margin of hind limb insertion;

HL Head length, measured from tip of snout to articulation of quadrate bone;

HW Head width, measured at widest part of head;

HH Head height, measured from occiput to underside of lover jaw;

SL Snout length, measured from tip of snout to anterior margin of orbit;

OEL Orbit-ear length, measured from posterior margin of eye to anterior margin of ear opening;

OD Orbit diameter, measured from anterior to posterior margin of orbit;

EL Ear length, measured from anterior to posterior margin of ear opening;

ML Mental length, maximum length of mental shield;

EN Eye-nostril distance, measured from anterior margin eye to posterior margin of external nares;

IN Internarial distance, measured between the nares across the rostrum;

FL Forearm length, measured from base of palm to elbow;

TBL Tibia length, measured from base of heel to knee;

TaL Tail length, measured from the vent to the tip of the tail, original or regenerated;

TaW Tail width, measured at widest part of tail.

Merisitic charatcers were counted on both right and left sides when possible. The characters were recorded as follows:

DTR Dorsal tubercles, number of tubercle rows on dorsum at midbody, counted in one row between lateral folds;

PVT Paravertebral tubercles, number of tubercles counted in a longitudinal row between posterior insertion of fore limb and anterior insertion of hind limb;

VS Ventral scales, number of ventral scales at midbody, counted in one row between lateral folds;

EPFS Enlarged precloaca and femoral scales, number of enlarged precloacofemoral scales, counted along lowest, pore-bearing row;

PFP Precloaca and femoral pores, number of precloaca and femoral pores;

PCT Postcloacal tubercles, number of postcloaca tubercles, right and left;

LT4 Subdigital lamellae under 4th toe, subdigital scales under 4th toe, counted from first enlarged scale (true lamellae) on lower side of toe to scale proximal to apical scale;

SL Supralabial scales, labial scales of upper jaw, beginning with first enlarged scale bordering rostral shield, ending with last enlarged scale bordering labial angle for right and left side;

IL Infralabial scales, labial scales of lower jaw, beginning with first scale bordering mental shield, ending with last enlarged scale bordering labial angle for right and left side;

IN Internasal scales, number of scales between rostronasals, bordering rostral shield;

GUL Gular scales, number of gular scales bordering pair of 1st postmentals (excluding enlarged second 2nd postmentals)

To make clear the counting of some scales (supra and infralabials, precloacafemoral scales, gular scales) and detection of the presence of pores, we used a staining technique with methylene blue in 70% alcohol (Harvey et al. 2015). For species comparison, we used the published descriptions of other species.

Laboratory protocols

Total genomic DNA was extracted from liver tissues using the Qiagen Dneasy tissue kit (Valencia, CA, USA) following the standard protocol for animal tissues. The Natrium Dehydrogense Subunit 2 (ND2) gene and partial flanking tRNAs was amplified by using polymerase chain reactions (PCRs) under the following conditions: a cycle of 9 min at 94 °C, then followed by 35 cycles of 45 s at 94 °C, 45 s at 60 °C and 1 min at 72 °C with a final extension step of 6 min at 72 °C. Amplifications were carried out in 25-µl volume consisting of 2.5 µl genomic DNA (approximately 100 ng), 0.4 µм each primer and 1× GoTaq Green Master Mix (Promega, Wisconsin, USA). The primers used in this study followed Oliver et al. (2012): M112F (5’- AAGCTTTCGGGGCCCATACC-3’) and M1123R (5’- GCTTAATTAAAGTGTYTGAGTTGC -3’). The PCR products were visualized using 1.5% agarose gel for confirmation of PCR amplification. The PCR product was then sent to the sequencing service 1st BASE Singapore ( through PT. Genetika Science Indonesia. The PCR primers were also used for sequencing.

Phylogenetic reconstruction

Sequences were uploaded, assembled and editted in Geneious software 2022.2.2 ( The protein coding-region of the sequences were aligned and translated to amino acid to verify that the targeted sequences were assembled correctly. After that confirmation, all sequences were submitted to GenBank. We followed Grismer et al. (2021) for the classification of species groups. For comparison, ND2 sequences from the C. darmandvillei, C. marmoratus, C. lateralis, C. sworderi and C. agamensis groups that were available in GenBank were used for phylogenetic tree reconstruction with our samples (Table 1). Phylogenetic relationship was constructed using maximum likelihood (ML) analysis performed using RaxML HPC Black Box with 1000 bootstrap replicates implemented in CIPRES Sience Gateway portal (Miller et al. 2010; accessed through We used the GTR+Γ model of sequence evolution. Nodal support with bootstrap (BS) values ≥ 70 was considered as strongly supported (Hillis and Bull 1993). The tree resulted from RAxML was then visualized and editted in iTOL v6 (Letunic and Bork 2021; available at and further improvement in Photoshop C6 64-bit. We also calculated uncorrected p-distances using MEGA 7 (Kumar et al. 2016).

Table 1.

Species used in the phylogenetic reconstruction including localities and GenBank accession numbers of the mitochondrial NADH dehydrogenase subunit 2 gene. PM = Peninsular Malaysia; Gn.= Gunung.

Species Locality Museum number Accession number Source
agamensis group
Cyrtodactylus gonjong sp. nov. Agam, Sumatra Barat, Indonesia UNP193 OR208777 This study
Cyrtodactylus gonjong sp. nov. Agam, Sumatra Barat, Indonesia UNP194 OR208778 This study
Cyrtodactylus gonjong sp. nov. Agam, Sumatra Barat, Indonesia UNP199 OR208779 This study
Cyrtodactylus gonjong sp. nov. Agam, Sumatra Barat, Indonesia UNP203 OR208780 This study
Cyrtodactylus gonjong sp. nov. Agam, Sumatra Barat, Indonesia UNP146 OR208781 This study
Cyrtodactylus gonjong sp. nov. Tanah Datar, Sumatra Barat, Indonesia UNP061 OR208782 This study
Cyrtodactylus gonjong sp. nov. Padang, Sumatra Barat, Indonesia UNP053 OR208783 This study
Cyrtodactylus gonjong sp. nov. Padang, Sumatra Barat, Indonesia UNP055 OR208784 This study
Cyrtodactylus gonjong sp. nov. Tanah Datar, Sumatra Barat, Indonesia UNP062 OR208785 This study
Cyrtodactylus gonjong sp. nov. Padang Pariaman, Sumatra Barat, Indonesia UNP045 OR208786 This study
Cyrtodactylus gonjong sp. nov. Padang Pariaman, Sumatra Barat, Indonesia UNP047 OR208787 This study
Cyrtodactylus gonjong sp. nov. Padang Pariaman, Sumatra Barat, Indonesia UNP048 OR208788 This study
Cyrtodactylus gonjong sp. nov. Padang, Sumatra Barat, Indonesia UNP053 OR208789 This study
Cyrtodactylus gonjong sp. nov. Tanah Datar, Sumatra Barat, Indonesia UNP060 OR208790 This study
Cyrtodactylus gonjong sp. nov. Padang Pariaman, Sumatra Barat, Indonesia UNP165 OR208791 This study
Cyrtodactylus gonjong sp. nov. Padang Pariaman, Sumatra Barat, Indonesia UNP167 OR208792 This study
C. metropolis Batu caves, Selangor, PM LSUHC 11343 KU253579 Grismer et al. 2016
C. payacola Bukit Panchor, Penang, PM LSUHC 10070 JQ889190 Johnson et al. 2012
C. majulah Nee Soon Swamp, Singapore ZRC 26951 JX988529 Grismer et al. 2013
C. pantiensis Gn. Panti, Johor, PM LSUHC 8905 JQ889186 Johnson et al. 2012
C. tiomanensis Pahang, PM LSUHC 6251 JX440563 Wood et al. 2012
C. rosichonariefi Bunguran, Great Natuna, Indonesia MZB Lace 12132 KP256187 Riyanto et al. 2015
C. psarops Indonesia MZB 9687 MH248931 O’Connell et al. 2019
C. sp. 3 Indonesia ENS 18140 MH248911 O’Connell et al. 2019
C. sp. 4 Indonesia ENS 18591 MH248912 O’Connell et al. 2019
C. sp. 5 Indonesia ENS 18659 MH248916 O’Connell et al. 2019
C. sp. 6 Indonesia ENS 18719 MH248917 O’Connell et al. 2019
C. semenanjungensis Gn. Panti, Johor, PM LSUHC 8900 JQ889177 Johnson et al. 2012
C. semicinctus Indonesia ENS 14749 MH248925 O’Connell et al. 2019
C. cf. agamensis Indonesia ENS 19636 MH248910 O’Connell et al. 2019
C. cf. agamensis Indonesia ENS 19635 MH248909 O’Connell et al. 2019
C. cf. agamensis Indonesia ENS 19634 MH248908 O’Connell et al. 2019
C. cf. agamensis Indonesia ENS 19694 MH248907 O’Connell et al. 2019
sworderi group
C. quadrivirgatus Bukit Larut, Perak, PM LSUHC 8859 JQ889241 Johnson et al. 2012
C. guakanthanensis Gua Kanthan, Perak, PM LSUHC 11323 KU253577 Grismer et al. 2016
C. tebuensis Gn. Tebu, Terengganu, PM LSUHC 10902 JX988527 Wood et al. 2012
C. sworderi Sungai Kawal, Peta, PM LSUHC 7685 JQ889189 Johnson et al. 2012
C. gunungsenyumensis Hutan Lipur Gn. Senyum, Pahang, PM LSUHC 12201 KU253585 Grismer et al. 2016
lateralis group
C. lateralis Indonesia UTA 62916 KU893163 Harvey et al. 2016
C. rubidus CES 131445 KM255203 Agarwal et al. 2014
C. durio Malaysia LSUHC 9725 KU893159 Harvey et al. 2016
marmoratus group
C. marmoratus Indonesia ENS 15932 KR921721 Harvey et al. 2015
C. papuensis SAMA R62652 JQ820320 Oliver et al. 2012
C. sp. 1 Indonesia ENS 15813 KR921697 Harvey et al. 2015
C. sp. 2 Indonesia ENS 15784 KR921689 Harvey et al. 2015
darmandvillei group
C. batucolus Pulau Besar, Melaka, PM LSUHC 8934 JQ889179 Johnson et al. 2012
C. petani Pasuruan, Jawa Timur, Indonesia MZB Lace 11706 KU232620 Grismer et al. 2016
C. kimberleyensis Siuna, Sulawesi Tengah, Pulau Sulawesi, Indonesia WAM R164144 JX440544 Wood et al. 2012
C. jellesmae Siuna, Sulawesi Tengah, Pulau Sulawesi, Indonesia RMB 1672 GU550721 Siler et al. 2010
C. sadleiri Christmas island, Australia SAMA R34810 JQ820309 Oliver et al. 2012
C. seribuatensis Pulau Nangka Kecil, Johor, PM LSUHC 6349 JQ889187 Johnson et al. 2012
C. darmandvillei Nusa Tenggara Barat, Indonesia WAM R98393 JX440533 Wood et al. 2012
Hemidactylus frenatus LLG 4871 GQ458049 Murthy et al. 2015
Gekko gecko Thailand: Patong Beach, Kathu District, Phuket Island, Phuket Province MVZ 215314 AF114249 Macey et al. 1999


Phylogenetic relationships of Cyrtodactylus from Sumatra Barat

After assembling sequences, we obtained 609 to 1027 basepairs of ND2 gene for subsequent alignment and phylogenetic tree construction. In general, the ML tree topology was congruent with a previous study (Grismer et al. 2021) in term of inter-relationship among major groups; i.e., C. agamensis, C. sworderi, C. lateralis, C. marmoratus and C. darmandvillei. We obtained a new putative species that was strongly supported as a sister clade to Cyrtodactylus semenanjungensis Grismer & Leong, 2005 from Peninsular Malaysia (BS = 100). Together they nested within the C. agamensis group (Grismer et al. 2021) along with C. semicinctus, C. psarops and other non-Sumatran species (e.g., Cyrtodactylus payacola Johnson, Quah Anuar, Muin, Wood, Grismer, Greer, Onn, Ahmad, Bauer & Grismer, 2012 and Cyrtodactylus majulah Grismer, Wood & Lim, 2012) (Fig. 2). The genetic distance within the new putative species group was 0–4.2% (mean = 1.7%) and the distance between them to C. semenanjungensis was 18.3–20% (mean = 19.3%). The p-distance of new putative species to other species is more than 17% (Table 2). We herein describe this population as new to science. Although the bootstrap value separating the new putative species from C. semenanjungensis is significant, the bootstrap value between the new putative species and other agamensis group species (C. semenanjungensis, C. semicinctus, C. cf. agamensis) is less than 70.

Table 2.

Uncorrected pairwise genetic distance (%) within the new putative species and with the closely related lineages based on the ND2 gene.

No. Species 1 2 3 4
1. C. sp. 0–4.2
2. C. semenanjungensis 18.3–20.0
3. C. semicinctus 18.7–19.7 18.6
4. C. cf. agamensis 24.6–26.0 27.6 17.6
5. C. psarops 31.9–38.4 31.9 27.3 30.5
Figure 2. 

The maximum likelihood (ML) tree topology of the new species with other Cyrtodactylus inferred by the ND2 gene sequences. The light orange box indicates the C. agamensis group members (the sample indicated by an asterisk denotes the holotype). The numbers on the branches are bootstrap values.


Cyrtodactylus gonjong sp. nov.

Recommended English common name: Gonjong Bent-toed Gecko Recommended Indonesia common name: Cicak Jari Lengkung Gonjong

Type material

Holotype. UNP193 (Fig. 3) adult female, collected from Langkuik Tamiang waterfall (0°26'0.7404"S, 100°16'44.5404"E), Nagari (Village) Malalak Selatan, Kecamatan (District) Malalak, Kabupaten (Regency) Agam, Sumatra Barat Province, Indonesia, on 22 July 2022 by Fitra Arya Dwi Nugraha (FADN), Yunico Amardi (YA) and Mahesa Rafi. Paratypes (Fig. 4). One adult male (UNP045) collected from Anai Dempoland tourism object (0°29'46.8312"S, 100°20'7.638"E), village Kayu Tanam, district 2×11 Kayu Tanam, regency Padang Pariaman, Sumatra Barat Province on 3 June 2020 by FADN and YA. Two adult males (UNP052, UNP053) collected from village Sungai Barameh (1°1'54.71"S, 100°24'43.69"E), district Padang Selatan, Padang city, Sumatra Barat Province on 31 January 2021 by FADN and YA. One adult male (UNP060) collected from Lembah Anai Nature Reserve (0°29'24"S, 100°20'24"E), district X Koto, regency Tanah Datar, Sumatra Barat Province on 10 June 2021 by Mahesa Rafi, Katon Agusdi, Fadhil Raid and Fachrul Rozi Octavian. One adult male (UNP148) collected from Sarasah Uwak Waterfall (0°54'28"S, 100°28'54"E), village Limau Manis, district Pauh, Padang city, Sumatra Barat Province on 31 May 2022 by FADN. Two adult females (UNP165, UNP167) collected from village Sungai Sirah (0°24'8.8128"S, 100°8'37.6728"E), district Sungai Geringging, regency Padang Pariaman, Sumatra Barat Province on 7 June 2022 by FADN and YA. One juvenile (UNP194), same data as holotype. Three adult males (UNP196, UNP199, UNP203) collected from village Koto Malintang (0°17'56.3748"S, 100°7'42.5064"E), district Tanjung Raya, regency Agam, Sumatra Barat Province.

Figure 3. 

The holotype (UNP193, adult female, SVL 65.1 mm) A rostral view B gular view C precloacal scales D dorsal E ventrolateral F ventral view. A–D, F were taken after 1 month preservation E is a live individual.

Figure 4. 

The representative paratypes of C. gonjong sp. nov.


The specific ephitet gonjong is taken from the name of the roof style of the typical house in Sumatra Barat created by its ethnic people, called Minang. The house itself, called Rumah Gadang, and the unique style of the roof shape was inspired by the horn of buffalo, the most respected animal in Minang ethnology. The gonjong has become a symbol used by the Minang people to show their ethnic identity outside Sumatra Barat and now is used not only for the house but also for government buildings, restaurant buildings, hotels or other public venues. This name is given as an honour to the Minang people because they were helpful during our survey.


Cyrtodactylus gonjong sp. nov. is assigned to the C. agamensis group on the basis of its recovered phylogenetic position (Fig. 2). This species can be differentiated from all congeners within the C. agamensis group by having the following combination of characters: (1) a medium-sized Cyrtodactylus, 54.1–77.7 mm in adult males, 65.1–76.7 mm in adult females; (2) 10–13 enlarged supralabial and 9–13 enlarged infralabial scales; (3) 3–5 internasal scales; (4) antebrachium tuberculated, brachium not tuberculated; (5) 32–41 paravertebral tubercles; (6) 16–19 longitudinal rows of dorsum tubercles; (7) 38–46 ventral scales; (8) 22–24 subdigital lamellae on the fourth toe; (9) 34–46 enlarged femoroprecloacal scales; (10) femoroprecloacal pores present in both sexes, 13–36 in adult males, 0–18 in adult females; (11) precloacal groove or depression present; (12) enlarged median subcaudals absent; (13) distinct ventrolateral folds; (14) subconical prominent tubercles on body that extend to the base of the tail; (15) 2–3 postcloacal tubercles; (16) two postocular stripes fused to a U-shaped mark on occiput; (17) postocular stripes extended beyond the arms; (18) 7–8 dark bands on trunk; (19) tail ringed by beige and black; and (20) labials yellow or dark with yellow/beige spots.

Description of holotype

Adult female with 65.1 mm SVL; head moderately in length (HL/SVL 0.29), wide (HW/HL 0.60), slightly flattened (HH/HL 0.34), distinct from neck, and triangular shape from dorsal view; lores concave; canthus rostralis rounded; snout elongated (SL/HL 0.40), rounded in rostral region, eye to snout distance larger than head depth; eyes large (OD/ HL 0.22), obtrusive and appeared beyond labials in dorsal view, eye diameter less than the eye to ear distance, pupil vertical; ear oppening small (EL/HL 0.03), elliptical, oriented obliquely leaning posteriorly; rostral large, subrectangular in shape, medial posterior edge interupted by an subhexagonal internasal scale that embedded within rostral, posteriorly bordered by three internasal scales, laterodorsally by nostril opening and lateroventrally by first supralabial; external nares directed lateroposteriorly, bordered anteriorly by rostral, posteriorly by two postnasals: one subcircular, one crescent shaped, dorsally by large supranasal, and ventrally by first supralabial; supranasal subrectangular, separated by one subhexagonal scale and a smaller subrectangular scale that piled up, the smaller one above the subhexagonal one, supranasal laterally bordered by nostril, right supranasal anteriorly bordered by five significantly smaller scales, left supranasal anteriorly bordered by four significantly smaller scales; one internasal scale, subhexgonal in shape, embedded within rostral, bordered by two supranasal scales, one smaller scale right posteriorly, and rostral anteriorly; 9/10 (right/left) supralabial scales to below center of the eye, 10/11 (right/left) to the posteriormost enlarged scale, 8 infralabial scales to below center of the eye, 10/11 (right/left) to the posteriormost enlarged scale; scales of frontonasal, prefrontal and lores small, juxtaposed, relatively raise; weak tubercles on the supraorbital region; prominent tubercles above the ear opening, larger than those on supraorbital, the tubercles gradually increased in size posteriorly.

Body slender, relatively short (AX/SVL 0.47), with distinct ventrolateral fold; scales on dorsum small, homogenous, interspersed by rounded to trihedral tubercles; tubercles present from the occiput region to the base of the tail but no further than 1/3 of the tail, being more dense gradually posteriorly until the base of the tail; 37 paravertebral tubercles; 17 tubercles transversally in the middle of the trunk; imbricate and smooth ventral scales, scales in middle larger than those on lateral and dorsal, 42 scales across the center of the trunk from one ventrolateral fold to another; femoral scales enlarged, extending until 2/3 portion of femora, contiguous with enlarged precloacal scales, forming a reverse V-shaped mark in the middle of hindlimb; a greatly enlarged scale at the apex, larger than other precloacal and femoral scale, rather long than wide, each right and left bordered by three smaller scales; 46 continuous enlarged precloacal and femoral scales, all similar in size except the scale at apex; precloacal groove or depression absent.

Limbs moderately slender; forelimbs relatively short (FL/SVL 0.15); scales on forelimbs dorsum larger than on body dorsum, rostrum or frontal, domed to subconical in shape, scales near the junction of limbs with trunk being smaller; round to subconical tubercles present on antebrachium, concentrated in the middle of antebrachium, similar in size with those on nape; brachium not tuberculated; fingers relatively long, well developed, no webbing; claw well developed, relatively short; hind limbs more robust than fore limbs, moderate in length (TBL/SVL 0.19); scales on dorsum domed to subconical, interspersed with trihedral tubercles; tubercles on dorsum approximately similar in size with those on posterior trunk; anterior ventral scales small, gradually increase in size posteriorly, the scales after enlarged femoral scales much smaller than those on anterior part; 22 subdigital lamellae on fourth finger; toes relatively long, claw well developed, relatively short.

Tail 78.9 mm in length, longer than SVL (TL/SVL 1.21), 5.4 mm in width at base, cylindrical, decreasing in size posteriorly; scales on tail dorsum small, approximately similar in size with those on the distal of femoral; tail dorsum tuberculated at its base, no more than 1/3 of tail; postcloacal tubercles at each side, left tubercles slightly searated, right tubercles in contact one another; no enlarged median subcaudal scale; subcaudal scales larger than on tail dorsum.

Coloration in life

Dorsal ground color of head, neck, trunk, limbs and tail beige to weak yellow; yellow mottling along the ventrolateral fold, on the lateral part of neck, on supra and infralabial scales; ventral of head, neck, trunk, limbs whitish to grey pale; palmar, metatarsal, fingers and toes darker; basal subcaudal ground color whitish to pale grey with dense yellow mottling; most of the original tail except basal subcaudal encircled by beige and black; labials yellow or dark with beige or yellow spots; weak black stripe between nostril and eye (sometimes absent); lateral stronger black stripe extends from posterior margin of the eye to approximately second black band on dorsum; wide U-shaped band around occiput connects with lateral stripe; the U-shaped band usually continuous (only one exception in UNP165 which is discontinuous in the middle); the Y-shaped pattern on occiput present in two individuals (UNP045, UNP193), the others lack Y-shaped pattern instead of irregular network or spots; bands on trunk dorsum sometimes clearly lying transversally and sometimes blotched; strong nape black spot only present in one individual (UNP045) where left portion fused with lateral stripe, fainted nape black spot presents in two individuals (UNP148, UNP165), the others lack nape spot; black lateral spots present positioned parallel with dorsum bands or between them; 5–6 irregular black bands on forelimbs and hindlimbs; beige to yellow spots on the base fingers (except finger 1); dorsum first finger lighter than other fingers; the clear beige and black bands on subcaudal appear after the one third portion from basal.


Cyrtodactylus gonjong sp. nov. can be differentiated from all other congeners in the C. agamensis group based on a combination of morphological characters.

Cyrtodactylus gonjong sp. nov. differs from its most close relative, C. semenanjungensis from Peninsular Malaysia by having a larger body size, SVL maximum 77.7 in adult male and 76.7 in adult female (vs 62.1 mm in adult male and 69 mm in adult female); 8–10 supralabial scales to center of eye (vs 11–15); 16–19 DTR (vs 18–20); maximum PVT reachs 41 (vs 37); 38–46 ventral scales (vs 49–53); enlarged femoral scales present (vs absent); femoral and precloacal pores present in both sexes (vs absent in both sexes); 22–24 subdigital lamellae under fourth toe (vs 20–21); a black spot on the nape absent (vs present); posterior end of lateral stripe far beyond the arm insertion to the body, approximately reaching the second band on dorsum (vs between the arms insertion); posterior end of lateral stripe in contact with dorsum band that is separated (vs lateral stripe in contact with clear, not separated band or non-blotched band, creating a box encircled the nape spot); mostly lack nape spot, if present the spot is faint (vs strong black nape spot present).

Cyrtodactylus gonjong sp. nov. differs from C. semicinctus by being smaller in the adult female (76.7 mm vs 89 mm) and larger in the adult male (77.7 mm vs 75 mm); having fewer DTR (16–19 vs 29–35); having more PVT (32–41 vs 24–27); maximum ventral scales 46 (vs 44); 0–36 precloacal and femoral pores (vs 36–38); 22–24 subdigital lamellae under the fourth toe (vs 19–22);10–13 enlarged supralabial scales (vs 8–11); 10–13 enlarged infralabial scales (vs 8–10); lateral stripe extending from behind the eye to behind arm present (vs absent); labials with beige or yellow spot (vs without spot); beige and black band encircled tail and subcaudal (ringed tail) (vs only dorsum tail – not ringed tail).

Cyrtodactylus gonjong sp. nov. differs from C. psarops by being smaller in the adult female (76.7 mm vs 82 mm) and larger in the adult male (77.7 mm vs 74 mm); 16–19 dorsal tubercles transversally (vs 28–38); 32–41 longitudinal row tubercles on the middle of the body (vs 23–26); maximum number of ventral scales 46 (vs 49); 0–36 pores on precloacal and femoral region (vs 28–32); 2–3 postcloacal tubercles (vs usually single); 10–13 enlarged supralabials (vs 9–12); 10–13 enlarged infralabial scales (vs 8–11); brachium not tuberculated (vs tuberculated); postocular stripes, left and right, fused to form U-shaped mark on the occiput (vs usually does not fuse to form U-shaped mark); labials usually pale grey or yellow (vs charcoal or dark).

Cyrtodactylus gonjong sp. nov. differs from C. agamensis by being being smaller in the adult female (76.7 mm vs 86.8 mm) and larger in the adult male (77.7 mm vs 74.9 mm); 10–13 infralabial scales (vs 9–12); maximum number of dorsal tubercles transversally (21 vs 19); maximum number of paravertebral tubercles 41 (vs 37); 38–46 ventral scales (vs 50–67); 13–36 precloacal and femoral pores in males (vs 9–10), 0–18 in females (vs 0–7); 7–8 body black bands on trunk (vs 6–7); postocular stripe extends beyond the arms present (vs absent).


Dorsal ground color of head, neck, trunk, limbs and tail beige to weak yellow. Labials yellow (UNP148, UNP165, UNP167) or dark on the first three supralabials but pale grey for the rest (UNP193, UNP194, UNP199, UNP203) or dark on first eight supralabials (UNP196) (Fig. 5). Stripe between eye and nostril absent (UNP045, UNP052, UNP060, UNP148, UNP165, UNP193), faint (UNP167, UNP194, UNP199, UNP203), or stronger (UNP196). Lateral stripe that extends beyond the arm could be slightly discontinuous right in the arm insertion (UNP167), while the others are continuous. The medium part of U-shaped mark is discontinuous in UNP165, while the others are continuous (Fig. 4). The bands on the trunk continuous (UNP045, UNP193, UNP167, UNP194, UNP196), while others are irregular (UNP052, UNP199, UNP203) or blotched (UNP060, UNP148, UNP165).

Figure 5. 

Variation of labial color, stripe between eye and nostril and body bands in lateral view A UNP167 B UNP165 C UNP196.

Distribution, habitat and natural history

All individuals were captured in Sumatra Barat Province, from the location between c. 83 meter above sea level (m asl) to c. 700 m asl. In Sungai Barameh, UNP052 was captured when it was sticking on a vertical cement wall approximately 70 cm above the ground; UNP053 was found on a vertical metal roofing sheet which was similar to UNP052 in height; UNP055, a juvenile, was found perching on a fern leave. In Malibo Anai, UNP045, UNP047, UNP048 were found in a similar habitat, sticking on a vertical stem tree approximately 1 m from the ground. In Lembah Anai Nature Reserve, UNP060-062 were found similar to those found in Malibo Anai. However, in Lembah Anai Nature Reserve, we observed an uncollected individual on the forest floor with leaf litter. In Sarasah Uwak Waterfall, we found UNP146 perching on a horizontal branch of a herb approximately 1 m above the ground. In village Sungai Sirah, UNP165 and UNP167 were found perching on a vertical stem of a tree approximately 1 m above the ground. In Langkuik Tamiang waterfall, UNP192 was found on a rock in the wall of a small rocky stream; UNP193 was found on herb branch; and UNP194 was found on a horizontal dead tree approximately 1.2 m above the ground. In village Koto Malintang, we found UNP199 and UNP203 perching on a herb branch approximately 1 m above the ground. There was one female carrying two eggs that can be seen inside the abdomen (UNP165; see Fig. 4).


We found significant support for separation of the new species from the sister clade. However, our results also showed a low bootstrap (<70) branch between the C. semicinctus clade and the C. semenanjungensis clade which indicate the topology of this branch remains unstable.

Genetically, the new species is closely related to C. semenanjungensis which was supported by the similar morphological characters between them such as the U-shaped mark on occiput, yellow labials, and the brachial tuberculation. Other characters such as the postocular stripe extending beyond the arm and yellow spots on the labials are more similar to C. psarops. The greatly enlarged scale at the apex of the pore-bearing series, as mentioned by a previous study (Harvey et al. 2015), is also possessed by C. gonjong sp. nov. indicating that this character is a synapomorphy within Sumatran Cyrtodactylus. The absence of the enlarged scale at the apex of the pore-bearing series in Malaysian C. semenanjungensis is a variation shared between Sumatran species with Malay Peninsular species.

Compared to Malaysia, Myanmar or Vietnam, herpetological research on Sumatra island is far from enough, especially for Cyrtodactylus taxa. According to our results, the diversity of Sumatran Cyrtodactylus remains unresolved. Despite the additional species of Cyrtodactylus, Sumatra still has lower diversity compared to Peninsula Malaysia (7 vs 26). It strongly suggests that the sampling effort has not been sufficient to uncover the diversity. For example, O’Connell et al. (2019) revealed that there are at least six undescribed lineages of Cyrtodactylus along Sumatra island. Our study also revealed that even in lowland forest, there is a commonly encountered, but undescribed taxon new to science. This discovery supported the dispersal scenario of Cyrtodactylus suggested by O’Connell et al. (2019) in which Sumatra was invaded by lowland species from the Thai-Malay Peninsula (indicated by sister-taxon groupings between C. gonjong sp. nov. and C. semenanjungensis). Perhaps, the lowland connection occurred during the Miocene which then facilitated dispersal of this species. This discovery also explains O’Connell’s study which predicted lowland species from Sumatra likely exist but were unsampled.


The authors would like to thank (1) Lembaga Penelitian dan Pengabdian Masyarakat Universitas Negeri Padang for funding this work with a contract number 1688/UN35.13/LT/2022 to Yuni Ahda, (2) Universitas Brawijaya with a contract number 1074.3/UN10.C10/PN/2022 to Nia Kurniawan and (3) Universitas Andalas with a contract number T/17/UN.16.17/PT.01.03/IS-RKI-A(MITRA/2022) to Djong Hon Tjong. We also thank M. Rafi, K. Agusdi and F. R. Octavian for helping us in the fieldwork. The permission to carry out survey within conservation area was issued by Balai Konservasi Sumberdaya Alam Sumatra Barat (letter number: SI.496/K.9/TU/DTN/3/2021).

Additional information

Conflict of interest

The authors have declared that no competing interests exist.

Ethical statement

No ethical statement was reported.


This work was supported by Lembaga Penelitian dan Pengabdian Masyarakat Universitas Negeri Padang, Universitas Brawijaya, Universitas Andalas.

Author contributions

Conceptualization: NK, SML. Data curation: FADN, DHT, MAF. Formal analysis: FADN, MAF. Funding acquisition: NK, YA, DHT. Investigation: FADN. Methodology: NK, AR. Project administration: FADN, MAF. Resources: SML, YA. Software: FADN. Supervision: SML, AR. Validation: AR. Visualization: FADN. Writing – original draft: FADN. Writing – review and editing: SML, YA, DHT, NK, AR, FADN.

Author ORCIDs

Fitra Arya Dwi Nugraha

Yuni Ahda

Djong Hon Tjong

Awal Riyanto

Muhammad Alif Fauzi

Si-Min Lin

Data availability

All of the data that support the findings of this study are available in the main text or Supplementary Information.


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