Research Article |
Corresponding author: Yuni Ahda ( ahdayuni@fmipa.unp.ac.id ) Academic editor: Anthony Herrel
© 2023 Yuni Ahda, Fitra Arya Dwi Nugraha, Djong Hon Tjong, Nia Kurniawan, Yunico Amardi, Muhammad Alif Fauzi, Si-Min Lin.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Ahda Y, Nugraha FAD, Hon Tjong D, Kurniawan N, Amardi Y, Fauzi MA, Lin S-M (2023) A new species of the Cyrtodactylus quadrivirgatus complex (Chordata, Reptilia, Squamata, Gekkonidae) from Sumatra Barat, Indonesia. ZooKeys 1168: 367-386. https://doi.org/10.3897/zookeys.1168.98724
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Among the six species of Cyrtodactylus occurring in Sumatra, two species were described based on non-Sumatran type series, C. consobrinus and C. quadrivirgatus. The latter species was described originally from Thailand thus the wider distribution in Sumatra should be clarified taxonomically. Cyrtodactylus quadrivirgatus from Sumatra Barat was examined using both morphology and the Natrium Dehydrogenase Subunit 2 (ND2) gene to clarify its taxonomic status and phylogenetic placement. It was found that these specimens form a sister clade to all other species of the sworderi group from Peninsular Malaysia and the genetic distance ranges from 20–24.3%. This subset is herein described as a new species. The new species is readily distinguished from C. quadrivirgatus and other Sumatran species by a combination of characters: small size SVL 37.5–53.78 mm; longitudinal rows of dorsal tubercles 16–19; paravertebral tubercles 31–41; ventral scales 32–43; 24–49 enlarged precloacal and femoral scales; precloacal pores rarely present; no precloacal depression; two postcloacal tubercles on each side; 14–19 subdigital lamellae on forth toe; 9–15 supralabial scales; 9–12 infralabial scales; three or four internasal scales; and 3–6 gular scales that border the first pair of postmental scales. This work underscores the importance of clarifying widely distributed species for taxonomic validation.
Distribution, evolution, molecular, morphology, ND2 gene, systematics, taxonomy
Cyrtodactylus quadrivirgatus Taylor, 1962 was originally described from Khao Chong Forest Experiment Station, Trang Province, Thailand. It ranges from southern Thailand, Peninsular Malaysia and adjacent islands, Singapore to northern Sumatra (
Along Peninsular Malaysia, the populations of C. quadrivirgatus exhibit coloration differences among different localities. The south population has four dark dorsal stripes, the upland population has two dorsolateral stripes and medial blotches, and the other populations possess only blotches instead of stripes. Although there was obvious variation among populations, the ND2 p-distance showed that they were separated from each other by 3.3%–5.8% (
Meanwhile, the population from Sumatra was not examined either morphologically or molecularly, leaving this population unknown in term of its taxonomic status and phylogenetic placement. We began surveying Cyrtodactylus Gray, 1827 in Sumatra Barat Province in 2020 and found them from lower elevations, approximately 8 m a.s.l. to 712 m a.s.l. Through careful examination, we wanted to establish whether C. quadrivirgatus from Sumatra Barat should be treated as a distinct species and into which lineage it fell.
Field surveys were undertaken in the province of Sumatra Barat: Lembah Anai Nature Reserve (LANR) (0°29'24"S, 100°20'24"E), around Sarasah Gasang waterfall (SG) (0.31°S, 100.23°E), around Sungai Sirah village (SS) (0°24'8.8128"S, 100°8'37.6728"E), around Sarasah Uwak waterfall (0°54'28"S, 100°28'54"E) and in Bungus Selatan village (1°02'20"S, 100°24'50"E). Individuals were all collected during the night from 19.00–23.00 hours by hand. Anaesthetization and euthanization were done using benzocaine and fixation using 10% formalin. The specimens were stored in 70% alcohol and the livers were stored in 95% ethanol. All photographs were deposited at the
Department of Biology, Universitas Negeri Padang, Indonesia (UNP). All specimens will be deposited at
Museum Zoologicum Bogoriense, Bogor, Indonesia (
Color notes were observed from digital images of living individuals prior to preservation. If in case the individual displayed stress coloration, we placed them in the cage mimicking the natural habitat and waited until the natural coloration appeared. The individuals under the stress condition showed black coloration along their dorsum, causing the disappearance of the black stripes and bands on the dorsum.
The following measurements were taken with a dial caliper to the nearest 0.5 mm following
SVL Snout-vent length, measured from the tip of snout to the vent;
AX Axial length, measured from the posterior margin of the forelimb at its insertion point on the body to the anterior margin of the hind limb at its insertion point on the body;
TL Tail length, measured from the vent to the tip of the tail, original or regenerated;
AL Arm length, measured insertion of antebrachium with body wall to claw of longest finger;
LL Leg length, measured insertion of femur with body wall to claw of longest toe;
HL Head length, measured from tip of snout to articulation of quadrate bone;
HW Head width, measured at level of ear openings;
HH Head height, measured at level of ear opening;
SL Snout length, measured from tip of snout to anterior margin of orbit;
OEL Orbit-ear length, measured from posterior margin of orbit 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;
IN Internarial distance, measured between the nares across the rostrum;
EN Eye to nostril distance, measured between the anterior margin of the eyeball to the posterior margin of the external nares.
Meristic counts included:
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 precloacal and femoral scales, number of enlarged precloaco-femoral scales, counted along lowest, pore-bearing row;
PP Precloacal pores, number of precloacal pores;
PFP Precloacal and femoral pores, number of precloaco-femoral pores;
PCT Postcloacal tubercles, number of postcloacal tubercles;
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;
SLL Left supralabial, labial scales of upper jaw, beginning with first enlarged scale bordering rostral shield, ending with last enlarged scale bordering labial angle for left side;
SLR Right supralabial, labial scales of upper jaw, beginning with first enlarged scale bordering rostral shield, ending with last enlarged scale bordering labial angle for right side;
ILL Left infralabial, labial scales of lower jaw, beginning with first scale bordering mental shield, ending with last enlarged scale bordering labial angle for left side;
ILR Right infralabial, labial scales of lower jaw, beginning with first scale bordering mental shield, ending with last enlarged scale bordering labial angle for right 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 scales (supralabials and infralabials, precloaco-femoral scales) and detecting the presence of pores, we used a staining technique with methylene blue in 70% alcohol (
Total genomic DNA was extracted from the livers using the Qiagen DNeasy tissue kit (Valencia, CA, USA) following the standard protocol for animal tissue. The amplification of the Natrium Dehydrogenase Subunit 2 (ND2) gene and partial flanking tRNAs was done by using Polymerase Chain Reaction (PCR) under the following condition: 2 min at 95 °C followed by 33 cycles of 95 °C for 35 s, annealing at 54 °C for 35 s, extension at 72 °C for 35 s and a final extension step of 10 min at 72 °C. Amplifications were carried out in 25-µl volume vials consisting of 2.5 µl genomic DNA (concentration: approximately 100 ng), 0.4 µм each primer and 1× GoTaq Green Master Mix (Promega, Wisconsin, USA). The primers used in this study followed
Sequences were uploaded, assembled, and edited in Geneious Prime 2022.2.2 (http://www.geneious.com/). All sequences, ingroup and outgroup (Table
Species of Cyrtodactylus 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 |
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agamensis group | ||||
C. metropolis | Batu caves, Selangor, PM | LSUHC 11343 | KU253579 |
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C. payacola | Bukit Panchor, Penang, PM | LSUHC 10070 | JQ889190 |
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C. majulah | Nee Soon Swamp, Singapore | ZRC 26951 | JX988529 |
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C. pantiensis | Gn. Panti, Johor, PM | LSUHC 8905 | JQ889186 |
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C. tiomanensis | Pahang, PM | LSUHC 6251 | JX440563 |
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C. rosichonariefi | Bunguran, Great Natuna, Indonesia |
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KP256187 |
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C. psarops | Indonesia |
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MH248931 |
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C. sp. 3 | Indonesia | ENS 18140 | MH248911 |
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C. sp. 4 | Indonesia | ENS 18591 | MH248912 |
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C. sp. 5 | Indonesia | ENS 18659 | MH248916 |
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C. sp. 6 | Indonesia | ENS 18719 | MH248917 |
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C. semenanjungensis | Gn. Panti, Johor, PM | LSUHC 8900 | JQ889177 |
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C. semicinctus | Indonesia | ENS 14749 | MH248925 |
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C. cf. agamensis | Indonesia | ENS 19634 | MH248908 |
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sworderi group | ||||
C. quadrivirgatus | Bukit Larut, Perak, PM | LSUHC 8859 | JQ889241 |
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C. guakanthanensis | Gua Kanthan, Perak, PM | LSUHC 11323 | KU253577 |
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C. tebuensis | Gn. Tebu, Terengganu, PM | LSUHC 10902 | JX988527 |
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C. sworderi | Sungai Kawal, Peta, PM | LSUHC 7685 | JQ889189 |
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C. gunungsenyumensis | Hutan Lipur Gn. Senyum, Pahang, PM | LSUHC 12201 | KU253585 |
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C. awalriyantoi sp.nov. | Sungai Geringging, Padang Pariaman | UNP 153 | OR122991 | This study |
UNP 161 | OR122987 | |||
UNP 162 | OR122988 | |||
UNP 163 | OR122989 | |||
UNP 164 | OR122990 | |||
lateralis group | ||||
C. lateralis | Indonesia | UTA 62916 | KU893163 |
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C. rubidus | – | CES 131445 | KM255203 |
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C. durio | Malaysia | LSUHC 9725 | KU893159 |
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marmoratus group | ||||
C. marmoratus | Indonesia | ENS 15932 | KR921721 |
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C. papuensis | – | SAMA R62652 | JQ820320 |
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C. sp. 1 | Indonesia | ENS 15813 | KR921697 |
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C. sp. 2 | Indonesia | ENS 15784 | KR921689 |
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darmandvillei group | ||||
C. batucolus | Pulau Besar, Melaka, PM | LSUHC 8934 | JQ889179 |
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C. petani | Pasuruan, Jawa Timur, Indonesia |
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KU232620 |
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C. kimberleyensis | Siuna, Sulawesi Tengah, Pulau Sulawesi, Indonesia | WAM R164144 | JX440544 |
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C. jellesmae | Siuna, Sulawesi Tengah, Pulau Sulawesi, Indonesia | RMB 1672 | GU550721 |
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C. sadleiri | Christmas island, Australia | SAMA R34810 | JQ820309 |
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C. seribuatensis | Pulau Nangka Kecil, Johor, PM | LSUHC 6349 | JQ889187 |
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C. darmandvillei | Nusa Tenggara Barat, Indonesia | WAM R98393 | JX440533 |
|
Outgroup | ||||
Hemidactylus frenatus | – | LLG 4871 | GQ458049 |
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Gekko gecko | Thailand: Patong Beach, Kathu District, Phuket Island, Phuket Province | MVZ 215314 | AF114249 |
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We used 969–1005 bases of ND2 gene sequence from the new putative species to build a ML phylogenetic tree. Our ML tree (Fig.
Uncorrected p-distance (in %) of the ND2 gene calculated for the new species and sworderi and agamensis groups. For the accession numbers for each species, refer to Table
No. | Species | 1 | 2 | 3 | 4 | 5 | 6 | 7 | 8 |
---|---|---|---|---|---|---|---|---|---|
1 | C. awalriyantoi sp. nov. | 0–0.5 | |||||||
2 | C. quadrivirgatus | 20–20.6 | |||||||
3 | C. guakanthanensis | 22.8–23.5 | 19.5 | ||||||
4 | C. sworderi | 23.7–24.3 | 21.7 | 15.2 | |||||
5 | C. tebuensis | 20.9–21.5 | 20.2 | 13.4 | 17.1 | ||||
6 | C. gunungsenyumensis | 22.5–23.2 | 20.4 | 14.3 | 17.1 | 7.7 | |||
7 | C. semenanjungensis | 32.3–33.1 | 26.5 | 31.4 | 32.1 | 30.4 | 28.3 | ||
8 | C. semicinctus | 30.4–31.1 | 23.4 | 25.4 | 28.6 | 29.2 | 26.8 | 17.3 | |
9 | C. psarops | 33.6–34.3 | 27.4 | 28.3 | 35 | 31.3 | 26.8 | 26.1 | 22.7 |
Holotype. Adult male, UNP070 (Fig.
Cyrtodactylus awalriyantoi sp. nov. is assigned to the sworderi group based on its phylogenetic position (Fig.
This species is the smallest Cyrtodactylus species inhabiting Sumatra with the maximum SVL of adult individual of 53.78 mm. It can be distinguished from other Cyrtodactylus as follows:
C. awalriyantoi sp. nov. has unique morphological combination and can be separated from other congeners within the sworderi group as follows:
Small-sized Cyrtodactylus with SVL of 37.5–53.78 mm; the length of the tail is 31.4–54.77 mm including the original or regenerated tip; the axial body length is 16.65–24.31 mm (Fig.
The nares are oval, bordered by rostral anteriorly, by supranasals and internasals dorsally, by 1st supralabial ventrally. Supranasal scales larger than post-nasal scales. The supranasal scales as large as intersupranasals and separated from each another by three or four intersupranasal scales (Fig.
The triangular mental is bordered laterally by first infralabial and posteriorly by right and left first postmental. First postmentals medially connected each other for ~ 30% of their length. Second postmentals in contact with 1st and 2nd infralabials (N = 2) (Fig.
Body moderate in length (AX/SVL 0.38–0.53); defined ventrolateral fold with tubercles smaller than dorsal tubercles; dorsum with small scales interspersed with large conical or pyramidal, tubercles most dense on flanks; tubercles extending from occipital region to the base of tail, tubercles on tail largest; 16–19 tubercles between lateral fold in middle of body; 31–41 tubercles of paravertebral from posterior insertion of arm to body to anterior of femur insertion to body; 32–43 ventral scales larger than dorsal scales; ventral scales in middle part slightly larger than those near the ventrolateral folds; from middle of body, scales are smaller anteriorly to the head, ventrum, and posteriorly until groin region (Fig.
Forelimbs medium length (AL/SVL 0.33–0.4); granular scales on upper arm larger than those on dorsum of body (~ 2–3 ×larger); without tubercles; lower arm with smaller scales than upper arm scales, intermixed with weak tubercles slightly larger than weak tubercles on parietal parts; hindlimbs also moderate in size (LL/SVL 0.40–0.52); more robust than forelimbs; covered dorsally by granular scales intermixed with large, rounded tubercles; ventral scales of thigh larger than dorsals; 14–19 subdigital lamellae on 4th toe. Continuous enlarged precloacal and femoral scales present (N = 24–49); no specimen has precloacal groove/ depression; enlarged post-precloacal scales present; two post-cloaca tubercles on left and right base of tail, mostly connected to each another (Fig.
Tail length ~ 1.1 × of SVL, circular in cross-section but tapering at the end portion; tubercles on base of tail dorsally similar in size to those on body dorsum; 4–11 black dorsoventral stripes separated by white stripes; black stripes on venter more faded than on dorsal; part; no median, transversely enlarged, series of scales on the subcaudal; subcaudal cycloidal scales relatively larger than dorsal (Fig.
Ground color of body dorsum dark grey to brown; top of head blackish with irregular broken spots scattered on parietal region to nostril; on occipital regions three short black lines extending longitudinally: one in the middle, two begin behind each side of eyes almost parallel to the supraorbital regions; those three short black bands stop at approximately parallel to ears, after which there is a transverse white line extending from each pre-ear region; after the white line, there are two black lines at the nape of the neck that extend backwards, then some meet at an angle and some remain separate, as if these two lines continue the black line originating from the back of the eye parallel to the supraorbital area; after the meeting, there are two lines that separate to the back of the tail, and some are still united to the tail so that it tends to look like a black transverse band; in individuals with the two midlines converging, the confluence of the two lines begins just before the anterior part of the upper arm; there are eight or nine rows of black transverse bands that are counted from the beginning of the union of the two lines to the base of the tail; on the dorsolateral, there are two black lines that extend from behind the eyes to the base of the tail; unpatterned black blotches or obscure irregular black banding on limbs; black and white bands on tails; the width of the black line increases towards the posterior; and the white is opposite; in some individuals, the above-mentioned black stripes are not clear and not strong along the dorsal and dorsolateral body. Ventral surface of head, trunk, and limbs are white, pale grey to cream; ventral surface of tail cream in the first third at the anterior, then the rest to the posterior tends to black with narrow white rings (Fig.
Ground color of dorsal trunk, limbs, and tail brown to dark; parietal part to the tip of snout paler than any other parts of dorsum; the individuals with unclear or weak black lines on middle dorsum and dorsolateral tend to be dark from the nape to the base of the tails; black lines on nape and trunk still visible; tail with black and white bands; ventral head, trunk, limbs whitish to dark brown. Fresh specimens darker than the others both in ventral or dorsal parts of the body (Fig.
We collected the type series in the primary forest of LANR, SG, and SS with elevation ~ 380–767 m a.s.l. and we encountered non-vouchered individuals from ~ 7 m a.s.l. At SG, this species was found on leaves measuring ~ 7–10 cm width and on twigs, ~ 1 m above the ground, 1–3 m from the edge of the rocky stream. The stream that empties into the waterfall has a breadth of ~ 2 m with a heavy flow. Fewer specimens were found closer to the waterfall. At LANR and SS, this species occupied the same microhabitat as the SG population, but the stream at this location is wider (~ 5–7 m width; Fig.
Currently, this new species is found only in Sumatra.
The specific epithet awalriyantoi is in reference to the Indonesian herpetologist, Awal Riyanto. He has dedicated much of his time researching Indonesian Cyrtodactylus from Indonesia, as well as patiently and continuously supervising many younger amphibian and reptile taxonomists from both academic institutions and independent positions. Moreover, his contribution to the study of amphibians and other reptiles is significant for Indonesian herpetological knowledge and conservation.
Previously, the sworderi group of Cyrtodactylus contained five species of which four are endemic to Peninsular Malaysia: living in lowland swampy habitats (C. sworderi;
Widely distributed species in Cyrtodactylus are most likely questionable, for example, two potentially new species have been detected within the C. marmoratus complex from southern Sumatra (
The authors would like to thank Lembaga Penelitian dan Pengabdian Masyarakat Universitas Negeri Padang for funding this work under contract number 1688/UN35.13/LT/2022 to Yuni Ahda, Universitas Brawijaya support with contract number 1074.3/UN10.C10/PN/2022 to Nia Kurniawan, and 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 the survey within a conservation area was issued by Balai Konservasi Sumberdaya Alam Sumatra Barat (letter reference SI.496/K.9/TU/DTN/3/2021).
The authors have declared that no competing interests exist.
No ethical statement was reported.
Lembaga Penelitian dan Pengabdian Masyarakat Universitas Negeri Padang under contract number 1688/UN35.13/LT/2022; Yuni Ahda, Universitas Brawijaya support with contract number 1074.3/UN10.C10/PN/2022; Nia Kurniawan, and Universitas Andalas with a contract number T/17/UN.16.17/PT.01.03/IS-RKI-A(MITRA/2022) to Djong Hon Tjong.
Conceptualization: YA, DHT. Data curation: MAF, YA. Formal analysis: NK, DHT, MAF. Funding acquisition: SML, DHT, NK, YA. Investigation: FADN, YA. Methodology: YA. Project administration: MAF, FADN. Resources: FADN. Software: FADN. Supervision: SML. Validation: SML. Visualization: NK. Writing – original draft: YA, FADN. Writing – review and editing: SML, YA, FADN, YA, DHT.
Yuni Ahda https://orcid.org/0000-0003-0545-2965
Fitra Arya Dwi Nugraha https://orcid.org/0000-0002-0048-8515
Djong Hon Tjong https://orcid.org/0000-0002-4743-9964
Muhammad Alif Fauzi https://orcid.org/0000-0003-0975-9681
Si-Min Lin https://orcid.org/0000-0001-7080-706X
All of the data that support the findings of this study are available in the main text or Supplementary Information.
Morphometric and meristic data
Data type: Morphology (.xlsx file)
Explanation note: This file contain the measurement on morphometric characters and meristic, examined on the type series specimen.
Comparison with C. quadrivirgatus
Data type: Morphology (.xlsx file)
Explanation note: This file contains the comparison between C. awalriyantoi sp. nov. with C. quadrivirgatus from type series (