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Research Article
A new species of the genus Calamaria H. Boie in F. Boie, 1827 (Squamata, Calamariidae) from Xishuangbanna, Yunnan Province, China
expand article infoTierui Zhang, Yuhao Xu§, Tan Van Nguyen|, Nikolay A. Poyarkov, Gernot Vogel#, Xinge Wang, Song Huang
‡ Anhui Normal University, Anhui, China
§ Qinghai University, Xining, China
| Duy Tan University, Da Nang, Vietnam
¶ Lomonosov Moscow State University, Moscow, Russia
# Society for South East Asian Herpetology, Heidelberg, Germany
Open Access

Abstract

A new species of reed snake, Calamaria synergis sp. nov., is described based on two specimens collected from Mountain Jinuo, Jinghong City, Xishuangbanna Dai Autonomous Prefecture, Yunnan Province, China. The new species is distinguished from its congeners by a unique combination of morphological characters, including eight enlarged maxillary teeth; rostral higher than wide; prefrontal shorter than frontal and contacting the first two supralabials; mental not in contact with anterior chin shields; single preocular and postocular; four supralabials, with the 2nd and 3rd contacting the eye; five infralabials; five scales surrounding the paraparietal; ventrals 161–166; subcaudals 20–23, paired; dorsal scales reduced to six rows at the tail base and further to four rows near the terminal subcaudals; tail relatively short (6.6–9.2% of total length), abruptly tapering at the tip; dorsum blackish-brown with a distinct pale nuchal ring; two outermost dorsal scale rows pale khaki with upper margins darkened; ventral surface uniform pale khaki. Phylogenetic analysis of the mitochondrial cytochrome b gene places the new species as sister to C. andersoni and C. yunnanensis, from which it differs by an uncorrected p-distance of 8.7% and 7.9%, respectively. Calamaria synergis sp. nov. is currently known only from tropical evergreen forests of Xishuangbanna at elevations around 1,050 m asl. We propose its conservation status as Data Deficient (DD) following the IUCN Red List categories.

Key words:

Calamaria synergis sp. nov., morphology, Mountain Jinuo, phylogenetics, taxonomy

Introduction

The genus Calamaria H. Boie in F. Boie, 1827, commonly known as Southeast Asian reed snakes, currently comprises 69 valid species distributed from northeastern India and southern Japan to the Maluku Islands of eastern Indonesia (Inger and Marx 1965; Uetz et al. 2025). Of these, 18 species have been currently recorded from China and the Indochinese region (including Laos, Cambodia, Thailand, and Vietnam), namely: C. abramovi Orlov, 2009; C. arcana Yeung, Lau & Yang, 2022; C. andersoni Yang & Zheng, 2018; C. buchi Marx & Inger, 1955; C. berezowskii Günther, 1896; C. concolor Orlov, Nguyen, Nguyen, Ananjeva & Ho, 2010; C. dominici Ziegler, Tran & Nguyen, 2019; C. gialaiensis Ziegler, Nguyen & Nguyen, 2008; C. jinggangensis Cai, Jiang, Wu, Huang, Fei & Ding, 2023; C. lovii Boulenger, 1887; C. lumbricoidea Boie, 1827; C. nebulosa Lee, 2021; C. pavimentata Duméril, Bibron & Duméril, 1854; C. sangi Nguyen, Koch & Ziegler, 2009; C. schlegeli Duméril, Bibron & Duméril, 1854; C. septentrionalis Boulenger, 1890; C. strigiventris Poyarkov, Nguyen, Orlov & Vogel, 2019; C. thanhi Ziegler & Le, 2005; and C. yunnanensis Chernov, 1962 (Poyarkov et al. 2019, 2023; Cai et al. 2023; Liang et al. 2024; Uetz et al. 2025). Notably, six species have been described or revalidated within the past eight years, suggesting that the diversity and taxonomy of Calamaria in the montane regions of China and Indochina remain incompletely understood.

During recent fieldwork in Mountain Jinuo, Jinghong City, Xishuangbanna Dai Autonomous Prefecture, Yunnan Province, China, we collected two unidentified snake specimens assignable to the genus Calamaria based on the following diagnostic characters: a cylindrical, vermiform body; a short, thick tail; a head not distinct from the neck; small eyes with round pupils; eight subequal maxillary teeth, often modified in shape; incomplete upper head scalation typical of colubroid snakes, with internasal, loreal, and temporal scales always absent; four or five supralabials, the posterior-most broadly contacting the parietal; a large paraparietal scale behind the last supralabial; dorsal scales in 13 rows throughout the body, all smooth; a single, entire cloacal plate; and paired subcaudal scales (following Inger and Marx 1965; David et al. 2023). Subsequent morphological and molecular analyses revealed that these two individuals are distinct from all seven species of Calamaria currently known from China, as well as all other congeners reported from the Indochina region, based on a unique combination of morphological traits and significant genetic divergence. We herein describe these specimens as a new species of the genus Calamaria.

Materials and methods

Sampling

One adult male specimen and one juvenile male specimen of Calamaria were collected from Xishuangbanna Dai Autonomous Prefecture, Yunnan Province, China. These specimens were humanely euthanised with 0.7% tricaine methane sulfonate (MS-222) solution. Fresh liver tissue was extracted and immediately preserved in 95% ethanol for the subsequent molecular analysis. Specimens were preserved in 75% ethanol for permanent storage and deposited in Anhui Normal University Museum, Anhui, China (ANU). Sampling procedures involving live snakes were approved by the Animal Ethics Committee of Anhui Normal University and complied with the Wild Animals Protection Law of China.

Molecular phylogeny

Total genomic DNA was extracted from preserved liver tissue with the OMEGA Tissue DNA Kit D3396 (Omega Bio-Tek, Norcross, GA, USA). A fragment of the mitochondrial cytochrome b (Cyt b) gene was amplified via Polymerase Chain Reaction (PCR) using the primer pair L14910 (5’-GACCTGTGATMTGAAACCAYCGTTGT-3’) and H16064 (5’-CTTTGGTTTACAAGAACAATGCTTTA-3’) (Burbrink et al. 2000). The double-stranded PCR products were sequenced by General Biosystems (Anhui) Corp. Ltd. (Chuzhou, China), and raw sequences were assembled using SeqMan in the DNASTAR software package (Burland 2000).

A total of 42 sequences from 14 known Calamaria species and three out-group species, including Elaphe quatuorlineata (Lacépède), Orientocoluber spinalis (Peters), and Lycodon rufozonatus Cantor, were obtained from GenBank or newly sequenced and incorporated into our dataset (see Table 1). DNA sequences were aligned by the Clustal W algorithm with default parameters (Thompson et al. 1997) and trimmed with gaps partially deleted in MEGA X (Kumar et al. 2018). Bayesian inferences (BI) were conducted in MRBAYES v. 3.2.7a (Ronquist et al. 2012) under the GTR + I + G model on Phylosuite v. 1.2.3 (Zhang et al. 2020; Xiang et al. 2023). In the BI analysis, three independent runs were conducted with 1 × 107 generations and sampled every 1,000 generations, with the first 25% of samples discarded as burn-in. In the ML analysis, the bootstrap consensus tree was inferred from 1000 replicates. Maximum likelihood (ML) was conducted under the best-fit substitution model (GTR + I + G) in RaxmlGUI 1.3 (Silvestro and Michalak 2012). Bootstrap proportions (BSP) were investigated with 1,000 bootstrap replicates using the fast-bootstrapping algorithm. Uncorrected pairwise genetic distances (p-distance) of the Cyt b gene among Calamaria species examined were calculated with MEGA X (Kumar et al. 2018).

Table 1.

DNA sequences, voucher specimens, and GenBank accession numbers of the genus Calamaria and outgroup taxa used in this study.

Species Specimen voucher no. Locality GenBank Sources
Calamaria synergis sp. nov. AHNU ZR24046 Mt. Jinuo, Xishuangbanna, Yunnan, China PV745121 This study
Calamaria synergis sp. nov. AHNU ZR25021 Mt. Jinuo, Xishuangbanna, Yunnan, China PV745122 This study
C. alcalai PNM 9873 Sitio Palbong, Barangay Batong Buhay, Sablayan, Mindoro, Philippines MT819383 Weinell et al. 2020
C. andersoni SYS r001699 Yingjiang, Yunnan, China MH445955 Yang and Zheng 2018
C. andersoni HS R20101 Dehong, Yunnan, China OQ354844 Cai et al. 2023
C. andersoni HS R20181 Tengchong, Yunnan, China OQ354845 Cai et al. 2023
C. andersoni AHNU ZR25021 Mangshi, Dehong, Yunnan, China PV745123 This study
C. arcana KFBG 14611 Mt. Dadongshan, Guangdong, China ON482335 Yeung et al. 2022
C. arcana HS 17082 Mt. Dawu, Guangdong, China OQ354835 Cai et al. 2023
C. arcana GP 9975 Yongxing, Hunan, China OP980549 Cai et al. 2023
C. arcana DL R199 Mt. Wuyi, Fujian, China OQ354834 Cai et al. 2023
C. berezowskii GXNU DLR194 Mt. Gongga, Sichuan, China PP747047 Liang et al. 2024
C. berezowskii GXNU DLR195 Mt. Gongga, Sichuan, China PP747048 Liang et al. 2024
C. berezowskii GXNU 20221215002 Mt. Gongga, Sichuan, China PP747049 Liang et al. 2024
C. gervaisii KU 324661 Puguis, La Trinidad, Benguet, Luzon, Philippines MT819384 Weinell et al. 2020
C. gervaisii KU 334485 Narvacan, Ilocos Sur, Luzon, Philippines MT819385 Weinell et al. 2020
C. jinggangensis DL 20200725 Mt. Jinggangshan, Jiangxi, China OQ354830 Cai et al. 2023
C. jinggangensis DL 20200625-2 Mt. Jinggangshan, Jiangxi, China OQ354831 Cai et al. 2023
C. jinggangensis DL 20200625-3 Mt. Jinggangshan, Jiangxi, China OQ354832 Cai et al. 2023
C. jinggangensis DL 20200625-4 Mt. Jinggangshan, Jiangxi, China OQ354833 Cai et al. 2023
C. lumbricoidea KU 315159 Pasonanca NP, Zamboanga, Philippines MT819388 Weinell et al. 2020
C. lumbricoidea KU 334479 Mt. Lumot, Gingoog, Misamis, Philippines MT819389 Weinell et al. 2020
C. cf. lumbricoidea USMHC 1560 Air Itam Dam, Penang, Malaysia MN338526 Quah et al. 2020
C. muelleri TNHC 58955 Gowa, South Sulawesi, Indonesia MT819390 Weinell et al. 2020
C. muelleri RMB 1283 Gowa, South Sulawesi, Indonesia MT819391 Weinell et al. 2020
C. nebulosa FMNH 258666 Phongsaly, Laos MN338524 Quah et al. 2020
C. palavanensis KU 309445 Barangay Irawan, Puerto Princessa, Palawan, Philippines MT819386 Weinell et al. 2020
C. palavanensis KU 311411 Mt. Mantalingahan, Rizal, Palawan, Philippines MT819387 Weinell et al. 2020
C. pavimentata KFBG 14507 Ningming, Guangxi, China MH445957 Yang and Zheng 2018
C. schlegeli LSUHC 10278 Bukit Larut, Perak, Malaysia MN338525 Quah et al. 2020
C. septentrionalis FTB 2839 unknown locality KR814699 Pyron unpublished data
C. septentrionalis KFBG 14506 Hainan, China MH445956 Yang and Zheng 2018
C. septentrionalis HS 11119 (CHS 116) Tunxi, Huangshan, Anhui, China MK201273 Li et al. 2020
C. septentrionalis HS 12055 (CHS 118) Huangshan, Anhui, China MK201274 Li et al. 2020
C. septentrionalis RE 30 (CHS 302) Mangshan, Hunan, China MK201384 Li et al. 2020
C. septentrionalis SYS r000932 (CHS 613) Guangdong, China MK201434 Li et al. 2020
C. septentrionalis HS R19100 Mt. Huangshan, Anhui, China OQ354842 Cai et al. 2023
C. septentrionalis HS 11145 Mt. Nanling, Guangdong, China OQ354840 Cai et al. 2023
C. septentrionalis DL 2021610-1 Huangsha, Guangxi, China OQ354838 Cai et al. 2023
C. cf. septentrionalis ROM 35605 Phia Oac-Phia Den NP, Cao Bang, Vietnam AF471081 Lawson et al. 2005
C. cf. septentrionalis ROM 35597 Phia Oac-Phia Den NP, Cao Bang, Vietnam KX694890 Alencar et al. 2016
C. yunnanensis ROM 41547 Simao, Yunnan, China KX694891 Zaher et al. 2009
C. yunnanensis YPx 503 Yunnan, China JQ598922 Grazziotin et al. 2012
C. yunnanensis QHU R2024054 Mt. Wanzhangshan, Simao, Yunnan, China PV755783 This study
Outgroup
Orientocoluber spinalis MVZ 211019 Ningxia, China AY486924 Nagy et al. 2004
Elaphe quatuorlineata LSUMZ 40626 Hungary AY486931 Nagy et al. 2004
Lycodon rufozonatus LSUMZ 44977 China AF471063 Lawson et al. 2005

Morphological analysis

Terminology and measurements follow Inger and Marx (1965). Measurements were taken with a digital slide caliper to the nearest 0.1 mm, except for body and tail lengths, which were measured to the nearest 1.0 mm with a measuring tape. The number of ventral scales was counted according to Dowling (1951). The numbers of dorsal scale rows are given at one head length behind the head, at midbody, and at one head length before the vent. The sex was determined by dissection (inspection of gonads) and/or by tail shape when dissection was not possible. Maxillary teeth of the specimens were counted by examining both maxillae using a dissecting pin under a binocular microscope prior to preservation.

The following measurements (all in mm) and counts were taken: head length (HL, from snout tip to jaw angles) ; head width (HW) ; interorbital distance (IOD) ; eye-nostril distance (EN, from anterior edge of orbit to posterior edge of nostril) ; eye diameter (EyeD, horizontal) ; eye-mouth distance (Eye-MouthD, measured from the lowest point of the eye to the mouth gap) ; snout length (SnL, from the tip of rostral to the anterior margin of the eye) ; the number of dorsal scales reducing to six rows above the position of the subcaudal anterior to the tail tip (DSR6R) ; the number of dorsal scales reducing to four rows above the position of the subcaudal anterior to the tail tip (DSR4R) ; snout-vent length (SVL) ; tail length (TaL) ; total length (TL) ; ratio of tail length/total length (TaL/TL) ; dorsal scale rows number (DSR) ; supralabial scales (SL) ; number of supralabials touching the eye (SL-E) ; infralabial scales (IL) ; preocular scales (PrO) ; postocular scales (PoO) ; subcaudal scales (SC) ; ventral scales (VEN). Asymmetric characters are given in left/right order. Other abbreviations: Mt.: Mountain; NR.: Nature Reserve; NP.: National Park; Is.: Island; asl.: above sea level. For museum abbreviations, see Suppl. material 1: table S1.

Morphological and chromatic characters of the examined specimens were compared in detail to other species of the genus Calamaria known to occur in mainland Southeast Asia and other congeners. The examined comparative material is listed in Suppl. material 1: table S2. For comparison with other taxa, we relied on previously published data (Duméril et al. 1854; Inger and Marx 1965; Darevsky and Orlov 1992; Ziegler and Le 2005; Ziegler et al. 2007, 2008, 2019; Orlov 2009; Orlov et al. 2010; Yang and Zheng 2018; Poyarkov et al. 2019; Lee 2021; Yeung et al. 2022; Cai et al. 2023; Liang et al. 2024; Nguyen et al. 2025; Zhang et al. 2025).

Results

Molecular results

A total of 1079 base pairs (bp) of the mitochondrial Cyt b gene were successfully sequenced and aligned, including two sequences from the putative new species. Both ML and BI analyses yielded congruent topologies (Fig. 1), which are consistent with those reported in previous studies (Cai et al. 2023; Liang et al. 2024). Within the current taxon sampling, the genus Calamaria was recovered as a monophyletic group with strong support (BPP = 1.00; BSP = 100). The two individuals representing Calamaria sp. from Xishuangbanna formed a distinct and well-supported clade (BPP = 1.00; BSP = 100), which was recovered as the sister lineage to a clade comprising C. andersoni and C. yunnanensis (BSP = 100; BPP = 1.00). Uncorrected p-distances between the Xishuangbanna Calamaria sp. and other congeners ranged from 7.9% (vs C. yunnanensis) to 21.6% (vs C. cf. lumbricoidea), with a p-distance of 8.7% from the closely related C. andersoni (see Table 2).

Table 2.

Uncorrected p-distances (percentage) between Calamaria species based on 1079 base pairs from the mitochondrial gene Cyt b. Remark: The GenBank accession numbers are consistent with those provided in Table 1.

No. GenBank Species 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16
1 PV745121 Calamaria synergis sp. nov.
2 MT819383 Calamaria alcalai 20.8
3 MH445955 Calamaria andersoni 8.7 21.8
4 ON482335 Calamaria arcana 14.1 19.6 16.0
5 PP747049 Calamaria berezowskii 16.5 21.1 16.2 15.3
6 MT819384 Calamaria gervaisii 20.8 17.3 20.0 19.7 20.7
7 OQ354830 Calamaria jinggangensis 14.7 20.5 16.3 5.7 14.6 19.2
8 MN338526 Calamaria cf. lumbricoidea 21.6 16.5 21.2 20.4 19.9 18.6 20.9
9 MT819389 Calamaria lumbricoidea 19.9 14.9 19.7 19.2 19.8 17.3 18.7 13.3
10 MT819391 Calamaria muelleri 19.1 14.5 20.2 17.2 19.9 15.1 15.9 15.1 12.5
11 MN338524 Calamaria nebulosa 16.4 21.4 17.2 16.4 16.6 21.0 16.5 20.0 18.4 18.0
12 MT819386 Calamaria palavanensis 19.5 14.9 20.7 21.0 23.4 15.8 18.2 16.4 16.5 15.5 19.1
13 MH445957 Calamaria pavimentata 16.8 21.8 18.6 17.0 15.8 22.0 15.7 21.5 20.4 21.8 17.7 22.2
14 MN338525 Calamaria schlegeli 18.7 14.7 20.6 19.9 21.1 17.8 19.0 16.7 14.4 15.6 18.2 16.7 19.7
15 MK201434 Calamaria septentrionalis 13.7 21.1 14.9 11.0 15.2 21.5 9.7 21.0 20.5 20.3 18.1 19.5 15.0 20.2
16 KX694890 Calamaria cf. septentrionalis 14.2 21.7 14.8 10.6 14.2 21.2 10.4 20.1 20.1 20.2 17.1 20.1 16.6 19.8 4.4
17 KX694891 Calamaria yunnanensis 7.9 21.7 9.8 16.3 17.1 22.0 14.9 21.2 20.6 22.3 16.9 22.2 17.7 20.6 14.9 14.2
Figure 1. 

The Bayesian tree of Calamaria derived from analyses of Cyt b gene. Bayesian posterior probabilities (BPP) from BI analyses/ bootstrap supports (BSP) from ML analyses are listed beside the nodes, the ones lower than 0.70 or 75 are displayed as “-” or omitted.

Given the substantial genetic divergence and the presence of clear morphological differentiation from all known congeners (see the Comparisons section below), we recognise the examined specimens as representing a distinct previously unknown species, which we herein describe as a new species of the Calamaria.

Taxonomic account

Calamaria synergis sp. nov.

Figs 2, 3A, 4; Suppl. Figs S1A, S2A; Table 3

Type material.

Holotype : ANU ZR24046, adult male from Mountain Jinuo, Jinghong City, Xishuangbanna Dai Autonomous Prefecture, Yunnan Province, China (22.009245°N, 101.014120°E; elevation 1,050 m asl), collected on 5 June 2024, by TRZ. Paratype: ANU ZR25021, one juvenile male from the same location as the holotype, collected on 7 August 2023, by TRZ.

Diagnosis.

Calamaria synergis sp. nov. can be distinguished from all other congeners by the following combination of morphological characters: eight enlarged maxillary teeth; rostral higher than wide; prefrontal shorter than frontal and contacting the first two supralabials; mental not in contact with anterior chin shields; single preocular and postocular; four supralabials, with the 2nd and 3rd contacting the eye; five infralabials; five scales surrounding the paraparietal; ventrals 161–166; subcaudals 20–23, paired; dorsal scales reduced to six rows at the tail base and further to four rows near the terminal subcaudals; tail relatively short (6.6–9.2% of total length), abruptly tapering at the tip; dorsum blackish-brown with a distinct pale nuchal ring; two outermost dorsal scale rows pale khaki with upper margins darkened; ventral surface uniform pale khaki.

Description of the holotype

(Fig. 2). Specimen ANU ZR24046 is in excellent condition. Body slender and cylindrical (SVL 258 mm, TL 284 mm); body thickness ~4.3–5.9 mm; tail not as thick as body, base of tail 3.6 mm thick. Tail short (TaL 26 mm, TaL/TL 9.2%); tail uniformly cylindrical in the anterior part, then abruptly tapering at tip; tip of tail obtusely pointed; head small, elliptical in dorsal view (HL 7.3 mm, HW 4.6 mm, HH 3.7 mm); eye small and round (EyeD 1.2 mm), larger than eye-mouth distance (Eye-MouthD 0.9 mm), ED/HL 16.4%.

Figure 2. 

Calamaria synergis sp. nov. in preservative, holotype (ANU ZR24046, adult male). A. Dorsal view of body; B. Ventral view of body; C. Dorsal view of head; D. Ventral view of head; E. Right lateral view of head; F. Left lateral view of head. Photographs by TRZ.

Rostral higher than wide (width 1.4 mm, height 1.6 mm), portion visible from dorsal aspect almost equal to the length of the prefrontal suture; prefrontal (length 1.8 mm) shorter than frontal (length 3.1 mm), not entering orbit, in contact with 1st and 2nd supralabial; frontal hexagonal, longer (length 3.1 mm) than wide (width 2.4 mm), ~3.0 times maximum width (0.8 mm) of supraocular; paraparietal surrounded by five scales; preocular 1/1 (left/right, hereinafter), higher than wide, slightly higher than postocular, not as high as eye diameter; postocular 1/1, higher than wide; nasals small, barely surrounding nostrils, surrounded by the 1st supralabial, rostral and prefrontal; supralabials 4/4, 2nd and 3rd entering orbit, 4th largest, relative supralabial width 4>2>1>3; mental triangular, not in contact with the anterior chin shields; infralabials 5/5, the first three pairs touching anterior chin shields, the first pair meeting in the midline, 4th largest; anterior chin shields longer than wide (length 2.1 mm, width 1.1 mm), pentagonal, meeting in the midline; posterior chin shields shorter than the anterior ones (length 1.6 mm), touching anteriorly and separated posteriorly by the first gular scales.

Dorsal scales in 13 rows throughout the body, reducing to six rows above the 7th subcaudal and to four rows above the penultimate pair of subcaudals. Dorsal scales homogeneous in size and entirely smooth; vertebral row not enlarged. Ventrals 161. Anal plate undivided. Subcaudals 20, paired, smooth; terminal scale single and rigid.

Dentition.

Maxillary teeth modified (enlarged), eight on each side (8/8).

Colouration of the holotype in life

(Fig. 3A). Dorsum of head and upper parts of supralabials brownish black, scattered with indistinct white flecks; ventral surface of head pale khaki, with black speckles on the infralabials and anterior portion of the anterior chin shields. The dorsal surface of the body and the tail brownish black with iridescent white flecks distributed over most dorsal scales. Two outermost dorsal scale rows are slightly pale khaki, with upper margins partially darkened, forming two indistinct, narrow black stripes along each side of the body. A distinct pale khaki nuchal ring is present, extending dorsally from the level of the 2nd–4th ventral scales, equivalent in width to two dorsal scale rows. No pale rings or blotches are present on the neck or tail. Ventral scales of the body and subcaudal scales of the tail are pale khaki, with black pigmentation on the outermost corners; small black flecks are present along the median line of the ventral tail surface, extending from the 2nd subcaudal scale to the tail tip.

Figure 3. 

In situ photographs of Calamaria synergis sp. nov. and its congeners in life. A. Calamaria synergis sp. nov. (ANU ZR24046); B. C. andersoni (ANU ZR25022); C. C. yunnanensis (QHU R2024054). Photographs by TRZ (A, B) and T.Y. Zhang (C).

Colouration of the holotype in preservative

(Fig. 2). After nearly one year in 75% ethanol, the dorsum of the body and tail remains brownish black; white flecks on the dorsal scales and narrow black stripes along the flanks are still visible. The nuchal ring and ventral surfaces of the body have faded to greyish white.

Variation.

Specimen ANU ZR25021 (paratype), a juvenile male (see Fig. 4), has a smaller body size (SVL 127 mm, TaL 9 mm, HL 6.0 mm, HW 3.1 mm, EyeD 0.8 mm, Eye-MouthD 0.5 mm). Compared to the holotype, it exhibits a higher number of ventrals (VEN 166 vs 161), subcaudals (SC 23 vs 20), and lacks black flecks along the median line of the ventral tail surface (vs present in the holotype).

Figure 4. 

Calamaria synergis sp. nov. in preservative, paratype (ANU ZR25021, juvenile male). A. Dorsal view of body; B. Ventral view of body. Photographs by TRZ.

Etymology.

The specific name synergis is a Latin noun given in the apposition, derived from the Greek synergos (συνεργός), meaning “working together”. It emphasises that the resolution of the taxonomic status of the new species is the outcome of coordinated international scientific cooperation. The name is given in reference to both the collaborative effort involved in describing this new species and the broader need for joint action to address the complex taxonomic problems within the genus Calamaria in the future. We suggest the following common names: “Mountain Jinuo reed snake” (English), “基诺两头蛇” (Chinese), “Rắn mai gầm hiệp lực” (Vietnamese), and “Цзинхунская карликовая змея” (Tszinhunskaya karlikovaya zmeya, Russian).

Comparisons.

Comparative morphological data for the new species and the currently recognised members of the genus Calamaria from Indochina to southern China are presented in Table 3 and Suppl. material 1: figs S1, S2.

Table 3.

Comparison of morphological characters of Calamaria synergis sp. nov. with those of congeners which occur in China and Indochina. Symbols: (1) = Supralabials; (2) = Supralabials contacting orbit; (3) = Preocular (1 = present, 0 = absent); (4) = Mental touching chin shields (1 = yes, 0 = no); (5) = Number of scales contacting paraparietal; (6) = VEN in males; (7) = VEN in females; (8) = SC in males; (9) = SC (females); (10) = max TL (mm) in males; (11) = max TL (mm) in females; (12) = TaL/TL (%) in males; (13) = TaL/TL (%) in females; (14) = Tail: tapering (2), slightly tapered (1), or not (0); (15) = End of tail; (16) = Dorsal colour; (17) = Ventrals colour. Notes: N/a: Not available; diagnostic characters (with respect to Calamaria synergis sp. nov.) are highlighted in bold.

Species (1) (2) (3) (4) (5) (6) (7) (8) (9) (10) (11) (12) (13) (14) (15) (16) (17) Sources
C. synergis sp. nov. 4 2/3 1 0 5 161–166 n/a 20–23 n/a 284 n/a 6.6–9.2 n/a 0 obtusely pointed blackish-brown pale khaki, unspotted This study
C. abramovi 4 2/3 1 0 6 159 174 26 20 139 482 13.3 7.1 2 sharply pointed uniformly black yellow-orange spots Orlov et al. 2009
C. andersoni 4 2/3 1 0 6 164–172 186 20–23 n/a 351 312 8.8–9.2 5.8 0 obtusely pointed brownish with faint narrow black lateral stripes bright orange to orange-yellow, unspotted Yang and Zheng 2018; Zhang et al. 2025
C. arcana 4 2/3 1 0 6 170176 192 20–22 12 303 n/a 7.2–11.8 4.7 0 obtusely pointed brownish orange-red, unspotted Yeung et al. 2022; Zhang et al. 2023
C. berezowskii 4 2/3 1 0 6 149155 153–171 22–25 12–16 290 245 6.6–10.5 6.5–6.9 0 obtusely pointed blackish-brown or brown pale khaki or white Inger and Marx 1965; Liang et al. 2024
C. buchi 4 2/3 1 1 5 n/a 221–236 n/a 13–14 n/a 466 n/a 3.9–4.1 1 obtusely pointed blackish with small pale spots yellow, unspotted Inger and Max 1965; Ziegler et al. 2008
C. concolor 5 2/3 1 1 5 209 n/a 19 n/a 578 n/a 7.2 n/a 1 obtusely pointed uniformly orange-red bright red, unspotted Orlov et al. 2010; Nguyen et al. 2025
C. dominici 4 or 5 2/3 or 3/4 1 1 or 0 6 n/a 174 n/a 17 or 18 n/a 421 n/a 6.2 1 obtusely pointed dark with irregular yellow blotches dark with few yellow blotches & bands Ziegler et al. 2019
C. gialaiensis 4 2/3 1 1 5 191 n/a 23 n/a 457 n/a 8.1 n/a 0 rounded pale greyish-brown with a faint dark neck collar and a few dark blotches along the posterior vertebral region yellowish beige, unspotted Ziegler et al. 2008
C. jinggangensis 4 2/3 1 0 6 157158 179 20 12 260 364 15 3.6 0 obtusely pointed brownish black dark orange Cai et al. 2023; Zhang et al. 2024
C. lovii ingermarxorum 4 2/3 0 1 6 205 n/a 23 n/a 318 n/a 7.4 n/a 0 blunt unspotted bluish-grey with pale spots on four lateral neck scales dark gray, unspotted Darevsky and Orlor 1992
C. lumbricoidea 5 3/4 1 1 4 or 5 144196 137–229 17–27 13–21 498 642 6.3–11.4 3.9–8.3 1 sharply pointed black with narrow cream or yellow rings; head red or pink in juveniles yellow with black ventral scales forming bands Inger and Marx 1965
C. nebulosa 4 2/3 0 0 6 n/a 179 n/a 22 n/a 354 n/a 7.9 0 obtusely pointed bluish-grey yellow, unspotted Lee 2021
C. pavimentata 4 2/3 1 0 n/a 151 n/a 27 n/a n/a n/a n/a n/a 2 sharply pointed with narrow dark longitudinal stripes and a solid black collar behind the neck yellowish white, unspotted Duméril et al. 1854; Inger and Marx 1965
C. sangi 4 2/3 1 1 5 or 6 190 n/a 19 n/a 373 n/a 6.2 n/a 1 obtusely pointed greyish-brown with fine dark mottling cream with narrow dark transverse bands Nguyen et al. 2009
C. schlegeli 5 3/4 1 or 0 0 5 or 6 129–161 136–180 2544 19–37 391 395 11.121.3 7.3–14.4 0 blunt unspotted grey or brown; head variably pink, yellow, and/or brown cream, unspotted Inger and Marx 1965
C. septentrionalis 4 2/3 1 0 6 148–166 168–188 1519 6–11 344 384 6.3–8.6 2.6–4.3 0 broadly rounded dark brown or black dorsally, usually with a narrow yellow ring ~68 scales behind the head yellow, with small black spots Inger and Marx 1965
C. strigiventris 4 2/3 1 0 6 130–168 176–183 2931 20–22 362 367 11.217.9 8.4–0.6 2 pointed uniform grey-brown bright yellow with longitudinal black stripes Poyarkov et al. 2019
C. thanhi 4 2/3 0 0 6 or 7 184 198 28 21 461 455 9.9 6.8 2 gradually to a point dark with 46 pale body bands yellow, unspotted Ziegler et al. 2005, 2007
C. yunnanensis 4 2/3 0 0 6 167184 189–199 15–21 16–19 296 516 5.4–8.4 5.0–5.5 2 obtusely pointed blue-grey to olive-brown bright orange to orange-yellow, not spotted Lee 2021; Zhang et al. 2025

Calamaria synergis sp. nov. is readily distinguished from C. lumbricoidea (distributed in Thailand, Malaysia, Singapore, Indonesia, Brunei, and the Philippines) and C. schlegeli (Thailand, Malaysia, Indonesia, and Singapore) by having four supralabials, with the second and third in contact with the eye (vs 5 supralabials with the third and fourth in contact with the eye). Furthermore, both species are restricted to the areas south of the Isthmus of Kra in Peninsular Malaysia and have not been recorded from mainland Indochina.

The new species also differs from C. lovii ingermarxorum Darevsky & Orlov (Gia Lai Province, Vietnam), C. nebulosa (Phongsaly Province, Laos; possibly northern Thailand), C. thanhi (Quang Binh Province, Vietnam), and C. yunnanensis (Yunnan Province, China; possibly northwestern Vietnam) by the presence of a preocular scale (vs absent).

Calamaria synergis sp. nov. differs from C. abramovi (Kon Tum Province, Vietnam) in having the paraparietal surrounded by five shields and scales (vs six), fewer subcaudal scales in males (SC 20–23 vs 26), a maximum total length in males (max TL 284 mm vs 139 mm), shorter relative tail length in males (6.6–9.2% vs 13.3%), a non-tapering tail (vs gradually tapering), an obtusely pointed tail tip (vs sharply pointed), a blackish-brown dorsum (vs uniformly black), and pale khaki unspotted ventrals (vs yellow-orange spots).

Compared to C. andersoni (Fig. 3B; Suppl. material 1: Figs S1B, S2B), Calamaria synergis sp. nov. differs by having five shields surrounding the paraparietal (vs 6 in C. andersoni), a smaller maximum total length in males (max TL 284 mm vs 351 mm), eight maxillary teeth (vs 9), unspotted pale khaki ventrals (vs bright orange to orange-yellow), and the presence of light neck rings (vs absent).

The new species is distinct from C. arcana (Zhejiang, Guangdong, Fujian, and Hunan provinces, China) by five shields surrounding the paraparietal (vs 6), fewer ventrals in males (VEN 161–166 vs 170–176), unspotted pale khaki ventrals (vs orangish-red), and the presence of light neck rings (vs absent).

Calamaria synergis sp. nov. is further separated from C. berezowskii (Sichuan Province, China) by having more ventrals in males (VEN 161–166 vs 149–155), five shields surrounding the paraparietal (vs six), and the presence of light neck rings (vs absent).

The new species differs from C. buchi (Lam Dong Province, Vietnam) by the mentals not contacting the chin shields (vs contacting), fewer ventrals (161–166 vs 221–236), a blackish-brown dorsum (vs blackish with small pale spots), and unspotted pale khaki ventrals (vs yellow, unspotted).

From C. concolor (Thua Thien-Hue and Quang Nam provinces, Vietnam), the new species differs in having the mental not contacting the chin shields (vs contacting), fewer ventrals in males (VEN 161–166 vs 209), a smaller maximum total length in males (max TL 284 mm vs 578 mm), a non-tapering tail (vs slightly tapered), a blackish-brown dorsum (vs uniform orangish-red), unspotted pale khaki ventrals (vs bright red), and pale neck blotches (vs absent).

The new species is distinct from C. dominici (Dak Nong Province, Vietnam) in having five shields surrounding the paraparietal (vs 6), a non-tapering tail (vs slightly tapering), a blackish-brown dorsum (vs dark with irregular yellow blotches), and unspotted pale khaki ventrals (vs dark with few yellow blotches and bands).

Compared to C. gialaiensis (Gia Lai Province, Vietnam), the new species differs by the mental not contacting the chin shields (vs contacting), fewer ventrals in males (VEN 161–166 vs 191), a smaller maximum total length in males (max TL 284 mm vs 457 mm), an obtusely pointed tail tip (vs rounded), a blackish-brown dorsum (vs pale greyish-brown with a faint dark neck collar and posterior blotches), and unspotted pale khaki ventrals (vs yellowish beige).

Calamaria synergis sp. nov. can be distinguished from C. jinggangensis (Jiangxi and Guizhou, possibly Hunan provinces, China) by five shields surrounding the paraparietal (vs 6), slightly more ventrals in males (VEN 161–166 vs 157–158), shorter relative tail length in males (ratio TaL/TL 6.6–9.2% vs 15%), and unspotted pale khaki ventrals (vs dark orange).

The new species differs from C. pavimentata (Suppl. material 1: figs S1C, S2C), which is distributed from India and Myanmar through Indochina to Peninsular Malaysia and China (including Taiwan) and southernmost Japan, by having a higher number of ventrals in males (VEN 161–166 vs 151 in the holotype of C. pavimentata), fewer subcaudals in males (SC 20–23 vs 27 in the holotype), a non-tapering tail (vs tapered), and an obtusely pointed tail tip (vs sharply pointed).

Calamaria synergis sp. nov. is distinct from C. sangi (Kon Tum Province, Vietnam) by the mental not contacting the chin shields (vs contacting), fewer ventrals in males (VEN 161–166 vs 190), a smaller maximum total length in males (max TL 284 mm vs 373 mm), a non-tapering tail (vs slightly tapered), and the presence of neck blotches (vs absent).

The new species differs from C. septentrionalis (southern China and northern Vietnam) by the mental not contacting the chin shields (vs contacting), five shields surrounding the paraparietal (vs six), slightly more subcaudals in males (20–23 vs 15–19), a smaller maximum total length in males (max TL 284 mm vs 344 mm), an obtusely pointed tail tip (vs broadly rounded), a blackish-brown dorsum (vs dark brown or black, usually with a narrow yellow ring behind the head), and unspotted pale khaki ventrals (vs yellow with small black spots).

Calamaria synergis sp. nov. differs from C. strigiventris (Lam Dong and Khanh Hoa provinces, Vietnam) by the mental not contacting the chin shields (vs contacting), five shields surrounding the paraparietal (vs 6), slightly fewer subcaudals in males (SC 20–23 vs 29–31), a smaller maximum total length in males (max TL 284 mm vs 362 mm), a shorter relative tail length in males (ratio TaL/TL 6.6–9.2% vs 11.2–17.9%), a non-tapering tail (vs tapering), an obtusely pointed tail tip (vs abruptly pointed), a blackish-brown dorsum (vs uniform grey-brown), and unspotted pale khaki ventrals (vs bright yellow with longitudinal black stripes).

Finally, the new species differs from C. yunnanensis (see Fig. 3C, Suppl. material 1: figs S1D, S2D) by the presence of a preocular scale (vs absent), fewer ventral scales in males (VEN 161–166 vs 167–184), eight maxillary teeth (vs 9), unspotted pale khaki ventrals (vs bright orange to orange-yellow), and the presence of pale neck rings (vs absent).

Natural history, distribution, and conservation status.

Calamaria synergis sp. nov. is currently known only from tropical evergreen forest at its type locality (see Fig. 5), at an elevation of ca 1,050 m asl. This region is characterised by high herpetofaunal diversity, with several snake species occurring sympatrically in the same habitat, including Oligodon fasciolatus (Günther), Lycodon fasciatus (Anderson), Plagiopholis nuchalis Boulenger, and Trimeresurus lanna Idiiatullina, Nguyen, Pawangkhanant, Suwannapoom, Chanhome, Mirza, David, Vogel & Poyarkov (see also Idiiatullina et al. 2024; Vogel et al. 2024). It seems highly likely that the new species also occurs in the adjacent mountainous areas of eastern Myanmar, northern Laos, and northeastern Vietnam. Due to the limited information available on the ecology and potential threats of Calamaria synergis sp. nov., we recommend that the new species be classified as Data Deficient (DD) according to the IUCN Red List Categories and Criteria (IUCN 2025).

Figure 5. 

Distribution ranges of Calamaria synergis sp. nov., C. andersoni, and C. yunnanensis. Numbers indicate localities where the species have been recorded (see Suppl. material 1: table S3 for locality details).

Discussion

The discovery of Calamaria synergis sp. nov. brings the total number of recognised Calamaria species in China to eight: C. andersoni, C. arcana, C. berezowskii, C. jinggangensis, C. pavimentata, C. septentrionalis, C. synergis sp. nov., and C. yunnanensis. Among these, six species – C. andersoni, C. arcana, C. berezowskii, C. jinggangensis, C. synergis sp. nov., and C. yunnanensis – are currently considered endemic to China. Notably, the type locality of Calamaria synergis sp. nov. lies in close proximity to the borders of Myanmar, Laos, and Vietnam, suggesting the potential for a broader distribution encompassing Shan State (Myanmar), Phongsaly Province (northern Laos), and the northwestern provinces of Vietnam, such as Dien Bien, Lai Chau, and Lao Cai. These transboundary montane forests may represent a previously overlooked biogeographic corridor for regionally endemic fossorial taxa, and future surveys in these areas will be essential to delimit the species true distribution.

In Yunnan Province, China, four species of Calamaria are currently recorded: C. andersoni, C. cf. pavimentata, C. yunnanensis, and C. synergis sp. nov. (Yang and Rao 2008; Yang and Zheng 2018; this study). Among them, C. synergis sp. nov. exhibits several morphological similarities with C. pavimentata, a widespread species reported from multiple localities in Indochina and southern China. Yang and Rao (2008) reported populations of C. pavimentata from Jingdong (specimens KIZ 75II0225 [adult female] and KIZ 75II0230 [juvenile]) and from Fugong, Nujiang (KIZ 78II0007, adult female). The Jingdong specimens differ from Calamaria synergis sp. nov. in relative tail length (6.6–9.2% in males of C. synergis sp. nov. vs 5.5% in Jingdong females), ventral scale counts (161–166 in males vs 180–182 in females), and subcaudal scale counts (20–23 in males vs 14 in females). However, these differences fall within the expected range of sexual dimorphism in Calamaria (see Table 3), and the close geographic proximity of Jingdong to the type locality of C. synergis sp. nov. (ca 140 km) suggests that specimens KIZ 75II0225 and KIZ 75II0230 may be conspecific with C. synergis sp. nov. In contrast, the specimen from Fugong (KIZ 78II0007), also previously assigned to C. pavimentata, exhibits a relative tail length of 5.7%, 174 ventral scales, and 15 subcaudal scales, traits inconsistent with those of any currently recognised species in Yunnan Province. This individual may represent an undescribed species from the extreme north of the province. These findings highlight the need for a systematic re-evaluation of historical museum specimens, especially those from taxonomically ambiguous populations, using both morphological and genetic data to refine species boundaries and distribution records.

More broadly, our results underscore that species of Calamaria remain among the least understood snake lineages in mainland Southeast Asia. Their cryptic appearance, secretive habits, and strongly fossorial lifestyle make them particularly difficult to detect during field surveys. As a result, many species are known from very limited material, often a single specimen or a small series (e.g., Orlov et al. 2010; Yang and Zheng 2018; Lee 2021). This sampling bias likely contributes to an underestimation of the genus true diversity, especially in ecologically diverse but poorly explored montane zones of southern China and the Indochinese Peninsula.

To advance taxonomic knowledge and conservation assessments for this elusive group, targeted surveys using appropriate methods (e.g., systematic leaf-litter sifting, pitfall traps, and microhabitat-based searches) should be prioritised in mid-elevation evergreen forests across border regions. Furthermore, integrative approaches combining detailed morphology, molecular phylogenetics, and ecological niche modelling will be key to resolving complex species groups and identifying overlooked lineages. The sister-group relationship recovered between Calamaria synergis sp. nov., C. andersoni, and C. yunnanensis also suggests the existence of a regionally restricted clade within montane southwestern China, possibly shaped by shared ecological constraints or historical barriers to gene flow. Additional phylogeographic studies may help elucidate the origin, divergence, and evolutionary trajectories of this lineage within the broader context of Calamaria diversification.

The following key to the species of Calamaria known from China is adapted from Yeung et al. (2022), Cai et al. (2023), and Liang et al. (2024). It is provided as a practical reference for preliminary identification in the field or collection. However, given the morphological similarity among species and the presence of cryptic diversity within the genus, accurate species determination should be confirmed through detailed morphological examination and, where possible, supported by molecular data.

1 Preocular present 2
Preocular absent Calamaria yunnanensis
2 Dorsal scales reduced to 4 rows on the tail at last the subcaudals 3
Dorsal scales reduced to >4 rows on the tail at last the subcaudals 6
3 Pale rings or blotches on neck present 4
Pale rings or blotches on neck absent Calamaria andersoni
4 Pale rings/blotches on tail present 5
Pale rings/blotches on tail absent Calamaria synergis sp. nov.
5 Tail tapering gradually to a point Calamaria pavimentata complex
Tail broadly rounded at tip (not tapering) Calamaria septentrionalis
6 Less than half of posterior chin shields meeting at midline; indistinct nuchal ring or paired spots present 7
Posterior chin shields meeting extensively at midline; collar or nuchal spots absent Calamaria jinggangensis
7 VEN 170–176 in males, VEN 192 in females Calamaria arcana
VEN 149–155 in males, VEN 153–171 in females Calamaria berezowskii

Acknowledgements

We thank Patrick David (MNHN, France) for kindly sharing data and literature relevant to Calamaria, and Tianyou Zhang and Fan Gao (China) for their assistance with specimen collection and for providing photographs of Calamaria yunnanensis. We are also grateful to Ngoc Quynh Nguyen and Duc Trong Nguyen (SIFASV, Vietnam) for their assistance in preparing the figures and map. Special thanks are extended to Robert Jadin (Lawrence University, USA) and Jeffrey L. Weinell (University of Kansas, USA) for their valuable comments and suggestions, which greatly improved the manuscript.

Additional information

Conflict of interest

The authors have declared that no competing interests exist.

Ethical statement

No ethical statement was reported.

Use of AI

No use of AI was reported.

Funding

This research was supported by the National Natural Science Foundation of China (NSFC 31471968), the Science and Technology Projects of Xizang Autonomous Region, China (XZ202301ZY0036G), the Rufford Foundation (Grant No. 45888-2; data analysis), and the Russian Science Foundation (RSF Grant No. 22-14-00037-P: data analysis).

Author contributions

All authors have contributed equally.

Author ORCIDs

Tierui Zhang https://orcid.org/0009-0000-1869-9313

Yuhao Xu https://orcid.org/0000-0001-6094-6680

Tan Van Nguyen https://orcid.org/0000-0001-5413-968X

Nikolay A. Poyarkov https://orcid.org/0000-0002-7576-2283

Gernot Vogel https://orcid.org/0000-0002-4542-518X

Xinge Wang https://orcid.org/0000-0002-8938-1377

Song Huang https://orcid.org/0000-0001-6786-8523

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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Supplementary material

Supplementary material 1 

Supplementary figures and tables

Tierui Zhang, Yuhao Xu, Tan Van Nguyen, Nikolay A. Poyarkov, Gernot Vogel, Xinge Wang, Song Huang

Data type: pdf

Explanation note: fig. S1. Comparative head shape and body coloration of Calamaria synergis sp. nov., C. andersoni, C. pavimentata, and C. yunnanensis. (A) C. synergis sp. nov. (ANU ZR24046); (B) C. andersoni (SYS r001699); (C) C. pavimentata (KFBG 14507); (D) C. yunnanensis (QHU R2024054). Each row shows: (1) dorsal view of head; (2) dorsal view of body; (3) dorsolateral view of body. Photographs by T.R. Zhang (A), reproduced from Yeung et al. (2022) (B, D), and Y.H. Xu (C). fig. S2. Comparison of tail shape and coloration of Calamaria synergis sp. nov., C. andersoni, C. pavimentata, and C. yunnanensis. (A) C. synergis sp. nov. (ANU ZR24046); (B) C. andersoni (SYS r001699); (C) C. pavimentata (KFBG 14507); (D) C. yunnanensis (QHU R2024054). Each row shows: (1) dorsal view of tail; (2) lateral view of tail; (3) ventral view of tail. Photographs by T.R. Zhang (A), reproduced from Yeung et al. (2022) (B, D), and Y.H. Xu (C). table S1. Museums, private collections and biorepository abbreviations mentioned in this study. table S2. List of specimens examined of Calamaria. table S3. List of localities of the Calamaria synergis sp. nov., C. andersoni, and C. yunnanensis appearing on Fig. 5.

This dataset is made available under the Open Database License (http://opendatacommons.org/licenses/odbl/1.0/). The Open Database License (ODbL) is a license agreement intended to allow users to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited.
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