Research Article |
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Corresponding author: Tan Van Nguyen ( tan.sifasv@gmail.com ) Corresponding author: Song Huang ( snakeman@ahnu.edu.cn ) Academic editor: Robert Jadin
© 2025 Tierui Zhang, Yuhao Xu, Tan Van Nguyen, Nikolay A. Poyarkov, Gernot Vogel, Xinge Wang, Song Huang.
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:
Zhang T, Xu Y, Nguyen TV, Poyarkov NA, Vogel G, Wang X, Huang S (2025) A new species of the genus Calamaria H. Boie in F. Boie, 1827 (Squamata, Calamariidae) from Xishuangbanna, Yunnan Province, China. ZooKeys 1253: 255-275. https://doi.org/10.3897/zookeys.1253.161412
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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.
Calamaria synergis sp. nov., morphology, Mountain Jinuo, phylogenetics, taxonomy
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 (
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
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 (
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’) (
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
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 |
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| C. andersoni | SYS r001699 | Yingjiang, Yunnan, China | MH445955 |
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| C. andersoni | HS R20101 | Dehong, Yunnan, China | OQ354844 |
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| C. andersoni | HS R20181 | Tengchong, Yunnan, China | OQ354845 |
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| C. andersoni | AHNU ZR25021 | Mangshi, Dehong, Yunnan, China | PV745123 | This study |
| C. arcana | KFBG 14611 | Mt. Dadongshan, Guangdong, China | ON482335 |
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| C. arcana | HS 17082 | Mt. Dawu, Guangdong, China | OQ354835 |
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| C. arcana | GP 9975 | Yongxing, Hunan, China | OP980549 |
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| C. arcana | DL R199 | Mt. Wuyi, Fujian, China | OQ354834 |
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| C. berezowskii | GXNU DLR194 | Mt. Gongga, Sichuan, China | PP747047 |
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| C. berezowskii | GXNU DLR195 | Mt. Gongga, Sichuan, China | PP747048 |
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| C. berezowskii | GXNU 20221215002 | Mt. Gongga, Sichuan, China | PP747049 |
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| C. gervaisii | KU 324661 | Puguis, La Trinidad, Benguet, Luzon, Philippines | MT819384 |
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| C. gervaisii | KU 334485 | Narvacan, Ilocos Sur, Luzon, Philippines | MT819385 |
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| C. jinggangensis | DL 20200725 | Mt. Jinggangshan, Jiangxi, China | OQ354830 |
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| C. jinggangensis | DL 20200625-2 | Mt. Jinggangshan, Jiangxi, China | OQ354831 |
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| C. jinggangensis | DL 20200625-3 | Mt. Jinggangshan, Jiangxi, China | OQ354832 |
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| C. jinggangensis | DL 20200625-4 | Mt. Jinggangshan, Jiangxi, China | OQ354833 |
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| C. lumbricoidea | KU 315159 | Pasonanca NP, Zamboanga, Philippines | MT819388 |
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| C. lumbricoidea | KU 334479 | Mt. Lumot, Gingoog, Misamis, Philippines | MT819389 |
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| C. cf. lumbricoidea | USMHC 1560 | Air Itam Dam, Penang, Malaysia | MN338526 |
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| C. muelleri | TNHC 58955 | Gowa, South Sulawesi, Indonesia | MT819390 |
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| C. muelleri | RMB 1283 | Gowa, South Sulawesi, Indonesia | MT819391 |
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| C. nebulosa | FMNH 258666 | Phongsaly, Laos | MN338524 |
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| C. palavanensis | KU 309445 | Barangay Irawan, Puerto Princessa, Palawan, Philippines | MT819386 |
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| C. palavanensis | KU 311411 | Mt. Mantalingahan, Rizal, Palawan, Philippines | MT819387 |
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| C. pavimentata | KFBG 14507 | Ningming, Guangxi, China | MH445957 |
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| C. schlegeli | LSUHC 10278 | Bukit Larut, Perak, Malaysia | MN338525 |
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| C. septentrionalis | FTB 2839 | unknown locality | KR814699 | Pyron unpublished data |
| C. septentrionalis | KFBG 14506 | Hainan, China | MH445956 |
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| C. septentrionalis | HS 11119 (CHS 116) | Tunxi, Huangshan, Anhui, China | MK201273 |
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| C. septentrionalis | HS 12055 (CHS 118) | Huangshan, Anhui, China | MK201274 |
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| C. septentrionalis | RE 30 (CHS 302) | Mangshan, Hunan, China | MK201384 |
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| C. septentrionalis | SYS r000932 (CHS 613) | Guangdong, China | MK201434 |
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| C. septentrionalis | HS R19100 | Mt. Huangshan, Anhui, China | OQ354842 |
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| C. septentrionalis | HS 11145 | Mt. Nanling, Guangdong, China | OQ354840 |
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| C. septentrionalis | DL 2021610-1 | Huangsha, Guangxi, China | OQ354838 |
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| C. cf. septentrionalis | ROM 35605 | Phia Oac-Phia Den NP, Cao Bang, Vietnam | AF471081 |
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| C. cf. septentrionalis | ROM 35597 | Phia Oac-Phia Den NP, Cao Bang, Vietnam | KX694890 |
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| C. yunnanensis | ROM 41547 | Simao, Yunnan, China | KX694891 |
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| C. yunnanensis | YPx 503 | Yunnan, China | JQ598922 |
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| C. yunnanensis | QHU R2024054 | Mt. Wanzhangshan, Simao, Yunnan, China | PV755783 | This study |
| Outgroup | ||||
| Orientocoluber spinalis | MVZ 211019 | Ningxia, China | AY486924 |
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| Elaphe quatuorlineata | LSUMZ 40626 | Hungary | AY486931 |
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| Lycodon rufozonatus | LSUMZ 44977 | China | AF471063 |
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Terminology and measurements follow
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
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
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.
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
| 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 |
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.
Holotype
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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.
(Fig.
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.
Maxillary teeth modified (enlarged), eight on each side (8/8).
(Fig.
(Fig.
Specimen
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).
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
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 |
|
| C. arcana | 4 | 2/3 | 1 | 0 | 6 | 170–176 | 192 | 20–22 | 12 | 303 | n/a | 7.2–11.8 | 4.7 | 0 | obtusely pointed | brownish | orange-red, unspotted |
|
| C. berezowskii | 4 | 2/3 | 1 | 0 | 6 | 149–155 | 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 |
|
| 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; |
| 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 |
|
| 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 |
|
| 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 |
|
| C. jinggangensis | 4 | 2/3 | 1 | 0 | 6 | 157–158 | 179 | 20 | 12 | 260 | 364 | 15 | 3.6 | 0 | obtusely pointed | brownish black | dark orange |
|
| 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 | 144–196 | 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 |
|
| 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 |
|
| 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 |
|
| 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 |
|
| C. schlegeli | 5 | 3/4 | 1 or 0 | 0 | 5 or 6 | 129–161 | 136–180 | 25–44 | 19–37 | 391 | 395 | 11.1–21.3 | 7.3–14.4 | 0 | blunt | unspotted grey or brown; head variably pink, yellow, and/or brown | cream, unspotted |
|
| C. septentrionalis | 4 | 2/3 | 1 | 0 | 6 | 148–166 | 168–188 | 15–19 | 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 ~6–8 scales behind the head | yellow, with small black spots |
|
| C. strigiventris | 4 | 2/3 | 1 | 0 | 6 | 130–168 | 176–183 | 29–31 | 20–22 | 362 | 367 | 11.2–17.9 | 8.4–0.6 | 2 | pointed | uniform grey-brown | bright yellow with longitudinal black stripes |
|
| 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 4–6 pale body bands | yellow, unspotted | Ziegler et al. 2005, |
| C. yunnanensis | 4 | 2/3 | 0 | 0 | 6 | 167–184 | 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 |
|
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.
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
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.
Calamaria synergis sp. nov. is currently known only from tropical evergreen forest at its type locality (see Fig.
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. (
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.,
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
| 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 |
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.
The authors have declared that no competing interests exist.
No ethical statement was reported.
No use of AI was reported.
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).
All authors have contributed equally.
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
All of the data that support the findings of this study are available in the main text or Supplementary Information.
Supplementary figures and tables
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. (