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Research Article
Cirrhimuraena taiwanensis sp. nov., a new species of cirri-bearing eel (Anguilliformes, Ophichthidae) from Yilan, northeastern Taiwan
expand article infoYen-Ting Lin‡, Yu-Hsiang Lin‡, Yu-San Han‡
‡ National Taiwan University, Taipei, Taiwan

Corresponding author: Yu-San Han ( yshan@ntu.edu.tw )

Academic editor: Yahui Zhao
© 2025 Yen-Ting Lin, Yu-Hsiang Lin, Yu-San Han.
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: Lin Y-T, Lin Y-H, Han Y-S (2025) Cirrhimuraena taiwanensis sp. nov., a new species of cirri-bearing eel (Anguilliformes, Ophichthidae) from Yilan, northeastern Taiwan. ZooKeys 1224: 129-140. https://doi.org/10.3897/zookeys.1224.141248
ZooBank: urn:lsid:zoobank.org:pub:60197A0C-E82B-486A-ACAE-A8698E6CCAAE
Open Access

Abstract

A new species of cirri-bearing eel, Cirrhimuraena taiwanensis sp. nov. (Anguilliformes, Ophichthidae), is described based on a specimen collected from the estuary of the Langyang River (Yilan County), northeastern Taiwan. The new species is distinct from all congeners, except C. odishaensis and C. orientalis, in possessing a single row of mandibular teeth. Cirrhimuraena taiwanensis sp. nov. differs from C. odishaensis in having significantly shorter pectoral fins and fewer vertebrae, and it is distinguished from C. orientalis by its larger head, notably more total vertebrae, and a dorsal fin that originates well behind the gill opening. In the neighbor-joining tree based on COI sequences, the new species forms a distinct monophyletic group; thus, it is clearly separable from congeners both morphologically and genetically. With this addition, there are now 13 species in the genus Cirrhimuraena.

Key words

Biodiversity, brackish water, COI analysis, Taiwanese fringe-lip eel, taxonomy

Introduction

The family Ophichthidae, commonly known as snake eels, represents the most varied group within the order Anguilliformes, containing two subfamilies (Myrophinae and Ophichthinae), with 62 genera and 361 species recorded (Fricke et al. 2024). In Taiwan, there have been reported of 19 genera and 60 species of snake eels identified (Chiu et al. 2018).

The genus Cirrhimuraena Kaup, 1856 belongs to the subfamily Ophichthinae and is known as cirri-bearing eels. This genus is notable for its unique morphological traits, particularly the presence of cirri, which are small, fleshy projections located on the upper jaw (Mohapatra et al. 2021). Cirri-bearing eels typically inhabit sandy or muddy substrates in coastal and estuaries waters (Mohapatra et al. 2021; Mohanty et al. 2023). Currently, 12 valid species are known, including Cirrhimuraena chinensis (Kaup, 1856), C. tapeinoptera (Bleeker, 1863), C. cheilopogon (Bleeker, 1860), C. calamus (Günther, 1870), C. playfairii (Günther, 1870), C. oliveri (Seale, 1910), C. paucidens (Herre & Myers, 1931), C. inhacae (Smith, 1962), C. orientalis (Nguyen, 1993), C. yuanding (Tang & Zhang, 2003), C. indica (Mohapatra, Mohanty, Ray, Mishra & Seth, 2021), and C. odishaensis (Mohanty, Behera, Patro & Mohapatra, 2023). Of these 12 species, only one, C. chinensis, has been recorded in Taiwan, and research on this genus remains scarce (Ho et al. 2015).

We conducted a survey of freshwater glass eels (juveniles of Anguilla spp.) in the the Langyang River estuary in northeastern Taiwan (24.7162°N, 121.8352°E) twice a month since 2010. Notably, this survey has yielded both a new species of Ophichthidae (Lamnostoma taiwanense Chiu, Huang & Shao, 2018) and new records of Anguilla borneensis Popta, 1924 and A. interioris Whitley 1938 (Chiu et al. 2018; Lin and Han 2024). In December 2023, a single Cirrhimuraena specimen was collected. A morphological analysis and molecular evidence indicated that this specimen represents an undescribed species. Although only a single specimen was obtained, its distinct morphology and genetic characteristics underscore its importance in advancing our understanding of Cirrhimuraena species in Taiwan. Furthermore, the discovery of new species and fish records highlights the need for conservation efforts to protect fish biodiversity in the Langyang River, one of the most critical habitats for Anguilliformes in Taiwan (Han et al. 2016).

Materials and methods

Sample collection

The specimen was collected from the estuary of the Langyang River in Yilan County, Taiwan (24.7162°N, 121.8352°E) on December 22, 2023. The environmental conditions of the collection site at the time of collection were as follows: substrate sandy, water depth 1 m, salinity 7‰, and water temperature 18 °C. A single, undescribed specimen of Cirrhimuraena was captured using a fyke net. Once collection, the specimen was photographed and radiographed, measured, and subsequently preserved in 95% ethanol. The specimen was deceased at the time of collection, and no live animals were included in this study.

Measurement and comparisons

The morphometrics were measured with digital calipers with an accuracy of 0.1 mm, and the meristic analysis and counting of head pores followed the protocol used by McCosker et al. (1989). The identified specimen was deposited in the collection of Biodiversity Research Museum of the Academia Sinica of Taiwan (ASIZP) under registration code ASIZP0082637. The specimen was compared with records of all congeners documented from Taiwan, Cirrhimuraena chinensis, and nearby waters, including C. yuanding from Pingtan, China (Tang and Zhang 2003), and C. playfairii from Okinawa, Japan (Hibino et al. 2021).

Molecular analysis

The dorsal muscle was dissected for the total genomic DNA extraction using the EasyPure Genomic DNA Spin Kit (Bioman Scientific, Taiwan). A polymerase chain reaction (PCR) was carried out to amplify the partial segment of the cytochrome c oxidase subunit I (COI) by using the forward primer FishF1+2 (5′-TCR ACY AAY CAY AAA GAY ATY GGC AC-3′) and the reverse primers FishR1 (5′-TAG ACT TCT GGG TGG CCA AAG AAT CA-3′) and FishR2 (5′-ACT TCA GGG TGA CCG AAG AAT CAG AA-3′) following the protocol adjusted from Chang et al. (2016). The final PCR product was sequenced using the primer FishF1+2 by Genomics Scientific, Taiwan.

The COI sequences were aligned and trimmed using BioEdit v. 7.7.1, resulting in partial sequences of 562 base pairs. Once aligned, the sequences were saved in FASTA format and imported into MEGA v. 11 (Tamura et al. 2021) for phylogenetic analysis. A neighbor-joining (NJ) tree was constructed using the Kimura 2-parameter (K2P) distance model, with 10,000 bootstrap replicates to assess the reliability of the branches. Among all congeners, only two valid species of Cirrhimuraena, C. chinensis and C. indica, were available in the NCBI GenBank. In total, 16 sequences were used to construct the NJ tree. In the ingroup, C. chinensis (GenBank numbers KY472820.1, KX215192.1, KX215193.1, KX215194.1, MK264639.1, MK264640.1, MK264641.1, GU674221.1, and GU674224.1), C. indica (GenBank number MT019886.1), and C. taiwanensis sp. nov. (GenBank number PQ558516.1) were included. In the outgroup, and following the study by Mohapatra et al. (2021) were Ophichthus lithinus Jordan & Richardson, 1908 (GenBank number KU94289.1), O. zophochir Jordan & Gilbert, 1882 (GenBank number GU440436.1), O. olivaceus McCosker & Bogorodsky, 2020 (GenBank number MN480448.1), and Pisodonophis cancrivorus Richardson, 1848 (GenBank number KU942788.1, and MK777102.1). The details of all COI sequences used are listed in Table 1.

Table 1.
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The detail of the sequences used in the phylogenetic analysis.

Species NCBI Accession number Source Voucher Number Sampling Locality
Cirrhimuraena taiwanensis sp. nov. PQ524198.1 This study ASIZP0082637 Yilan, Taiwan
Cirrhimuraena chinensis KY472820.1 GenBank PT011 China
Cirrhimuraena chinensis KX215192.1 GenBank JLJ050 China
Cirrhimuraena chinensis KX215193.1 GenBank JLJ051 China
Cirrhimuraena chinensis KX215194.1 GenBank JLJ052 China
Cirrhimuraena chinensis MK264639.1 GenBank PTD055 China
Cirrhimuraena chinensis MK264640.1 GenBank PT055 China
Cirrhimuraena chinensis MK264641.1 GenBank QZ053 China
Cirrhimuraena chinensis GU674221.1 GenBank BWA6863 Indonesia
Cirrhimuraena chinensis GU674224.1 GenBank BWA6862 Indonesia
Cirrhimuraena indica MT019886.1 GenBank EBRC/ZSI/11811 India
Ophichthus lithinus KU942789.1 GenBank ASIZP0801626 Taiwan
Ophichthus olivaceus MN480448.1 GenBank KAU17-80 Saudi Arabia
Ophichthus zophochir GU440436.1 GenBank MFC132 California, USA
Pisodonophis cancrivorus MK777102.1 GenBank DOS05154 Vietnam
Pisodonophis cancrivorus KU942788.1 GenBank ASIZP0800053 Taiwan

Results

Family Ophichthidae

Cirrhimuraena taiwanensis sp. nov.

https://zoobank.org/3214769E-E179-46DD-B800-CE12D2A318B0
Figs 1, 2, 3, 4, Table 2

Material examined

Holotype : Taiwan • ASIZP0082637, 178.1 mm total length (TL); Yilan; 24.7162°N, 121.8352°E; 22 Dec. 2023; caught by fyke net, ca 1 m, Yu-San Han & Yen-Ting Lin leg.

Diagnosis

A new Cirrhimuraena species with the combination of following characteristics: pectoral fin very small, only 15.2% of head length (HL) (in congeners > 21% HL); HK 9.7% of TL dorsal fin originates 1½ pectoral-fin length behind gill opening; tooth pattern unique, with only a single row of mandibular teeth; cirri on upper jaw 11; vertebrae 150, vertebral formula 13-53-150.

Figure 1. 

Cirrhimuraena taiwanensis sp. nov., ASIZP0082637, 178.1 mm total length. Scale bar: 30 mm.

Description

The morphometric and meristic measurements of the holotype are shown in Table 2. Body very elongate, cylindrical; body height is almost consistent from gill opening to anus, with depth at gill opening of 2.2% of TL and depth at anus of 2.3% of TL. Head moderate, with head length (HL) 9.7% of TL. Tail longer than trunk, 63.6% of TL. Anal fin low, situated right after anus, with pre-anal length 36.8% of TL. Dorsal fin also low, originating far behind gill opening and pectoral fin; pre-dorsal length 13.2% of TL. Pectoral fin very small, 15.2% of HL, 1.5% of TL; pectoral-fin base positioned at same vertical as gill opening; gill opening positioned on latero-ventral side, length 16.4% of HL.

Figure 2. 

Head profile of Cirrhimuraena taiwanensis sp. nov. A origin of dorsal fin well behind gill opening B arrow indicates cirri on upper jaw.

Table 2.
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Morphometric and meristic data of Cirrhimuraena taiwanensis sp. nov.

Cirrhimuraena taiwanensis sp. nov. Holotype, ASIZP0082637
Total length (SL, mm) 178.1
Head length (HL, mm) 17.4
Pre-anal length (PAL, mm) 65.6
Pre-dorsal length (PDL, mm) 23.6
% in HL
Snout length 19.1
Eye diameter 8.9
Interorbital length 6.1
Upper jaw length 35.7
Lower jaw length 26.9
Gill opening length 16.4
Pectoral-fin length 15.2
% in TL
Head length 9.7
Pre-anal length 36.8
Pre-dorsal length 13.2
Trunk length 27.1
Tail length 63.6
Depth at gill opening 2.2
Depth at anus 2.3
Pores
Supraorbital 1 + 3
Infraorbital 3 + 2
Preoperculomandibular 7 + 4
Pores before pectoral fin 11
Pores before dorsal fin 16
Pores before anus 48
Vertebrae
Pre-dorsal 13
Pre-anal 53
Total 150

Eye relatively large, positioned nearer to snout tip than rictus; eye diameter 8.9% of HL; interorbital space slightly wider; interorbital length 6.1% of HL. Anterior nostril tubular, positioned at snout tip, while posterior nostril lies slightly behind orbit. Snout long, pointed, 19.1% of HL. Upper jaw longer than lower jaw, 35.7% and 26.9% of HL, respectively.

Five small cirri on edge of upper jaw between anterior and posterior nostrils; 6 cirri behind posterior nostril. No cirri on lower jaw and tip of jaw in front of nostrils. Dentition pattern illustrated in Fig. 3. Teeth numerous, closely arranged in a band, and primarily small and pointed, with slightly larger teeth at ends. Vomerine teeth in 1–3 rows, extending to posterior of maxilla; 5 teeth form a small patch at prevomer. Maxillary teeth in 2–6 rows of small, conical teeth; mandibular teeth band in only a single row on both sides. Pre-dorsal vertebrae 13, pre-anal vertebrae 53, and total vertebral 150.

Figure 3. 

Head and lateral line pores in Cirrhimuraena taiwanensis sp. nov. Abbreviaitons: IO: infraorbital pores; LL: lateral-line pores; POM: preoperculomandibular pores; SO: supraorbital pores; ST: supra-temporal pores.

Head pores tiny and indistinct, with supraorbital pores 1 + 3, infraorbital pores 3 + 2, preoperculomandibular pores 7 + 4, and supra-temporal pores 1 (Fig. 4). Lateral line pores before pectoral fin/gill opening 12, before dorsal-fin origin 16, and before anus 48.

Figure 4. 

Tooth dentition pattern in upper and lower jaws of Cirrhimuraena taiwanensis sp. nov. (holotype, ASIZP0082637, 178.1 mm total length).

Dorsal surface of body grayish, with numerous tiny black spots; some melanophores concentrated at tip of snout. Ventral side whitish. Dorsal and anal fins translucent; pectoral fin whitish.

Distribution

Currently only known from the type locality, with sandy substrate.

Etymology

The specific epithet taiwanensis refers to the location of the type locality, which recently only known in Taiwan; it is used as an adjective.

Remarks

Compared to all 12 congeners, C. taiwanensis sp. nov. can be easily distinguished from 10 species, except C. odishaensis and C. orientalis, in having only a single row of mandibular teeth (Fig. 4). However, C. taiwanensis sp. nov. can be separated from these two species morphologically, with comparative details shown in Table 3. The new species differs from C. odishaensis in having a shorter pectoral fin, only 15.2% of HL (compared to 21.3–25.0% HL in C. odishaensis); fewer vertebrae, with 13 pre-dorsal, 53 pre-anal, and 150 total vertebrae (vs 10 pre-dorsal, 46–47 pre-anal, and 160–162 total vertebrae in C. odishaensis); and fewer rows of maxillary teeth (2–6 rows in C. taiwanensis sp. nov. vs 3–7 rows in C. odishaensis). Compared to C. orientalis, C. taiwanensis sp. nov. has a larger head at 9.7% of TL (vs 5.5–6.2% of TL in C. orientalis), significantly more vertebrae (150 vs 131–136 in C. orientalis), and more rows of maxillary teeth (2–6 rows in C. taiwanensis sp. nov. vs 2–3 rows in C. orientalis).

Table 3.
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Morphometric comparisons of Cirrhimuraena taiwanensis sp. nov. with congeners with only a single row of mandibular teeth.

Cirrhimuraena taiwanensis sp. nov. (This study) C. odishaensis (Mohanty et al., 2023) C. orientalis (Nguyen., 1993)
HL % in TL 9.7 9.1–10.6 5.5–6.2
Pectoral fin % in HL 15.2 21.3–25.0 —
Mandibular teeth 1 row 1 row 1 row
Maxillary teeth 2–6 rows 3–7 rows 2–3 rows
Total vertebrate 150 160–162 131–136

— No data available.

Molecular results

Sixteen COI sequences from three taxa were analyzed, revealing nine unique haplotypes across 562 aligned base pairs, which included 196 variable sites and 151 parsimony-informative sites. The NJ tree analysis identified C. taiwanensis sp. nov. in a well-supported clade (bootstrap values 99%) with all other Cirrhimuraena species included in NCBI (Fig. 5). The average pairwise K2P genetic distance between C. taiwanensis sp. nov. and its congeners is 0.124, aligning with the average genetic distance typically found among congeneric fish species, as reported by Ward et al. (2005). Within the Cirrhimuraena group, C. taiwanensis sp. nov. and most C. chinensis specimens are clearly separated (bootstrap value 99%) from C. indica and two C. chinensis specimens collected in Indonesia (GU674221.1 and GU674224.1), which are suspected misidentifications of C. indica (Mohapatra et al. 2021). Cirrhimuraena taiwanensis sp. nov. also demonstrates a distinct separation from C. chinensis, with high bootstrap support of 82% and forming a unique monophyletic group. The distinct morphological characteristics and NJ tree results further support the separation of C. taiwanensis sp. nov. as a distinct species.

Figure 5. 

The neighbor-joining tree based on COI sequences of Cirrhimuraena taiwanensis sp. nov. and all the valid congeners in NCBI.

Discussion

Currently, there are 12 valid species in the genus Cirrhimuraena, and the distribution in the northwestern Pacific Ocean is primarily centered around the South China and Java Seas (Mohapatra et al. 2021; Mohanty et al. 2023). Only one species, C. chinensis, has been recorded from Taiwanese waters, from along the coast of Pingtung in southwestern Taiwan and Kinmen Island (Shao et al. 2008; Ho et al. 2015). Additional to C. chinensis recorded in Taiwanese waters, in the subtropical North Pacific there are two additional species: C. playfairii, recorded from Makiya, Okinawa Island, Japan (Hibino et al. 2021: fig. 6b, c), and C. yuanding, recorded from Pingtan County, Fujian Province, China (Tang and Zhang 2003) (Fig. 6).

Figure 6. 

Distribution map of Cirrhimuraena species found in Taiwanese waters and nearby regions.

In Table 4, we compare the new species, which exhibits distinct morphological differences from congeners found in Taiwanese waters (C. chinensis) and nearby regions (C. yuanding and C. playfairii). The two species recorded from China (C. yuanding) and Japan (C. playfairii) can be clearly distinguished from C. taiwanensis sp. nov. by the position of the dorsal fin, which originates in front of the gill opening in both C. playfairii and C. yuanding, with a pre-dorsal length (PDL) shorter than the HL (Table 4). Compared to the C. chinensis, C. taiwanensis sp. nov. has a significantly shorter pectoral fin at 15.2% of HL (vs 45.2–51.6% HL in C. chinensis); a smaller gill opening length at 16.4% of HL (vs 25.4–30.1% HL in C. chinensis); dorsal fin that originates well behind the gill opening, with a PDL of 13.2% TL (vs 9.6–11.2% TL in C. chinensis); and a slightly smaller head, at 9.7% TL (vs 10.9–11.8% TL in C. chinensis) (Table 4). Furthermore, molecular data confirm the distinction between C. chinensis and C. taiwanensis sp. nov., with a high bootstrap value (82%) supporting their separation (Fig. 5).

Table 4.
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Comparisons of Cirrhimuraena taiwanensis sp. nov. and C. chinensis in Taiwan, and two other congeners recorded from nearby waters.

Cirrhimuraena taiwanensis sp. nov. Holotype C. chinensis n = 10 C. yuanding n = 1 (Tang and Zhang 2003) C. playfairii n = 1 (Hibino et al. 2021)
Collection site Taiwan Taiwan China Japan (Okinawa)
Total length (SL, mm) 178.1 227–293 520.5 229
Head length (HL, mm) 17.4 23.5–28.9 30.0 —
Pre-anal length (PAL, mm) 65.6 89.3–94.2 161.3 —
Pre-dorsal length (PDL, mm) 23.6 25.5–28.2 20.5 —
% in HL
Snout length 19.1 18.9–22.7 16.0 15.3
Upper jaw length 35.7 37.1–40.0 24.0 31.3
Lower jaw length 26.9 34.5–43.5 — —
Gill opening length 16.4 25.4–30.1 12.7 8.6
Pectoral-fin length 15.2 45.2–51.6 28.3 23.3
% in TL
Head length 9.7 10.9–11.8 5.8 7.1
Pre-anal length 36.8 32.1–39.4 31.0 33.6
Pre-dorsal length 13.2 9.6–11.2 3.9 4.6
Depth at gill opening 2.2 2.5–3.3 1.8 2.0
Depth at anus 2.3 2.8–3.7 2.2 2.1
Vertebrate
Pre-dorsal 13 11 — 4
Pre-anal 53 49 — 60
Total 150 154 — 183

— No data available.

There are also notable morphological differences between Cirrhimuraena taiwanensis sp. nov. and other Indo-West-Pacific congeners. Cirrhimuraena calamus and C. oliveri both have significantly smaller heads, measuring 16.6% pre-anal length (PAL) in C. calamus (Günther 1870) and 16.4% of PAL in C. oliveri (Seale 1910), compared to 26.5% PAL in C. taiwanensis sp. nov.; C. tapeinoptera, C. cheilopogon, and C. inhacae have significantly larger pectoral fins, approximately 40–50% HL (Weber and de Beaufort 1916; Smith 1962; Smith and Heemstra 1986) vs 15.2% in C. taiwanensis sp. nov.; C. taiwanensis sp. nov. also has a shorter pre-anal length, at 36.8% TL, compared to 41.8% in C. paucidens (Catania and Fong 2020; Mohapatra et al. 2021). The combined morphological and molecular differences between the new species and all 12 congeners strongly support that the specimen we collected represents a distinct new species, Cirrhimuraena taiwanensis sp. nov.

The habitat of the Cirrhimuraena taiwanensis sp. nov. is at the estuary of the Langyang River, where the water is brackish year round and has an abundance plankton. The river estuary serves as an important habitat for the Anguilliformes and other brackish and freshwater fish species (Shih et al. 2008; Dahms et al. 2012; Han et al. 2016). The substrate is sandy, and the brackish-water environment is typical habitat for the Cirrhimuraena species (Mohanty et al. 2023). With the description of the new species, the ecological importance of the Langyang River estuary is enhanced; this estuary already serves as the type locality of Lamnostoma taiwanensis and habitat for other anguillid species in Taiwan (Chiu et al. 2018; Lin and Han 2024). In addition, the these new and recently described species suggests that many more unidentified species may be present in brackish waters, which highlight the importance of these environments for biodiversity.

Acknowledgements

We thank Zhen-Hui Chen (local fisherman) for assisting us in setting the fyke net and providing various sampling support, Chien-Hsiang Lin and Hsin-Wei Liu (Biodiversity Research Center, Academia Sinica) for photographing specimen and offering curatorial insights for this research, and Hong-Ji Veterinary Hospital for helping taking the radiograph for the specimen. We sincerely thank the academic editor, Dr. Yahui Zhao, two anonymous reviewers, and Dr. Mohapatra for their invaluable comments and suggestions to improve the quality of this manuscript.

Additional information

Conflict of interest

The authors have declared that no competing interests exist.

Ethical statement

No ethical statement was reported.

Funding

The authors extend our gratitude to the National Science and Technology Council, Executive Yuan, Taiwan (MOST 111-2313-B-002-016-MY3) for funding this project.

Author contributions

The sample collecting was performed by Yu-San Han and Yen-Ting Lin. Yen-Ting Lin conducted the measurement and write the manuscript; Yu-Hsiang Lin for the head profile and dentition drafting; Yu-San Han designed and supervised the experiments. All authors participated in manuscript writing and interpretation of results. All authors read and approved the final manuscript.

Data availability

All data that support the findings of this study are available in the main text, and the holotype is deposited in the collection of Biodiversity Research Museum of the Academia Sinica of Taiwan, under registration code ASIZP0082637.

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