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. 2025 Sep 23;1253:73–92. doi: 10.3897/zookeys.1253.153155

A new typical cavefish of the genus Triplophysa (Teleostei, Cypriniformes, Nemacheilidae) from the Jinsha River, Yunnan, China

Min Shi 1,2, Yuan-Chao Chen 1, Xing-Jin Che 1,2, Wei Dao 1,2, Wen-Ming Liu 3, Deng-Shan Wang 3, Jun-Xing Yang 1, Xiao-Ai Wang 1,✉
PMCID: PMC12485481  PMID: 41041658

Abstract

Triplophysa baishuijiangensissp. nov., a new cave-dwelling loach, was collected from an underground river outlet in Xiaoganxi Village, Yiliang County, Yunnan Province, China, situated within the core area of the Baishuijiang National Aquatic Germplasm Resources Reserve for Endemic Fish, part of the Hengjiang–Jinsha River system. The new species is distinguished from congeners by significant genetic divergence and the following combination of characteristics: vestigial eyes, absence of skin pigmentation, pelvic fin tip reaching the anus, complete lateral line, developed posterior chamber of the swim bladder, and dorsal fin rays iii-7. Phylogenetic analysis based on Cyt b positions this new species in the hypogean group of Triplophysa, increasing the documented cave species in the genus from 42 to 43. Triplophysa baishuijiangensissp. nov. represents the first documented typical cavefish in the Jinsha River basin, underscoring the region’s ecological significance for subterranean adaptation and suggesting unexplored cave biodiversity that merits systematic investigation.

Key words: Cavefish, Cyt b sequence, loach, molecular phylogeny, new species, Triplophysa

Introduction

Triplophysa Rendahl, 1933, the most species-rich genus in the family Nemacheilidae, comprises approximately 166 valid species, with 78% (130 species) distributed in China (Fricke et al. 2025). These species are primarily distributed across the Qinghai-Tibetan Plateau and adjacent regions but are also found in southwestern and northern China (Zhu 1989; Chen et al. 2009; Tang et al. 2021). Morphological characteristics distinguishing Triplophysa from other genera in the Nemacheilidae include: anterior and posterior nostrils closely set, a posterior wall of the bony swim bladder capsule, body cylindrical anteriorly, progressively compressed posteriorly, and sexual dimorphism (tubercles and thickened pads on the snout, cheeks, and dorsal surfaces of pectoral fins) (Zhu 1989). Within Triplophysa, sexual dimorphism presents various manifestations, Hou et al. (2010) categorized these into 12 distinct types, notably, of which one demonstrates sexual dimorphism in head and snout shape: males possess a squarish snout (in dorsal view), while females exhibit a rounded snout.

Triplophysa species are classified into hypogean (cave-dwelling) and epigean (surface-dwelling) groups based on ecological habits (Liu et al. 2022; Lu et al. 2022). Recent taxonomic studies have confirmed the monophyly of these two groups, with the epigean group occupying a basal position within Triplophysa (Yan 2018; Chen et al. 2019; Liu et al. 2022; Luo et al. 2023a; Lan et al. 2024; Cao et al. 2025). Among the hypogean group, varying degrees of ocular and pigment degeneration are observed, ranging from atypical cavefish to fully troglomorphic forms (Yan 2018; Liu et al. 2022). Typical cavefish exhibit advanced troglomorphic adaptations, characterized by regressed or completely degenerated eyes, translucent bodies, and well-developed sensory barbels (Li 2018).

Another Nemacheilid genus, Claea Kottelat, 2011, is morphologically similar to Triplophysa, with frequent confusion between the two genera (Kottelat, 2011; Chen et al. 2021; Zhang et al. 2024). Recent phylogenies confirm their close affinity: Claea forms a sister clade to the Triplophysa hypogean group and is embedded within Triplophysa (Yan 2017; Zhang et al. 2024; Lei et al. 2025). Claea species show a preference for cave habitats (Liao et al. 1997; Chen et al. 2021), with one troglomorphic species (C. scet Lei, He, Huang, Zhou & He, 2025) described. Thus, including Claea in phylogenetic analyses of putative new Triplophysa cave species is essential to avoid misidentification.

The karst region of southwestern China (Yunnan, Guizhou, Guangxi, and surrounding areas) is endowed with abundant river systems such as such as the Wujiang (a tributary of the Yangtze River), Nanpanjiang, Beipanjiang, and Hongshui River. Its diverse karst landforms (Liu et al. 2008) provide a favorable environment for the evolution of cavefish (Yan 2018; Luo et al. 2023b), and this area also serves as a biodiversity hotspot for cave-dwelling loaches, with 83 species of cave-adapted Nemacheilidae fishes currently documented (Luo et al. 2023c). Among these, the genus Triplophysa represents the largest group, comprising 43 species (Lan et al. 2024; Cao et al. 2025; Suppl. material 1).

Currently, six cave-dwelling Triplophysa species have been described from the Yangtze River basin, with their distributions restricted to the Wujiang River and Yuanjiang River systems. These include T. qingzhenensis Liu, Zeng & Gong, 2022, T. wudangensis Liu, Zeng & Gong, 2022, T. rosa Chen & Yang, 2005, and T. qini Deng, Wang & Zhang, 2022 from the Wujiang River, with T. xiangxiensis (Yang, Yuan & Liao, 1986) and T. erythraea Liu & Huang, 2019 from the Yuanjiang River.

In 2023, 11 cave loach specimens were collected from a karst cave in Xiaoganxi Village, Yiliang County, Yunnan Province, China. This cave is hydrologically connected to the Xiaoganxi stream, which drains into the Baishuijiang River (Jinsha River basin). Based on these specimens exhibiting: closely set anterior and posterior nostrils, cylindrical anterior body progressively compressed posteriorly, and presence of sexual dimorphism, we assign them to Triplophysa. Molecular phylogenetic analyses and morphological comparisons demonstrated that these specimens represent a distinct species within the genus Triplophysa. Here, we formally describe it as Triplophysa baishuijiangensis sp. nov.

Materials and methods

Specimens were collected using dip nets at a subterranean river outlet in the Baishuijiang Reserve. Samples were either fixed in 75% ethanol or transported live to the laboratory. After live photos were taken, live samples were fixed in 10% formalin. Before fixation, the right pelvic fin rays were clipped from all samples for molecular analysis and preserved in anhydrous ethanol. All specimens are preserved at the Kunming Institute of Zoology, Chinese Academy of Sciences, Kunming, Yunnan Province, China (KIZ).

Morphological examination. Ten specimens were measured with digital calipers, all measurable parameters were recorded as point-to-point distances to an accuracy of 0.01 mm, following the methodology of Li (2018) and Wang (2022). Sex differentiation is based on head shape, referring to the sexual dimorphism description of T. daqiaoensis Ding, 1993 in Hou et al. (2010). The count of vertebrae was taken from X-ray photographs (Digital Cabinet X-ray System Xpeart 80, Kubtec Scientific, Stratford, USA). One non-type specimen was dissected to observe and describe the morphology of the intestine and swim bladder, and to count the number of inner gill rakers on the first gill arch.

Comparative morphological data were acquired from specimen examination and literature (Suppl. material 2), type specimens of 14 cave-dwelling Triplophysa species were examined in this study, including: T. shilinensis Chen & Yang, 1992, T. anshuiensis Wu, Wei, Lan & Du, 2018, T. gejiuensis (Chu & Chen, 1979), T. huapingensis Zheng, Yang & Chen, 2012, T. nandanensis Lan, Yang & Chen, 1995, T. longliensis Ren, Yang & Chen, 2012, T. guizhouensis Wu, He, Yang & Du, 2018, T. rosa, T. tianlinensis Li, Li, Lan & Du, 2016, T. tianeensis Chen, Cui & Yang, 2004, T. macrocephala Yang, Wu & Yang, 2012, T. luochengensis Li, Lan, Chen & Du, 2017, T. longipectoralis Zheng, Du, Chen & Yang, 2009, and T. xichouensis Liu, Pan, Yang & Chen, 2017.

Phylogenetic analyses. Genomic DNA was extracted from pelvic fin tissues of four specimens using the Ezup column animal genome DNA extraction kit according to the manufacturer’s protocol. Primers L14724 (5’-GACTTGAAAAACCACCGTTG-3’) and H15915 (5’-CTCCGATCTCCGGATTACAAGAC-3’) were selected for PCR amplification of the mitochondrial cytochrome b (Cyt b) (Xiao et al. 2001). The amplification conditions were as follows: initial denaturation at 95 °C for 5 min, followed by 35 cycles of 94 °C for 1 min (denaturation), 48 °C for 1 min (annealing), and 72 °C for 1 min (extension), with a final extension at 72 °C for 10 min. The PCR products were sent to Sangon Biotech for sequencing.

A total of 43 Cyt b sequences were included in the molecular analysis, comprising four newly sequenced specimens and 39 sequences downloaded from GenBank (Table 1), with Homatula tigris Che, Dao, Chen, Pan, Hua, Liang & Wang, 2023 and H. laxiclathra Gu & Zhang, 2012 designated as outgroups. Cyt b sequences were aligned using the Clustal W algorithm in MEGA v.11.0 (Tamura et al. 2021), followed by manual trimming of the alignment. Genetic distances were calculated using the Kimura-2-parameter model in MEGA v.11.0. Phylogenetic trees were constructed with two methods: Maximum Likelihood (ML) analysis was conducted in IQ-TREE v. 2.1.4 (Minh et al. 2020) with 10,000 ultrafast bootstrap replicates (Hoang et al. 2018), model selection via ModelFinder identified GTR+F+I+G4 as the optimal evolutionary model based on BIC optimization (Kalyaanamoorthy et al. 2017). Bayesian Inference (BI) was performed using MrBayes v. 3.2.7 (Ronquist et al. 2012) with substitution models optimized through PartitionFinder v. 2.1.1 (Lanfear et al. 2017) under the Bayesian Information Criterion, the Cyt b gene was partitioned by codon position, with best-fit models assigned as: first codon: TRNEF+I+G, second codon: HKY+I, third codon: TIM+I+G. Two independent runs were executed for 20 million generations each, sampling trees every 1,000 generations. After discarding the initial 25% of samples as burn-in.

Table 1.

GenBank accession numbers for species used in the molecular phylogenetic analysis.

Species Localities Voucher ID Accession number Reference
T. anlongensis Xinglong Town, Anlong County, Guzihou, China GZNU20230112002 OQ754139.1 Luo et al. 2023
T. baishuijiangensis 1 Niujie Town, Yiliang County, Yunnan, China KIZ2025000165 PQ199313 This study
T. baishuijiangensis 2 Niujie Town, Yiliang County, Yunnan, China KIZ2025000166 PQ199314 This study
T. baishuijiangensis 3 Niujie Town, Yiliang County, Yunnan, China KIZ2025000175 PQ199315 This study
T. baotianensis Baotian Town, Panzhou City, Guzihou, China GZNU20180421005 NC_056365.1 Wang et al. 2021
T. cehengensis Rongbei Town, Ceheng County, Guzihou, China GZNU20230109001 OQ754132.1 Luo et al. 2023
T. erythraea Dalong Cave, Huayuan County, Hunan, China HY18011301 MG967615.1 Unpublished
T. fengshanensis Fengshan county, Guangxi, China SWU20160518002 OQ998929.1 Zhang et al. 2024
T. guizhouensis Lewang Town, Wangmo County, Guzihou, China GZNU20220313001 OQ241174.1 Luo et al. 2023
T. huapingensis Huaping Town, Leye County, Guangxi, China GZNU20230404004 OQ754125.1 Luo et al. 2023
T. langpingensis Longping Township, Tianlin County, Guangxi GZNU20230404001 OQ754122.1 Luo et al. 2023
T. longipectoralis Huanjiang county, Guangxi, China SWU20161110005 OQ998928.1 Zhang et al. 2024
T. longliensis Longli county, Guizhou, China SWU20160903003 MW582825.1 Cheng et al. 2021
T. macrocephala Lihu Town, Nandan County, Guangxi, China GZNU20230404002 OQ754123.1 Luo et al. 2023
T. nandanensis Liuzhai Town, Nandan County, Guangxi, China GZNU20230404007 OQ754128.1 Luo et al. 2023
T. nanpanjiangensis China KIZ20080361 MG238302.1 Min et al. 2023
T. nasobarbatula Dongtang Township, Libo County, Guizhou, China GZNU20220313011 OQ241176.1 Luo et al. 2023
T. orientalis Tibet, China H234 MK655279.1 Unpublished
T. panzhouensis Hongguo Town, Panzhou City, Guizhou, China GZNU20220513001 OQ754119.1 Luo et al. 2023
T. pappenheimi Gansu province, China CF2481hh KX373843.1 Unpublished
T. qingzhenensis Qingzhen County, Guiyang City, Guizhou, China IHB201911150004 MT700458.1 Liu et al. 2023
T. qini Houping Village, Wulong County, Chongqing, China WNHM10234 ON528185.1 Deng et al. 2022
T. qiubeiensis Nijiao Village, Qiubei County, Yunnan, China GZNU20230404006 OQ754127.1 Luo et al. 2023
T. robusta Tibet, China H30 OP616095.1 Unpublished
T. rongduensis Rongdu Town, Ceheng County, Guizhou, China GZNU20230110001 OQ754135.1 Luo et al. 2023
T. rosa HuoLuTown, Wulong County, Chongqing, China T21 OQ754130.1 Luo et al. 2023
T. sanduensis Zhonghe Town, Sandu County, Guizhou, China SWU20170613001 MW582822.1 Cheng et al. 2021
T. stolickai China H9 OP616074.1 Unpublished
T. tianeensis Bala Township, Tian ‘e County, Guangxi, China GZNU20230404003 OQ754124.1 Luo et al. 2023
T. wenshanensis Dehou Town,Wenshan County, Yunnan, China JWS2023003 PP661513 Cao et al. 2025
T. wudangensis Wudang District, Guiyang City, Guizhou, China IHB2019080904 MT700460.1 Liu et al. 2022
T. xiangxiensis Feihu Cave, Hunan Province / JN696407.1 Yao et al. 2012
T. xuanweiensis Tangtang Town, Xuanwei City, Yunnan, China ASIZB223820 OL675198.1 Lu et al. 2022
T. yaluwang Maoying Town, Ziyun City, Guizhou, China GZNU20240118005 PQ117067 Lan et al. 2024
T. yangi Wulong Town, Shizong County, Yunnan, China / PQ356185.2 Cao et al. 2025
T. zhenfengensis Xinlongchang Town, Xingren City, Guizhou, China GZNU20220313005 OQ241180.1 Luo et al. 2023
T. ziyunensis Maoying Town, Ziyun City, Guizhou, China GZNU20230529003 PQ117069 Lan et al. 2024
Claea wulongensis Wulong County, Chongqing, China SWU2019051308 MW582823.1 Cheng et al. 2021
C. dabryi 1 Niujie Town, Yiliang County, Yunnan, China KIZ20230516001 PQ199316 This study
C. dabryi 2 Kanding City, Sichuan, China CWNU201407121 KX289615.1 Unpublished
C. dabryi 3 Weixi County, Yunnan, China KIZ2009003611 MG238216.1 Min et al. 2023
Homatula tigris Panlong District, Kunming City, Yunnan, China KIZ202100004147 ON124934.1 Che et al. 2023
H. laxiclathra Jiangkou Town, Ningshan County, Shanxi, China KIZ2005014383 MG238220.1 Min et al. 2023
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Results

Phylogenetic analyses and genetic divergence

A total of 43 aligned Cyt b gene sequences were used for phylogenetic analysis and genetic distance calculation. The BI and ML phylogenetic trees showed a highly consistent topology, species in phylogenetic tree were classified into three distinct groups: hypogean group, epigean group, and trogloxene group (Fig. 1). The hypogean group comprises 31 cave-dwelling species of Triplophysa, while the trogloxene group includes C. dabryi (Sauvage, 1874) and C. wulongensis (Chen, Sheraliev, Shu & Peng, 2021). Phylogenetic analyses support a sister-group relationship between Claea and hypogean Triplophysa lineages. The epigean group comprises four surface-dwelling Triplophysa species: T. stolickai (Steindachner, 1866), T. orientalis (Herzenstein, 1888), T. robusta (Kessler, 1876), and T. pappenheimi (Fang, 1935). The hypogean group is further divided into two clades: Clade II includes 14 species primarily distributed in the upper reaches of the Pearl River (except for T. baishuijiangensis sp. nov. in the upper Yangtze River), while Clade I consists of 17 species occurring in the relatively lower reaches of the Yangtze and the Pearl River (Figs 1, 5). Notably, the three samples of T. baishuijiangensis sp. nov. form a monophyletic clade with a bootstrap value of 100 and are sister to T. xuanweiensis Lu, Li, Mao & Zhao, 2022 within Clade II.

Figure 1.

Figure 1.

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Phylogenetic tree based on 43 Cyt b sequences, Bayesian posterior probabilities (BPP) from BI analysis, and ultrafast bootstrap supports (UBP) from ML analysis are shown at nodes.

Figure 5.

Figure 5.

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Distribution map of 43 cave-dwelling Triplophysa species. Symbols denote species distribution: triangle (Yangtze River), circle (Pearl River), square (Red River). Colors represent phylogenetic clades: red (Clade I), black (Clade II), grey (lack of Cyt b data).

Pairwise genetic distances (Suppl. material 3) between the new species and its congeners within the hypogean group ranged from 10.00% (T. xuanweiensis) to 17.42% (T. anlongensis Lan, Song, Luo, Zhao, Xiao & Zhou, 2023). Within the hypogean group, the maximum pairwise genetic distance was 19.70% (between T. fengshanensis Lan, 2013 and T. anlongensis), while the minimum was 0.35% (between T. tianeensis and T. fengshanensis).

Taxonomy

. Triplophysa baishuijiangensis

Shi, Chen, Yang & X.-A. Wang sp. nov.

9CBCB7B0-7F40-5A0C-AAE6-3606589F7F02

https://zoobank.org/8DE249E6-3B59-46D6-BB24-8ECA30E874E5

Figs 1, 2, 3, 4, 5, Table 2

Figure 2.

Figure 2.

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Morphological characteristics of holotype KIZ2025000165 of Triplophysa baishuijiangensis sp. nov. A. Lateral view; B. Dorsal view; C. Ventral view; D. Lateral view of head; E. Dorsal view of head; F. Ventral view of head; G. Lateral and; H. Dorsal views of the X-ray graph.

Figure 3.

Figure 3.

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Sexual dimorphism of Triplophysa baishuijiangensis sp. nov. A. Female; B. Male.

Figure 4.

Figure 4.

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A. Habitat; B. Live photo of Triplophysa baishuijiangensis sp. nov. Red circles indicate the subterranean river outlet.

Table 2.

Morphometric characters data of Triplophysa baishuijiangensis sp. nov.

Characters Holotype Holotype + Paratypes (2025000165-175)
KIZ 2025000165 Min Max Mean SD
Standard length(SL)/mm 61,5 31,7 64,7 49,0
Head length(HL)/mm 15,2 7,9 16,1 12,5
Percent of SL (%)
Body depth 14,5 10,9 15,9 13,5 1,5
Body width 11,7 7,9 12,6 10,1 1,7
Length of caudal peduncle 17,4 15,1 18,7 16,8 1,3
Caudal peduncle depth 7,6 5,6 8,3 7,1 0,8
Predorsal length 55,7 52,6 57,2 55,2 1,3
Prepectoral length 25,0 25,0 28,5 26,8 1,3
Prepelvic length 56,4 54,6 59,3 56,8 1,3
Preanal length 76,7 73,3 79,1 76,9 1,8
Dorsal fin base length 12,3 10,5 14,0 12,4 0,9
Dorsal fin length 21,4 19,4 22,8 20,8 1,2
Pectoral fin base length 5,9 3,8 5,9 4,6 0,7
Pectoral fin length 22,1 20,2 23,8 21,8 1,0
Pelvic fin base length 4,8 3,0 4,8 3,6 0,5
Pelvic fin length 16,0 14,1 17,0 15,6 1,0
Anal fin base length 7,3 7,3 9,8 8,5 0,7
Anal fin length 16,8 14,0 19,0 16,8 1,3
Caudal fin Length 21,9 20,7 26,2 22,6 1,6
Δ (upper - lower caudal fin lobe length) 2,7 1,0 3,2 2,2 0,7
pectoral–pelvic distance 19,2 23,4 33,3 29,9 3,7
pelvic–anal distance 13,5 13,6 20,0 17,1 2,1
vent–anal fin origin distance 2,6 2,6 6,7 4,6 1,3
HL 24,7 24,7 27,0 25,5 0,8
Percent of HL (%)
Head depth at nape 50,1 41,0 50,3 45,7 3,3
Snout length 49,7 43,2 52,9 47,8 2,8
Eye diameter 6,1 2,9 6,1 4,8 1,2
Interorbital width 35,3 25,2 38,6 30,3 4,6
Preanterior nostril length 29,3 20,8 29,7 24,6 3,3
Distance between posterior nares 32,1 21,9 32,1 27,5 3,6
Upper jaw length 22,2 18,0 25,1 22,0 2,2
Lower jaw length 20,2 15,4 20,2 17,8 1,8
Mouth slit wide 26,6 23,5 30,5 26,5 2,3
Outer rostral barbel length 36,4 27,6 42,7 35,0 4,9
Internal rostral barbel length 23,3 19,2 25,0 22,1 2,1
Maxillary barbel length 34,9 25,3 35,0 30,7 3,3
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Type material.

Holotype. • KIZ2025000165 (Fig. 2), 75.64 mm total length, 61.5 mm standard length (SL), collected by Min Shi, Long Zhu, Yuan-Chao Chen, Jian-Fu Wei on May 17, 2023, at Xiaoganxi Village, Niujie town, Yiliang County, Yunnan Province, China, an outlet of a subterranean river belonging to the Baishui River system, Jingsha River drainage (27.82093589°N, 104.47385841°E; ca 628 m. a.s.l.; Fig. 5). Paratypes. • KIZ2025000175, collected on March 11, 2023; KIZ2025000166–174 (9 specimens) collected on May 17. 31.7–64.7 mm SL, other collection information same as holotype.

Diagnosis.

The diagnostic characters for cave-dwelling species of Triplophysa are provided in Suppl. material 2. Triplophysa baishuijiangensis sp. nov. is distinguished from its congeners by the following combination of characters:

  1. Vestigial eyes (vs absent or normal in T. nanpanjiangensis Zhu & Cao, 1988, T. huapingensis, T. sanduensis Chen & Peng, 2019, T. rongduensis Mao, Zhao, Yu, Xiao & Zhou, 2023, T. panzhouensis Yu, Luo, Lan, Xiao & Zhou, 2023, T. anlongensis, T. baotianensis Li, Liu & Li, 2018, T. xiangshuiqingensis Li, 2004, T. zhenfengensis Wang & Li, 2001, T. nandanensis, T. tianxingensis Yang, Li & Chen, 2016, T. guizhouensis, T. nasobarbatula Wang & Li, 2001, T. longliensis, T. fengshanensis, T. posterodorsalus (Li, Ran & Chen, 2006), T. qini, T. qiubeiensis Li & Yang, 2008, T. shilinensis, T. xuanweiensis, T. erythraea, T. anshuiensis, T. xiangxiensis, T. gejiuensis, T. flavicorpus Yang, Chen & Lan, 2004, T. yunnanensis Yang, 1990, and other epigean congeners);

  2. Skin pigment absent (vs reduced or covers the whole body in T. luochengensis, T. qingzhenensis, T. macrocephala, T. wudangensis, T. longipectoralis, T. xichouensis, T. ziyunensis Wu, Luo, Xiao & Zhou, 2024, T. yaluwang Lan, Liu, Zhou & Zhou, 2024, and T. wenshanensis Jiang, Cao, Song, Yi & Yang, 2025);

  3. Tip of pelvic fin reaching anus (vs not reaching anus in T. tianlinensis, T. aluensis Li & Zhu, 2000, and T. tianeensis);

  4. Lateral line complete (vs incomplete in T. langpingensis Yang, 2013);

  5. Posterior chamber of air bladder developed (vs reduced in T. yangi Jiang, Cao, Song, Yi & Yang, 2025);

  6. Dorsal fin iii, 7 (vs iv, 9 in T. cehengensis Luo, Mao, Zhao, Xiao & Zhou, 2023, and iii, 9 in T. rosa).

Description.

Detailed morphometric data of Triplophysa baishuijiangensis sp. nov. specimens are summarized in Table 2 and Suppl. material 4. Dorsal fin iii, 7; pectoral fin i, 9–10; pelvic fin i, 6; anal fin ii, 5; 15–16 branched rays in the caudal fin; vertebrae: 4+34.

Body elongated, anteriorly cylindrical, posteriorly compressed laterally behind dorsal-fin origin. Maximum body depth at tip of adpressed pectoral fins (corresponding to the swollen position of posterior swim bladder chamber). Snout slightly pointed. Eyes vestigial, reduced to black pigment spots.

Three pairs of barbels: outer rostral barbel longest, extending beyond posterior nostril; maxillary barbel shorter, extending past the eye spot; inner rostral barbel shortest, extending to the corner of mouth. Mouth inferior, arched, lower lip folded with medial V-shaped notch. Anterior and posterior nostrils closely connected, anterior nostril enclosed within a nasal valve that extends into a barbel-like tip, posterior nostril larger and without nasal valve.

Posterior edge of dorsal fin straight, distance from dorsal fin base to snout tip greater than to caudal fin base, dorsal fin starts before pelvic fins. Pectoral fins developed, arc-shaped, tips reaching or exceeding midpoint between dorsal and pectoral fin bases. Pelvic fins arc-shaped, slightly behind dorsal fin, extending to anus. Posterior edge of anal fin straight, separated from anus by a short distance. Caudal fin shallowly concave, upper lobe slightly longer than lower. Faint adipose crests on both dorsal and ventral sides of caudal peduncle, dorsal crest larger than ventral.

Two chambers of swim bladder, anterior chambers dumbbell-shaped and encased in bony capsule, posterior chamber developed, oval, and free within the abdominal cavity (Fig. 2G, H). Gill rakers not developed, nine gill rakers on first gill arch. Intestine short, length 52.0% of SL, bends downward behind the stomach and gradually narrows, stomach is U-shaped and enlarged.

Coloration.

Entire body lacks pigment and scaleless, lateral line complete. Live specimens semi-transparent, flesh-colored, internal organs vaguely visible ventrally. Fin rays white, fin membranes transparent (Fig. 4B). After fixation, body turns white, abdomen pale yellow. Eye spot color intensity varies, with some nearly invisible.

Sexual dimorphism.

All examined specimens lack thickened pads and furcella on the head and pectoral fins. However, males and females show significant differences in head shape. Male heads narrow sharply at the eye position, then taper gradually, resembling a “bell shape” with a blunt snout. Female heads taper gradually, resembling a “triangle” with a pointed snout (Fig. 3A, B). Consistent with the sexual dimorphism characteristics of T. daqiaoensis, described by Hou et al. (2010).

Distribution and habitat.

So far, the new species has only been found at the outlet of an underground river in Xiaoganxi Village, Yiliang County, Zhaotong City, Yunnan Province. Water of outlet is clear year-round (Fig. 4A). The new species is active and feeds near the outlet, rarely far from the outlet, responding to light but is insensitive to vibrations, and moves slowly and is easy to capture.

Etymology.

The specific epithet baishuijiangensis is in reference to the type locality of the new species: the core area of Baishuijiang National Aquatic Germplasm Resources Reserve for Endemic Fish. We propose the common English name “Baishuijiang high-plateau loach” and the Chinese name “Bái Shuǐ Jiāng Gāo Yuán Qiū (白水江高原鳅)”.

Discussion

Currently documented cave-dwelling species of Triplophysa are predominantly distributed in karst landscapes of eastern Yunnan, northern Guangxi, southwestern Guizhou, and the Hunan-Chongqing border region. Hydrologically, with the exception of T. wenshanensis, T. xichouensis, and T. gejiuensis within the Red River system, all documented species inhabit tributaries of the Yangtze River (including Wujiang, Yuanjiang, and Hengjiang on the right bank) and Pearl River systems (predominantly Beipanjiang, Nanpanjiang, Hongshuihe, and Liujiang). The Yangtze River system harbors seven species compared to 33 species in the Pearl River system (Lan et al. 2004; Cao et al. 2025; Suppl. material 1). This biogeographical pattern reveals marked spatial heterogeneity: northern populations (Yangtze system) demonstrate lower species richness with scattered distributions, whereas southern populations (Pearl River system) exhibit higher species diversity and aggregation density (Fig. 5). This pattern likely reflects differential development of karst geomorphology between regions or uneven historical dispersal processes.

High congruence exists between the phylogenetic structure and geographic distribution pattern in hypogean species of Triplophysa. Phylogenetic analyses delineate two major clades within the hypogean group. Species of Clade I are distributed across eastern Yunnan, western Guizhou, and northwestern Guangxi regions, while species of Clade II occupy eastern territories relative to Clade I. The two distribution ranges are allopatric, with the extension line of lower reaches of the Beipanjiang River approximately serving as the east-west demarcation boundary between them (Figs 1, 5).

The trogloxene group (Claea spp.), sister to these hypogean group (Fig. 1), is currently found only in upper Yangtze reaches (Zhang et al. 2024; Fricke et al. 2025). As a lineage embedded within Triplophysa, Claea retains generic validity due to absent secondary sexual traits and processus dentiformis on the upper jaw, current evidence thus suggests the polyphyly of Triplophysa (Zhang et al. 2024; Lei et al. 2025). With more than 160 documented species, the taxonomic revision of Triplophysa is a significant challenge. Consequently, both the taxonomic status of Claea and whether the hypogean group of Triplophysa represents a distinct genus require further systematic study. Among the 43 cave-dwelling Triplophysa species, 16 species inhabit cave entrances, subterranean river outlets, or surface water bodies connected to subterranean systems (Suppl. material 1). Moreover, 33 species (77%) retain normal or partially degenerated eyes (Luo et al. 2023a; Lan et al. 2004; Cao et al. 2025; Suppl. material 2). Notably, T. baishuijiangensis, despite exhibiting extreme ocular degeneration and pigment loss, retains the ability to survive briefly outside the cave near its outlets, suggesting that some cave-dwelling Triplophysa species retain limited adaptability to surface environments. These species may periodically leave caves independently or be displaced by flooding events, resulting in short-distance dispersal to other suitable subterranean habitats. Interspecific genetic distances among cave-dwelling Triplophysa are relatively low (Suppl. material 3), with 13 species pairs (15 species in total) exhibiting genetic distances below 2.00%: T. qingzhenensis and T. wudangensis (1.62%), T. macrocephala and T. longipectoralis (1.51%), T. wudangensis and T. rosa (1.44%), T. qingzhenensis and T. rosa (1.42%), T. longipectoralis and T. nasobarbatula (1.24%), T. rongduensis and T. zhenfengensis (0.97%), T. nasobarbatula and T. macrocephala (0.80%), T. longliensis and T. sanduensis (0.71%), T. tianeensis and T. fengshanensis (0.35%), T. guizhouensis and T. yaluwang (1.45%), T. ziyunensis and T. qingzhenensis (1.73%), T. ziyunensis and T. wudangensis (1.45%), T. ziyunensis and T. rosa (1.54%). The relatively low genetic distances and geographic distance (Fig. 5) of these species pairs may suggest relatively recent isolation, speciation or introgression events (Liu et al. 2023; Yuan et al. 2023). Collectively, these findings suggest that cave-dwelling Triplophysa may have undergone rapid adaptive radiation and dispersal within the southwestern karst region. Furthermore, the growth rate of lineage numbers in cave-dwelling Triplophysa reached its peak during the late stages of differentiation (Yan 2018), which further corroborates the aforementioned viewpoint.

Triplophysa baishuijiangensis sp. nov. marks the first recorded cave loach in the Yangtze River basin within Yunnan and simultaneously the first documented typical cavefish in the Jinsha River system. Its discovery validates the presence of environmental conditions conducive to cavefish evolution in this basin. Field investigations conducted by authors reveal extensive karst development in Hengjiang and Chishui River basins, with the Chishui River basin exhibiting particularly dense subterranean cave systems and underground river networks, highlighting the potential cave biodiversity in these areas and warranting further investigation and research.

Comparative material examined

T. shilinensis: KIZ1991000936–937, 59.1–59.3 mm SL, China: Yunan Province: Shilin County: Weiboyi Village.

T. anshuiensis: KIZ2012005746–5747, 64.7–68.4 mm SL, China: Guangxi Zhuang Autonomous Region: Lingyun County: Anshui Village.

T. gejiuensis: KIZ1978000982–985, 41.6–45.5 mm SL, China: Yunan Province: Gejiu City: Kafan Town.

T. huapingensis: KIZ2008007606–7608, 57.6–61.7 mm SL, China: Guangxi Zhuang Autonomous Region: Leye County: Huaping Town.

T. nandanensis: KIZ9110001135–1137, 53.2–79.0 mm SL, China: Guangxi Zhuang Autonomous Region: Nandan County: Liuzhai Town: Mayang Village: Xijiang River.

T. longliensis: KIZ2010002987–2988, KIZ2010003221, 75.9–97.9 mm SL, China: Guizhou Province: Longli County: Baisheng Village: Hongshuihe River.

T. guizhouensis: KIZ2017000346, KIZ2017000348, 73.5–110.3 mm SL, China: Guizhou Province: Huishui County: Baijin Village: Hongshuihe River.

T. rosa: KIZ2002005675, 53.8 mm SL, China: Chongqing City: Wulong County: Jiangkou Town: Tianxing Village: Wujiang River.

T. tianlinensis: KIZ2012005690–5691, KIZ2012005693, 48.7–84.7 mm SL, China: Guangxi Zhuang Autonomous Region: Tianlin County: Langping Town: Hongxing Village: Hongshui River.

T. tianeensis: KIZ2003005724–5729, 57.4–58.9 mm SL, China: Guangxi Zhuang Autonomous Region: Tiane County: Bala Town: Hongshui River.

T. macrocephala: KIZ2010003078, KIZ2010003080–3081, 50.4–54.7 mm SL, China: Guangxi Zhuang Autonomous Region: Nandan County: Lihu Town: Renguang Village: Xijiang River.

T. luochengensis: KIZ2014005967–5968, 45.0–53.8 mm SL, China: Guangxi Zhuang Autonomous Region: Luocheng County: Jicheng Village: Hongshuihe River.

T. longipectoralis: KIZ2001004573–4574, KIZ2001004579, 46.9–51.9 mm SL, China: Guangxi Zhuang Autonomous Region: Hechi City: Huanjiang County: Xunle Town: Liujiang River.

T. xichouensis: KIZ2013600381–382, 57.4–61.4 mm SL, China: Yunan Province: Xichou County: Xisa Town: Red River.

Supplementary Material

XML Treatment for Triplophysa baishuijiangensis

Acknowledgements

The Second Comprehensive Survey and Research on the Qinghai Tibet Plateau (2019QZKK05010103); The adjustment of central refined oil prices had provided subsidies for fisheries in 2023 – a survey of local resources of cave organisms and large benthic invertebrates in Zhenxiong County, Yunnan province; Program of Yunnan Provincial Science and Technology Department Foundation, Grant/Award Number: 202105AD160026, 202301AT070305 and 202303AC100011.

Citation

Shi M, Chen Y-C, Che X-J, Dao W, Liu W-M, Wang D-S, Yang J-X, Wang X-A (2025) A new typical cavefish of the genus Triplophysa (Teleostei, Cypriniformes, Nemacheilidae) from the Jinsha River, Yunnan, China. ZooKeys 1253: 73–92. https://doi.org/10.3897/zookeys.1253.153155

Funding Statement

The Second Comprehensive Survey and Research on the Qinghai Tibet Plateau(2019QZKK05010103);The adjustment of central refined oil prices had provided subsidies for fisheries in 2023—a survey of local resources of cave organisms and large benthic invertebrates in Zhenxiong County,Yunnan province;Program of Yunnan Provincial Science and Technology Department Foundation,Grant/Award Number：202105AD160026,202301AT070305 and 202303AC100011.

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

No funding was reported.

Author contributions

Min Shi: Investigation (equal); Writing-original draft (equal); Data curation (equal); Writing-review and editing (equal). Yuan-Chao Chen: Investigation (equal); Writing-original draft (equal); Writing-review and editing (equal). Xing-Jin Che: Formal analysis (equal); Validation (equal). Wei Dao: Data curation (equal); Formal analysis (equal). Wen-Ming Liu: Supervision (equal); Project administration (equal). Deng-Shan Wang: Supervision (equal); Project administration (equal). Jun-Xing Yang: Writing-review and editing (equal); Resources (equal). Xiao-Ai Wang: Writing-review and editing (equal); Resources (equal); Visualization (equal); Project administration (equal).

Data availability

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

Supplementary materials

Supplementary material 1

Species distribution table

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.

Min Shi

Data type

xlsx

Supplementary material 2

Morphological comparison table of cave-dwelling Triplophysa

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.

Min Shi

Data type

xlsx

Supplementary material 3

Cyt b Genetic K2p distance of 43 sequences

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.

Min Shi

Data type

xls

Supplementary material 4

Supplementary material 4-original measurement data of Triplophysa baishuijiangensis sp. nov.

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.

Min Shi

Data type

xlsx

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Associated Data

This section collects any data citations, data availability statements, or supplementary materials included in this article.

Supplementary Materials

XML Treatment for Triplophysa baishuijiangensis
zookeys.1253.153155-treatment1.xml (42.2KB, xml)
Supplementary material 1

Species distribution table

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.

Min Shi

Data type

xlsx

zookeys-1253-073_article-153155__-s001.xlsx (15.8KB, xlsx)
Supplementary material 2

Morphological comparison table of cave-dwelling Triplophysa

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.

Min Shi

Data type

xlsx

zookeys-1253-073_article-153155__-s002.xlsx (22.3KB, xlsx)
Supplementary material 3

Cyt b Genetic K2p distance of 43 sequences

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.

Min Shi

Data type

xls

zookeys-1253-073_article-153155__-s003.xls (48.5KB, xls)
Supplementary material 4

Supplementary material 4-original measurement data of Triplophysa baishuijiangensis sp. nov.

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.

Min Shi

Data type

xlsx

zookeys-1253-073_article-153155__-s004.xlsx (15.2KB, xlsx)

Data Availability Statement

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

Articles from ZooKeys are provided here courtesy of Pensoft Publishers

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