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Research Article
Sinocyclocheilus xiejiahuai (Cypriniformes, Cyprinidae), a new cave fish with extremely small population size from western Guizhou, China
expand article infoCui Fan‡, Man Wang§|, Jia-Jia Wang‡, Tao Luo‡, Jia-Jun Zhou, Ning Xiao#, Jiang Zhou‡
‡ Guizhou Normal University, Guiyang, China
§ Institute of Hydrobiology, Chinese Academy of Sciences, Wuhan, China
| University of Chinese Academy of Sciences, Beijing, China
¶ Zhejiang Forest Resource Monitoring Center, Hangzhou, China
# Guiyang Healthcare Vocational University, Guiyang, China

Corresponding author: Ning Xiao ( armiger@163.com )

Corresponding author: Jiang Zhou ( zhoujiang@ioz.ac.cn )

Academic editor: Maria Elina Bichuette
© 2024 Cui Fan, Man Wang, Jia-Jia Wang, Tao Luo, Jia-Jun Zhou, Ning Xiao, Jiang Zhou.
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: Fan C, Wang M, Wang J-J, Luo T, Zhou J-J, Xiao N, Zhou J (2024) Sinocyclocheilus xiejiahuai (Cypriniformes, Cyprinidae), a new cave fish with extremely small population size from western Guizhou, China. ZooKeys 1214: 119-141. https://doi.org/10.3897/zookeys.1214.127629
ZooBank: urn:lsid:zoobank.org:pub:15735CAD-D139-4C84-9109-DB7F1782C65A
Open Access

Abstract

This study describes a new species, Sinocyclocheilus xiejiahuai sp. nov., discovered within a cave located in Hongguo Town, Panzhou City, Guizhou Province, southwestern China, with the type locality in the Nanpanjiang River basin. Phylogenetic trees reconstructed based on mitochondrial genes show that the new species represents an independent evolutionary lineage with large genetic differences, 1.9%–13.8% in mitochondrial Cyt b, from congeners. Morphologically, this species can be differentiated from the 79 species currently classified under the genus Sinocyclocheilus by several characteristics: absence of horn-like structures and indistinct elevation at the head-dorsal junction, absence of irregular black markings on the body lateral and scaleless, eyes large, eye diameter 13% of head length, dorsal-fin rays, iii, 6½, last unbranched ray strong, with serrations along posterior margin, pectoral-fin rays, i, 13, anal-fin rays, iii, 5, pelvic-fin rays, i, 7, lateral line pores 74, gill rakers well developed, nine on first gill arch, pectoral fins short, tip not reaching to pelvic-fin origin. The number of Sinocyclocheilus species has been increased from 79 to 80 since the description of this new species.

Key words

Cavefish, new species, morphology, phylogeny, taxonomy

Introduction

The genus Sinocyclocheilus Fang, 1936 (Cypriniformes: Cyprinidae) is endemic to China, and is currently found only in the Yangtze, Pearl, Lancangjiang, and Yuanjiang rivers (Xu et al. 2023). Based on recent taxonomic and phylogenetic studies, the genus Sinocyclocheilus includes 79 valid species (Shan et al. 2000; Zhao and Zhang 2009; Zhang et al. 2016; Luo et al. 2023; Xu et al. 2023; Shao et al. 2024), most of which are classified into five species groups, i.e., S. angularis species group includes 21 valid species, S. cyphotergous species group includes 20 valid species, S. microphthalmus species group includes three valid species, S. jii species group includes five valid species, and S. tingi species group includes 26 valid species (Zhao and Zhang 2009; Wen et al. 2022; Luo et al. 2023; Xu et al. 2023; Shao et al. 2024) (Table 1). The S. angularis, S. cyphotergous, and S. microphthalmus species groups have stygobite morphology, and the S. jii and S. tingi species groups a mixture of stygobite and stygophile morphology (Zhao and Zhang 2009). There are 25 species currently recorded for the S. tingi species group, of which 18, four, two, and one are distributed in the Nanpanjiang, Yuanjiang, Jinshajiang, and Lancangjiang rivers, respectively, and the new species is distributed in the Beipanjiang River (Fig. 1) (Zhao and Zhang 2009; Luo et al. 2023; Xu et al. 2023).

Figure 1. 

Sampling collection localities and distribution of the Sinocyclocheilus xiejiahuai sp. nov. and 26 species of the S. tingi species group of the genus Sinocyclocheilus in southwest China. The base maps are from Standard Map Service website (http://211.159.153.75/).

Table 1.
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List of 79 currently recognized species of the genus Sinocyclocheilus endemic to China and references. Recognized species modified from Jiang et al. (2019) and Xu et al. (2023).

ID Species Species group Province Rivers Literature obtained
1 S. altishoulderus (Li & Lan, 1992) S. angularis group Guangxi Hongshui River Li and Lan 1992
2 S. anatirostris Lin & Luo, 1986 S. angularis group Guangxi Hongshui River Lin and Luo 1986
3 S. angularis Zheng & Wang, 1990 S. angularis group Guizhou Beipanjiang River Zheng and Wang 1990
4 S. aquihornes Li & Yang, 2007 S. angularis group Yunnan Nanpanjiang River Li et al. 2007
5 S. bicornutus Wang & Liao, 1997 S. angularis group Guizhou Beipanjiang River Wang and Liao 1997
6 S. brevibarbatus Zhao, Lan & Zhang, 2009 S. angularis group Guangxi Hongshui River Zhao et al. 2009
7 S. broadihornes Li & Mao, 2007 S. angularis group Yunnan Nanpanjiang River Li and Mao 2007
8 S. convexiforeheadus Li, Yang & Li, 2017 S. angularis group Yunnan Nanpanjiang River Yang et al. 2017
9 S. hyalinus Chen & Yang, 1994 S. angularis group Yunnan Nanpanjiang River Chen et al. 1994
10 S. longicornus Luo, Xu, Wu, Zhou & Zhou, 2023 S. angularis group Guizhou Nanpanjiang River Xu et al. 2023
11 S. jiuxuensis Li & Ran, 2003 S. angularis group Guangxi Hongshui River Li et al. 2003c
12 S. flexuosdorsalis Zhu & Zhu, 2012 S. angularis group Guangxi Hongshui River Zhu and Zhu 2012
13 S. furcodorsalis Chen, Yang & Lan, 1997 S. angularis group Guangxi Hongshui River Chen et al. 1997
14 S. mashanensis Wu, Liao & Li, 2010 S. angularis group Guangxi Hongshui River Wu et al. 2010
15 S. rhinocerous Li & Tao, 1994 S. angularis group Yunnan Nanpanjiang River Li and Tao 1994
16 S. simengensis Li, Wu, Li & Lan, 2018 S. angularis group Guangxi Hongshui River Wu et al. 2018
17 S. tianeensis Li, Xiao & Luo, 2003 S. angularis group Guangxi Hongshui River Li et al. 2003d
18 S. tianlinensis Zhou, Zhang, He & Zhou, 2004 S. angularis group Guangxi Nanpanjiang River Zhou et al. 2004
19 S. tileihornes Mao, Lu & Li, 2003 S. angularis group Yunnan Nanpanjiang River Mao et al. 2003
20 S. xingyiensis Luo, Tang, Deng, Duan & Zhang, 2023 S. angularis group Guizhou Nanpanjiang River Luo et al. 2023
21 S. zhenfengensis Liu, Deng, Ma, Xiao & Zhou, 2018 S. angularis group Guizhou Beipanjiang River Liu et al. 2018
22 S. anshuiensis Gan, Wu, Wei & Yang, 2013 S. microphthalmus group Guangxi Hongshui River Gan et al. 2013
23 S. longshanensis Li & Wu, 2018 S. microphthalmus group Yunnan Nanpanjiang River Li et al. 2018
24 S. microphthalmus Li, 1989 S. microphthalmus group Guangxi Hongshui River Li 1989
25 S. aluensis Li & Xiao, 2005 S. tingi group Yunnan Nanpanjiang River Li et al. 2005; Zhao and Zhang 2013
26 S. angustiporus Zheng & Xie, 1985 S. tingi group Guizhou; Yunnan Nanpanjiang River Zheng and Xie 1985; Zhao and Zhang 2009
27 S. anophthalmus Chen & Chu, 1988 S. tingi group Yunnan Nanpanjiang River Chen et al. 1988a
28 S. bannaensis Li, Li & Chen, 2019 S. tingi group Yunnan Lancangjiang River Li et al. 2019
29 S. grahami (Regan, 1904) S. tingi group Yunnan Jinshajiang River Zhao and Zhang 2009
30 S. guishanensis Li, 2003 S. tingi group Yunnan Nanpanjiang River Li et al. 2003a
31 S. huaningensis Li, 1998 S. tingi group Yunnan Nanpanjiang River Li et al. 1998
32 S. huizeensis Cheng, Pan, Chen, Li, Ma & Yang, 2015 S. tingi group Yunnan Jinshajiang River Cheng et al. 2015
33 S. lateristriatus Li,1992 S. tingi group Yunnan Nanpanjiang River Li 1992
34 S. longifinus Li, 1998 S. tingi group Yunnan Nanpanjiang River Li et al. 1998
35 S. macrocephalus Li,1985 S. tingi group Yunnan Nanpanjiang River Li 1985
36 S. macroscalus Li, 1992 S. tingi group Yunnan Nanpanjiang River Li 1992
37 S. maculatus Li, 2000 S. tingi group Yunnan Nanpanjiang River Zhao and Zhang 2009; Li et al. 2000a
38 S. maitianheensis Li,1992 S. tingi group Yunnan Nanpanjiang River Li 1992
39 S. malacopterus Chu & Cui, 1985 S. tingi group Yunnan Nanpanjiang River Chu and Cui 1985
40 S. oxycephalus Li, 1985 S. tingi group Yunnan Nanpanjiang River Li 1985
41 S. purpureus Li, 1985 S. tingi group Yunnan Nanpanjiang River Li 1985
42 S. qiubeiensis Li, 2002 S. tingi group Yunnan Nanpanjiang River Li et al. 2002b
43 S. qujingensis Li, Mao & Lu, 2002 S. tingi group Yunnan Nanpanjiang River Li et al. 2002c
44 S. robustus Chen & Zhao, 1988 S. tingi group Guizhou Nanpanjiang River Chen et al. 1988b
45 S. tingi Fang, 1936 S. tingi group Yunnan Nanpanjiang River Zhao and Zhang 2009
46 S. wenshanensis Li, Yang, Li & Chen, 2018 S. tingi group Yunnan Yuanjiang River Yang et al. 2018
47 S. wumengshanensis Li, Mao, Lu & Yan, 2003 S. tingi group Yunnan Yuanjiang River Li et al. 2003a
48 S. xichouensis Pan, Li, Yang & Chen, 2013 S. tingi group Yunnan Yuanjiang River Pan et al. 2013
49 S. yangzongensis Chu & Chen, 1977 S. tingi group Yunnan Nanpanjiang River Zhao and Zhang 2009
50 S. yimenensis Li & Xiao, 2005 S. tingi group Yunnan Yuanjiang River Li et al. 2005
51 S. brevis Lan & Chen, 1992 S. cyphotergous group Guangxi Liujiang River Chen and Lan 1992
52 S. cyphotergous (Dai, 1988) S. cyphotergous group Guizhou Hongshui River Huang et al. 2017
53 S. donglanensis Zhao, Watanabe & Zhang, 2006 S. cyphotergous group Guangxi Hongshui River Zhao et al. 2006
54 S. donglanensis Zhou, Liu & Wang, 2011 S. cyphotergous group Guizhou Liujiang River Zhou et al. 2011
55 S. gracilicaudatus Zhao & Zhang, 2014 S. cyphotergous group Guangxi Liujiang River Wang et al. 2014
56 S. guiyang Shao, Cheng, Lu, Zhou & Zeng, 2024 S. cyphotergous group Guizhou Yangtze River Shao et al. 2024
57 S. huanjiangensis Wu, Gan & Li, 2010 S. cyphotergous group Guangxi Liujiang River Wu et al. 2010
58 S. hugeibarbus Li, Ran & Chen, 2003 S. cyphotergous group Guizhou Liujiang River Li et al. 2003b
59 S. lingyunensis Li, Xiao & Lu, 2000 S. cyphotergous group Guangxi Hongshui River Li et al. 2000
60 S. longibarbatus Wang & Chen, 1989 S. cyphotergous group Guizhou; Guangxi Liujiang River Wang and Chen 1989
61 S. luopingensis Li & Tao, 2002 S. cyphotergous group Yunnan Nanpanjiang River Li et al. 2002a
62 S. macrolepis Wang & Chen, 1989 S. cyphotergous group Guizhou; Guangxi Liujiang River Wang and Chen 1989
63 S. macrophthalmus Zhang & Zhao, 2001 S. cyphotergous group Guangxi Hongshui River Zhang and Zhao 2001
64 S. multipunctatus (Pellegrin, 1931) S. cyphotergous group Guizhou; Guangxi Liujiang River; Hongshui River; Wujiang River Zhao and Zhang 2009
65 S. punctatus Lan & Yang, 2017 S. cyphotergous group Guizhou; Guangxi Liujiang River; Hongshui River Lan et al. 2017
66 S. ronganensis Luo, Huang & Wen, 2016 S. cyphotergous group Guangxi Liujiang River Luo et al. 2016
67 S. sanxiaensis Jiang, Li, Yang & Chang, 2019 S. cyphotergous group Hubei Yangtze River Jiang et al. 2019
68 S. xunlensis Lan, Zhan & Zhang, 2004 S. cyphotergous group Guangxi Liujiang River Lan et al. 2004
69 S. yaolanensis Zhou, Li & Hou, 2009 S. cyphotergous group Guizhou Liujiang River Zhou et al. 2009
70 S. yishanensis Li & Lan, 1992 S. cyphotergous group Guangxi Liujiang River Li and Lan 1992
71 S. brevifinus Li, Li & Mayden, 2014 S. jii group Guangxi Hejiang River Li et al. 2014
72 S. guanyangensis Chen, Peng & Zhang, 2016 S. jii group Guangxi Guijiang River Chen et al. 2016
73 S. guilinensis Ji, 1985 S. jii group Guangxi Guijiang River Zhao and Zhang 2009
74 S. huangtianensis Zhu, Zhu & Lan, 2011 S. jii group Guangxi Hejiang River Zhu et al. 2011
75 S. jii Zhang & Dai, 1992 S. jii group Guangxi Guijiang River Zhang and Dai 1992
76 S. gracilis Li, 2014 No assignment Guangxi Guijiang River Li and Li 2014
77 S. luolouensis Lan, 2013 No assignment Guangxi Hongshui River Lan et al. 2013
78 S. pingshanensis Li, Li, Lan & Wu, 2018 No assignment Guangxi Liujiang River Wu et al. 2018
79 S. wui Li & An, 2013 No assignment Yunnan Mingyihe River Li and An 2013

During our biodiversity survey in southwestern Guizhou Province, China, in October 2019, a specimen of Sinocyclocheilus with normal eyes, scaleless and absence of irregular black markings on the body lateral was collected in a completely dark cave. This specimen was identified to the S. tingi species group based on morphological characters. Subsequent morphological examination and molecular evidence suggest that this specimen represents an undescribed species of the S. tingi species group within the genus Sinocyclocheilus. However, between 2019 and 2023, we conducted 16 more surveys in this cave again, none of which revealed any new individuals. Considering the conservation of the species and the rescue of diversity, here we formally describe the species as Sinocyclocheilus xiejiahuai sp. nov. based on the single-numbered specimen. Although there is only one specimen of this species, the significance of its discovery is as important as that of the publication of Sinocyclocheilus sanxiaensis in terms of zoogeography and conservation biogeography (Jiang et al. 2019).

Materials and methods

Sampling and preservation

The single specimen of the genus Sinocyclocheilus were collected in southwestern Guizhou Province, China during a cave fish diversity survey in southern China in October 2019. Gill muscle tissue was preserved in 95% alcohol at -20 °C for molecular analysis. All specimens were fixed in 7% buffered formalin and then transferred to 75% ethanol for long-term storage. Specimen were preserved at Guizhou Normal University, Guiyang City, Guizhou Province, China.

Morphological comparison and statistical analysis

The new species can be placed in the S. tingi species group based on morphology and can be clearly distinguished from species in the S. angularis, S. cyphotergous, and S. microphthalmus, S. jii species groups, e.g., absence of horn-like structures and indistinct elevation at the head-dorsal junction; pectoral fins short, not reaching to pelvic-fin origin; and with serrations along posterior margin of the last unbranched fin of the dorsal fin (Zhao et al. 2009). Therefore, this study focused on morphological comparisons with 26 species within the S. tingi species group (Table 2).

Table 2.
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Comparison of the diagnostic features of the new species described here with those selected for the 26 species of the S. tingi group and four unassigned species (last four) within the genus Sinocyclocheilus. Grey shading indicates clear difference in character compared to that of Sinocyclocheilus xiejiahuai sp. nov.

Species Body lateral markings Gill rakers Dorsal-fin rays Pectoral-fin rays Anal-fin rays Pelvic-fin rays Caudal -fin rays Lateral-line scales/pores Tip of pectoral fin reaching to ventral fin Tip of pelvic-fin rays reaching to anus Source
S. xiejiahuai sp. nov. Absent 9 iii, 6½; i, 13 iii, 5 i, 7 17 74 No No This study
S. aluensis Present 5–7 iii, 7 i, 13–16 ii, 5 i, 7–9 15–17 71–75 No No Li et al. 2005
S. angustiporus Present 7–9 iv,7 i, 14–16 iii, 5 i, 8 15–16 68–80 NA No Zhao and Zhang 2009
S. anophthalmus Absent 7–9 iv,8 i, 15–16 iii, 5 i, 8 16 52–56 Yes No Chen et al. 1988a
S. grahami Present 5–8 iii, 7 i, 15–17 iii, 5 i, 8–9 16 61–69 No No Zhao and Zhang 2009
S. guishanensis Present 5–6 iii, 7 i, 13–16 iii, 5 i, 7–8 15–16 73–80 No No Li et al. 2003a
S. huaningensis Present 6 iii, 7 i, 16 iii, 5 i, 8 16 59–67 No Yes Li et al. 1998
S. huizeensis Present 5–6 iii, 7 i, 15–16 iii, 5 i, 10 18 70–73 No No Cheng et al. 2015
S. bannaensis Present 5 iii, 8 i, 9 ii, 5 i, 9 16 47 Yes No Li et al. 2019
S. maculatus Present 14–17 iii, 7 i, 14–15 iii, 5 i, 7~8 16 81–88 No No Zhao and Zhang 2009
S. maitianheensis Present 7–8 iii, 7 i, 14–15 iii, 5 i, 9 18 70–82 No Yes Li 1992
S. malacopterus Present 7–9 iii, 7 i, 14–18 iii, 5 i, 9 15–16 67–81 No No Chu and Cui 1985
S. longifinus Absent NA iii, 7 i, 16 ii, 5 i, 8 17 70–72 Yes Yes Li et al. 1998
S. longshanensis Present 15–18 iii, 7 i, 14–15 ii, 5 i, 7–8 16 59–62 No No Li et al. 2018
S. macrocephalus Absent 12 iv, 7 i, 15–17 iii, 5 i, 8 16 72–78 No No Li 1985
S. lateristriatus Present 7–10 iv, 7 i, 15–16 iii, 5 i, 8 17 75–91 No No Li 1992
S. purpureus Present 7–8 iv, 6–7 i, 16 iii, 5 i, 8 NA 63–67 No No Li 1985
S. qiubeiensis Present 6–7 iii, 7 i, 14–17 iii, 5 i, 8–9 16 67–81 No No Li et al. 2002b
S. qujingensis Absent 6–8 iii, 7 i, 16 iii, 5 i, 8 16 70–79 No No Li et al. 2002c
S. robustus Present 9 iv, 7 i, 13–16 iii, 5 i, 7–8 16 72 No No Chen et al. 1988b
S. wumengshanensis Present 5–6 iii, 7 16 ii, 5 i, 8 16 67–76 Yes Yes Li et al. 2003a
S. xichouensis Present 6 iii, 6–7 i, 14–16 iii, 5 i, 8–9 NA 74–88 Yes No Pan et al. 2013
S. tingi Present 7–9 iv, 7 i, 14–16 iii, 5 i, 6–8 16 62–73 No No Zhao and Zhang 2009
S. yangzongensis Absent 8–11 iii, 7 i, 16 iii, 5 i, 9 16 71–81 No No Zhao and Zhang 2009
S. yimenensis Present 5–7 iii, 7 i, 14–15 ii, 5 i, 8 16-17 70–79 No No Li et al. 2005
S. oxycephalus Present 6–7 iv, 7 i, 16 iii, 5 i, 8 17 74–78 No No Li 1985
S. wenshanensis Present 7–9 iii, 7 i, 13–15 ii, 5 i, 7–8 14–15 67–72 No Yes Yang et al. 2018
S. macroscalus Present 9-10 iv, 7 i, 15–16 iii, 5 i, 8 NA 70-79 No No Li 1992
S. gracilis Absent 12 NA NA NA NA NA NA No No Li and Li 2014
S. pingshanensis Absent 10–12 iii, 7 i, 13–15 ii, 5 i, 7–8 16 75–78 Yes No Wu et al. 2018
S. luolouensis Present 10 iii, 7 i, 13–14 iii, 5 i, 7–8 16–17 40–49 Yes Yes Lan et al. 2013
S. wui Absent 7 iii, 7 i, 14–15 ii, 5 i, 7–8 14–15 79–81 No No Li and An 2013

We measured 32 morphometric data points (Suppl. material 1) from a total of nine specimens of three species, referenced from Xu et al. (2023). Principal component analysis (PCA) of corrected morphometric measurements and two-dimensional scatter plots were used to explore the relative contribution of specific variables to morphological variation. Prior to PCA analysis, all included measurements were normalized using ratios to standard length (standard length is the ratio to full length) followed by log transformation (Shao et al. 2024). PCA analyses were performed in SPSS 21.0 (SPSS, Inc., Chicago, IL, USA).

DNA extraction and sequencing

We sequenced the mitochondrial genomes of 13 species of the genus Sinocyclocheilus. Total genomic DNA was extracted from each sample of 95% ethanol-preserved tissue using the Cetyltrimethylammonium Bromide method. For mitogenome sequencing, genomic DNA was fragmented to an appropriate size of 150–500 bp using a Covaris Ultrasonicator. A 400 bp DNA library was constructed based on the Whole Genome Shotgun and the library size. Sequencing was performed on an Illumina NovaSeq 6000 platform using a paired-end 150 bp protocol, generating an average of ~ 5.4 Gb of raw data. The raw data were cleaned using SOAPnuke v. 1.3 (Chen et al. 2018) based on the following criteria: removal of reads with more than 5% N-base content, reads with 50% low-quality bases, and reads with adapter contamination. The process yielded ~ 5.3 Gb of clean data. Mitogenome assembly was performed on the clean data using SPAdes v. 3.13 (parameter: -k 127) (Bankevich et al. 2012), and the assembled contigs were used for BLASTN analysis (BLAST 2.2.30+, parameter: e-5) using a reference mitogenome (Sinocyclocheilus rhinocerous: KR069119) to identify possible assembly errors. Assembled mitogenomes were annotated for genes using MITOS (Bernt et al. 2013). For mitochondrial cytochrome b (Cyt b) and NADH dehydrogenase subunit 4 (ND4) genes, PCR amplifications and sequencing followed Xu et al. (2023).

Phylogenetic reconstruction and divergence time estimate

In total, we collected 43 mitochondrial genomes and 76 mitochondrial gene fragments (36 Cytb, 34 ND4, five 16S, three ND5, and three CO1) for phylogenetic reconstruction and estimation of divergence times. Following Wen et al. (2022), we selected Carassius auratus, Cyprinus carpio, Schizothorax yunnanensis, Onychostoma simum, Barbus barbus, Puntius ticto, Neolissochilus hexagonolepis, Garra orientalis, Myxocyprinus asiaticus, and Danio rerio as outgroup species (Table 3). All sequences were assembled and aligned using the MUSCLE (Edgar 2004) module in MEGA v. 7.0 (Kumar et al. 2016) with default settings. The best-fit model was obtained based on the Bayesian information criterion computed with PartitionFinder v. 2.1.1 (Lanfear et al. 2017) (Suppl. material 2).

Table 3.
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Localities, voucher information, and GenBank numbers for all samples used.

ID Species Location (* type localities) Voucher number Mitogenome Cyt b ND4/16S/ND5/CO1
1 S. altishoulderus Mashan County, Guangxi – NC_013186
2 S. anatirostris – GZNU20210531002 NC_069226
3 S. angularis Panzhou City, Guizhou* GZNU20180420001 MZ636514
4 S. angularis Baotian Town, Panzhou City, Guizhou GZNU20180420001 PQ157935
5 S. angustiporus Xingren County, Guizhou GZNU20190504001 MZ636515
6 S. anophthalmus – – NC_023472
7 S. anshuiensis Lingyun County, Guangxi – KR069120
8 S. aquihornes Shuanglongjian town, Qiubei County, Yunnan* S28 – PQ155086 PQ155094
9 S. bicornutus XingrenCounty, Guizhou – KX528071
10 S. brevibarbatus – GX0064–L20–13 – MT373106 MW548423
11 S. brevifinus – – – OQ718395
12 S. brevis – GX0155 – MT373105 MW548424
13 S. cyphotergous Luodian County, Guizhou* GZNU20150819010 OQ319607
14 S. convexiforeheadus Wenliu Township, Qiubei County, Yunnan* S30 – PQ155090 PQ155091
15 S. donglanensis Donglan County, Guangxi CA139 AB196440 MW548425
16 S. furcodorsalis Tian’e County, Guangxi – GU589570
17 S. gracilicaudatus – – – OQ718398
18 S. grahami Kunming City, Yunnan – GQ148557
19 S. guanyangensis – GX0173 – MT373108 MW548426
20 S. guilinensis – GX0073–L17–2 – MT373104 MW548427
21 S. guishanensis Guishan, Shilin County, Yunnan XH5401 – AY854722 AY854779
22 S. huangtianensis – GX0175 – MT373109 MW548428
23 S. huaningensis Huaning County, Yunnan XH3701 – AY854718 AY854775
24 S. huanjiangensis – GX0124 MT373103 MW548429
25 S. hugeibarbus Libo County, Guizhou* GZNU20150120005 MW014319
26 S. huizeensis Huize County, Yunnan hrfri2018046 MH982229
27 S. hyalinus Alugudong, Luxi County, Yunnan XH4701 – AY854721 AY854778
28 S. jii Gongcheng County, Guangxi YNUSJ201308060038 MF100765
29 S. jiuxuensis Jiuxu Town, Hechi City, Guangxi XH8501 – AY854736 AY854793
30 S. lateristriatus Maojiachong, Zhanyi County, Yunnan XH1102 – AY854703 AY854760
31 S. lingyunensis – – MW411665
32 S. longibarbatus Libo County, Guizhou* GZNU2019102022 NC_056194
33 S. longihornes Hongguo Town, Panzhou City, Guizhou* GZNU20210503016 – MZ634123 MZ634125
34 S. longshanensis Shupi Township, Qiubei County, Yunnan* S22 – PQ155085 PQ155093
35 S. macrocephalus Heilongtan, Shilin County, Yunnan XH0103 AY854683 AY854740 DQ845925
36 S. macrolepis Nandan County, Guangxi XH8201 AY854729 AY854786
37 S. macrophthalmus Xiaao, Duan County, Guangxi XH8401 AY854733 AY854790 HM536754 HM536835 HM536889
38 S. maculatus Weiwei Township, Yanshan County, Yunnan 8 – EU366193 EU366183
39 S. malacopterus Wulong Township, Shizong County, Yunnan* S43 – PQ155088 PQ155095
40 S. maitianheensis Jiuxiang,Yiliang County, Yunnan XH2301 AY854710 AY854767
41 S. mashanensis – GX0026–L18–12 MT373107 MW548430
42 S. microphthalmus Lingyun County, Guangxi NNNU201712001 MN145877
43 S. multipunctatus Huishui County, Guizhou – MG026730
44 S. oxycephalus Shilin County, Yunnan YNUSO20160610002 MG686610
45 S. purpureus Luoping County, Yunnan IHB:2006638 MW548264
46 S. punctatus – – MW014318
47 S. purpureus Zhonghe Ying Township, Kaiyuan, Yunnan* S20 – PQ155083 PQ155097
48 S. qiubeiensis Songming, Yunnan IHB:2006624 NC_063104
49 S. qiubeiensis Qiubei County, Yunnan* S21 – PQ155084 PQ155098
50 S. qujingensis Huize County, Yunnan hrfri2018044 MH937706
51 S. rhinocerous Luoping County, Yunnan – KR069119
52 S. ronganensis Rong’an County, Guangxi – KX778473
53 S. sanxiaensis Guojiaba Town, Zigui County, Hubei* KNHM 2019000001 OP745534
54 S. simengensis – – OQ718406
55 S. tingi Fuxian Lake, Yunnan YNUST201406180002 MG323567
56 S. wenshanensis Xigu Town, Wenshan, Yunnan YNUSW20160703016 MW553076
57 S. wenshanensis Dehou Town, Wenshan City, Yunnan* S45 – PQ155089 PQ155100
58 S. wumengshanensis Xuanwei County, Yunnan YNUSM20160817008 MG021442
59 S. xunlensis Huanjiang, Guangxi IHB:04050268 EU366187 EU366184 HM536752 HM536833 HM536887
60 S. xiejiahuai sp. nov. Hongguo Town, Panzhou City, Guizhou* S46 PQ165088
61 S. xingyiensis XingyiCity, Guizhou, China* – ON573218
62 S. xichouensis Xingjie Town, Xichou County, Yunnan* S37 – PQ155087 PQ155099
63 S. yangzongensis Yangzonghai Lake, Yunnan XH6101 AY854725 AY854782 DQ845926
64 S. yimenensis Yimen, Yunnan IHB:2006646 EU366191 EU366180
65 S. yishanensis Liujiang County, Guangxi – MK387704
66 S. zhenfengensis Zhenfeng County, Guizhou* GZNU20150112021 MW014317
67 S. zhenfengensis Zhenfeng County, Guizhou* S17 – PQ155082 PQ155096
68 S. tianlinensis Longping Township, Tianlin County, Guangxi* S10 – PQ155081 PQ155092
69 S. tianlinensis Longping Township, Tianlin County, Guangxi* GZNU20210531003 PQ214929
Outgroup
70 Carassius auratus – – AB111951
71 Cyprinus carpio – – JN105357
72 Schizothorax yunnanensis – – KR780749
73 Onychostoma simum – – KF021233
74 Barbus barbus – – AB238965
75 Puntius ticto – – AB238969
76 Neolissochilus hexagonolepis – – KU380329
77 Garra orientalis – – JX290078
78 Myxocyprinus asiaticus – – AY526869
79 Danio rerio – – KM244705

Phylogenetic reconstruction and divergence time estimation were performed in BEAST v. 2.4.7 (Bouckaert et al. 2014). In the absence of a reference fossil calibration, we refer to Wen et al. (2022) and Yang et al. (2016): (1) Sinocyclocheilus originated at 43.96 million years (Ma) (sigma = 2.8); (2) the most recent common ancestor of Sinocyclocheilus occurred at 32.13 Ma (sigma = 2.8); (3) the divergence of the S. angularis species group and the S. tingi + S. cyphotergous species groups occurred at ~ 26.3 Ma (sigma = 4.2). BEAST analyses were run for 40 million generations under an uncorrelated relaxed clock model and a Yule tree prior, sampled every 5000 generations. All calibrations were performed using a normal prior and monophyly. Convergence of the run parameters was checked using Tracer v. 1.7.1 (Rambaut et al. 2018) to ensure that the effective sample size of all parameters was greater than 200. A maximum clade credibility tree was generated using Treeannotator v. 2.4.1 (Bouckaert et al. 2014) by applying a burn-in of 25%. Uncorrected p-distances (1000 replicates) based on Cyt b gene were calculated in MEGA v. 7.0 (Kumar et al. 2016).

Results

Phylogenetic analyses, genetic divergence, and divergence time

The length of the aligned sequence was 15671 base pairs (bp), including 16S (1718 bp), 12S (954 bp), tRNAs (1587 bp), ATP6 (684 bp), ATP8 (165 bp), COI (1551 bp), COII (691 bp), COIII (786 bp), Cyt b (1142 bp), ND1 (975 bp), BD2 (1045 bp), ND3 (349 bp), ND4 (1381 bp), ND4L (297 bp), ND5 (1824 bp) and ND6 (522 bp). Information on the evolutionary models used for phylogenetic reconstruction is shown in Suppl. material 2.

The phylogenetic tree reconstructed using BEAST shows that the living Sinocyclocheilus can be divided into five major clades, Clades I–V, and is highly resolved (BPP = 1.00) (Fig. 2A). The phylogenetic relationship between the four clades is (Clade I+(Clade II+(Clade III + (Clade IV+ Clade V)))) (Fig. 2A). New species clustered in Clade V, close to S. lateristriatus, had a genetic distance of 1.9% at the level of the mitochondrial Cyt b (Suppl. material 3).

Figure 2. 

A time tree based on mitochondrial genes assessment B type species for five species groups. Species photos B1, B3, B4, and B6 from Shan et al. (2000), B2 from Zhang and Dai (1992), and B5 from Li (1992).

Divergence time analyses indicate that Sinocyclocheilus originated 40.22 Ma (95% highest probability density (HPD): 35.58–44.92 Ma), with its most recent common ancestor occurring at 34.83 (95%HPD: 30.87–38.8 Ma). Divergence of the remaining four clades (Clades II–IV) occurred in the Oligocene to Early Miocene, ~ 23.64–28.93 Ma (95% HPD: 19.39–32.92 Ma). The divergence of the new species from its close relatives occurred at the Pliocene/Pleistocene boundary at ~ 2.56 Ma (95% HPD: 0.87–4.89 Ma), which is older than the divergence of the other sister species (Fig. 2A).

Morphological analyses

A total of five principal component factors with eigenvalues greater than two were extracted based on principal component analysis of the morphometric data (Suppl. material 3). These together accounted for 94.48% of the total variance, with the first principal component (PC1) and second principal component (PC2) accounting for 32.98% and 25.84% of the total variance. In the scatter plot of PC1 versus PC2, the new species Sinocyclocheilus xiejiahuai sp. nov. was distinguishable from S. lateristritus and S. qujingensis on the PC1 axis (Fig. 3). Major morphometric characters loaded on the PC1 axis included body depth, anal-fin length, prepectoral length, pectoral-fin base length, caudal peduncle length, caudal peduncle depth, head width, snout length, eye diameter, interorbital width, distance between anterior nostrils, mouth width, rostral barbel length, and maxillary barbel length (Table 4).

Table 4.
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PCA loadings of five principal components extracted from 34 morphometric data of S. xiejiahuai sp. nov. and its related species.

PC1 PC2 PC3 PC4 PC5
Standard length 0.389 0.508 -0.188 -0.616 0.164
Body depth 0.645 0.419 -0.614 -0.089 0.031
Predorsal length 0.351 0.531 -0.586 0.406 -0.159
Dorsal-fin base length -0.476 0.765 -0.010 -0.267 0.106
Dorsal-fin length -0.303 0.877 -0.217 -0.020 0.004
Pre-anal length -0.383 0.026 -0.791 0.005 0.336
Anal-fin base length -0.473 0.259 -0.066 0.155 0.648
Anal-fin length -0.640 0.612 0.265 0.221 0.034
Prepectoral length -0.820 0.378 -0.159 -0.275 -0.134
Pectoral-fin base length 0.683 0.056 0.499 -0.489 -0.080
Pectoral-fin length -0.031 0.280 0.695 -0.075 -0.505
Prepelvic length -0.544 0.741 0.325 0.096 0.030
Pelvic-fin base length 0.588 -0.293 -0.060 0.043 -0.662
Pelvic-fin length -0.243 0.880 0.029 0.175 -0.363
Caudal peduncle length 0.602 -0.479 0.583 0.112 0.211
Caudal peduncle depth 0.725 0.363 -0.429 -0.356 -0.103
Head length 0.261 0.774 0.511 -0.039 0.180
Head depth 0.571 0.583 -0.409 -0.291 -0.219
Head width 0.727 0.353 -0.257 0.512 -0.004
Snout length 0.625 0.386 0.457 0.259 0.197
Eye diameter 0.643 0.702 0.060 -0.233 0.041
Interorbital width 0.755 0.532 0.057 -0.001 0.191
Prenostril length 0.852 0.336 0.008 0.063 0.348
Distance between posterior nostrils 0.561 -0.141 0.460 -0.393 0.278
Upper jaw length -0.535 0.436 0.524 0.176 0.084
Lower jaw length -0.562 0.649 0.151 0.248 0.072
Mouth width 0.731 0.415 -0.384 0.202 0.022
Rostral barbel length 0.639 0.369 0.403 0.477 -0.010
Maxillary barbel length 0.634 -0.022 0.719 0.192 -0.146
Distance from the pectoral-fin origin to the pelvic-fin origin 0.459 -0.352 -0.581 0.445 -0.073
Distance from the pelvic-fin origin to the anal-fin origin 0.456 -0.582 0.054 0.046 0.460
Eigenvalues 6.887 10.361 1.049 0.689 0.979
Percentage of total variance 32.981 25.837 17.032 7.820 6.814
Cumulative percentage 32.981 58.817 75.850 83.669 90.483
Figure 3. 

Plot of principal component analysis, scores of Sinocyclocheilus xiejiahuai sp. nov., S. lateristritus, and S. qujingensis based on morphometric data.

Morphological comparison

Based on morphology and phylogeny, the new species Sinocyclocheilus xiejiahuai sp. nov. was assigned to the S. tingi group, and a detailed morphological comparison is shown in Table 2.

Sinocyclocheilus xiejiahuai sp. nov. can be distinguished from the 24 species belonging to the S. angularis and S. microphthalmus groups by the absence of horn-like structures and indistinct elevation at the head-dorsal junction, pectoral fins tip not reaching to pelvic-fin origin (vs presence of horn-like structures and pectoral fins long and not reaching to pelvic-fin origin); from the five species belonging to the S. jii species group by with serrations along posterior margin of the last unbranched fin of the dorsal fin (vs absent) (Zhao et al. 2009), and from the 21 species belonging to the S. cyphotergous species group by pectoral fins tip not reaching to pelvic-fin origin (vs usually reaching to pelvic-fin origin).

For the 26 species of the S. tingi group, new species can be distinguished by a series of morphological characters. By lacking irregular markings on the body lateral, the new species can be distinguished from S. aluensis, S. angustiporus, S. bannaensis, S. grahami, S. guishanensis, S. huaningensis, S. huizeensis, S. lateristriatus, S. longshanensis, S. macrocephalus, S. maculatus, S. maitianheensis, S. malacopterus, S. oxycephalus, S. purpureus, S. robustus, S. wumengshanensis, S. xichouensis, S. tingi, S. yimenensis, and S. wenshanensis. New species differs from S. anophthalmus by eyes present (vs absent) and lateral line pores 74 (vs 52–56); from S. longifinus, S. qujingensis, and S. yangzongensis by six branched dorsal-fin rays (vs 7) and seven branched pelvic-fin rays (vs 8 or 9). The new species can be further distinguished from S. qujingensis and S. yangzongensis by 13 branched pectoral-fin rays (vs 16), and from S. longifinus by the tip of the pectoral fin not reaching to pelvic-fin origin (vs reaching to pelvic-fin origin).

For the four species not placed in any species group, new species differed from S. luolouensis by eye normal (vs eyes reduced) and pectoral fins tip not reaching to pelvic-fin origin (vs .beyond pelvic-fin origin) (Lan et al. 2013), from S. gracilis by having nine rakers on the first gill arch (vs 12) and body depth of 13% of standard length (vs 21.0–23.8%), from S. pingshanensis and S. gracilis by with serrations along posterior margin of the last unbranched fin of the dorsal fin (vs absent) and nine rakers on the first gill arch (vs 10–12), and from S. wui by three unbranched anal-fin rays (vs 2), 13 branched pectoral-fin rays (vs 14–15), and 17 branched caudal-fin rays (vs 14–15).

Taxonomic account

Sinocyclocheilus xiejiahuai Luo, Fan, Xiao & Zhou, sp. nov.

https://zoobank.org/B3636299-814D-4EE8-92F4-CA48078A7581
Fig. 4, Table 5

Material examined

Holotype. GZNU20230304001, total length 242.8 mm (TL), standard length 201.8 mm (SL), • Hongguo Town, Panzhou City, Guizhou Province, China; 25.6576°N, 104.4044°E; ca 1852 m a.s.l.; collected on October 2, 2021.

Diagnosis

Sinocyclocheilus xiejiahuai sp. nov. can be distinguished from all other congeners by the following combination of characters: (1) absence of horn-like structures and indistinct elevation at the head-dorsal junction; (2) absence of irregular black markings on the body lateral and scaleless; (3) eyes large, eye diameter 13% of head length; (4) dorsal-fin rays, iii, 6½, last unbranched ray strong, with serrations along posterior margin; (5) pectoral-fin rays, i, 13; (6) anal-fin rays, iii, 5; (7) pelvic-fin rays, i, 7; (8) lateral line pores 74; (9) gill rakers well developed, nine on first gill arch; (10) pectoral fins short, tip not reaching to pelvic-fin origin (Table 5).

Table 5.
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Morphological characteristics and measurements of Sinocyclocheilus xiejiahuai sp. nov., S. qujingensis, and S. lateristritus.

S. xiejiahuai sp. nov. (n = 1) lateristritus (n = 2) S. qujingensis (n = 6)
Range Range Mean ± SD Range Mean ± SD
Total length 240.0 73.2–87.5 80.8 ± 5.2 53.3–140.1 96.7 ± 61.4
Standard length 201.0 59.3–71.0 64.9 ± 4.1 43.3–98.1 70.7 ± 38.7
Body depth 56.7 13.0–18.4 15.3 ± 1.9 9.9–19.2 14.6 ± 6.6
Predorsal length 108.6 30.9–39.4 34.1 ± 3.0 22.3–50.0 36.1 ± 19.6
Dorsal-fin base length 24.7 8.5–9.6 9.2 ± 0.4 5.9–12.3 9.1 ± 4.5
Dorsal-fin length 40.0 12.7–16.7 15.2 ± 1.5 10.2–19.4 14.8 ± 6.4
Pre-anal length 140.4 42.0–49.2 45.5 ± 2.4 30.9–68.9 49.9 ± 26.9
Anal-fin base length 17.4 5.2–6.8 5.8 ± 0.5 4.0–8.4 6.2 ± 3.1
Anal-fin length 28.4 9.8–13.3 11.6 ± 1.2 7.3–16.3 11.8 ± 6.4
Prepectoral length 53.0 17.1–19.7 18.4 ± 0.9 12.7–27.2 19.9 ± 10.3
Pectoral-fin base length 8.9 2.1–3.1 2.5 ± 0.5 1.4–3.0 2.2 ± 1.2
Pectoral-fin length 32.9 8.9–16.0 13.0 ± 2.9 7.2–17.5 12.3 ± 7.3
Prepelvic length 98.1 29.9–36.0 33.3 ± 2.1 21.8–49.1 35.5 ± 19.3
Pelvic-fin base length 8.9 2.0–3.3 2.5 ± 0.5 1.8–4.2 3.0 ± 1.7
Pelvic-fin length 25.5 8.9–13.5 11.0 ± 1.6 6.5–13.7 10.1 ± 5.1
Caudal peduncle length 49.5 12.3–16.6 14.0 ± 1.6 8.4–22.6 15.5 ± 10.1
Caudal peduncle depth 24.7 6.4–8.5 7.1 ± 0.7 4.6–9.2 6.9 ± 3.2
Head length 57.1 16.4–21.0 19.2 ± 1.6 11.8–26.3 19.0 ± 10.3
Head depth 40.3 11.3–15.2 12.5 ± 1.4 8.1–16.1 12.1 ± 5.6
Head width 33.1 7.8–11.2 9.3 ± 1.2 5.6–13.6 9.6 ± 5.7
Snout length 21.0 4.7–6.8 6.3 ± 0.8 3.3–8.7 6.0 ± 3.8
Eye diameter 6.7 1.8–2.4 2.1 ± 0.2 1.2–2.4 1.8 ± 0.9
Interorbital width 18.6 4.4–6.4 5.5 ± 0.7 3.4–7.1 5.2 ± 2.6
Prenostril length 13.1 2.3–3.5 3.0 ± 0.4 1.7–3.5 2.6 ± 1.3
Distance between posterior nostrils 12.9 2.9–4.8 3.6 ± 0.7 2.4–4.8 3.6 ± 1.7
Upper jaw length 13.7 4.8–5.7 5.3 ± 0.4 3.2–7.6 5.4 ± 3.1
Lower jaw length 12.1 4.4–5.4 4.9 ± 0.4 3.0–6.8 4.9 ± 2.7
Mouth width 17.2 4.4–6.1 5.0 ± 0.7 3.1–6.8 5.0 ± 2.6
Rostral barbel length 24.6 4.8–9.8 7.1 ± 1.6 2.2–9.5 5.9 ± 5.2
Maxillary barbel length 30.2 5.3–12.0 8.0 ± 2.8 2.5–11.8 7.1 ± 6.6
Distance from the pectoral-fin origin to the pelvic-fin origin 42.2 11.4–14.2 12.7 ± 0.9 8.7–20.1 14.4 ± 8.0
Distance from the pelvic-fin origin to the anal-fin origin 38.4 9.5–10.8 9.9 ± 0.5 6.2–16.4 11.3 ± 7.2

Description

Body fusiform, moderately elongate and compressed. Dorsal profile convex from nape to dorsal-fin, greatest body depth at dorsal-fin insertion, ventral profile slightly concave, tapering gradually toward the caudal-fin, greatest body depth slightly anterior to dorsal-fin insertion.

Head short, compressed laterally, length longer than maximum head width, depth longer than maximum head width. Eyes present, eye diameter 13% of head length (HL), interorbital distance larger than distance between posterior nostrils. Snout short, U-shaped, and projecting beyond lower jaw in dorsal view, length 37% of HL. Mouth subterminal, with slightly projecting upper jaw. Two pairs of nostrils, anterior and posterior nostrils close set, nares at 2/3 between snout tip and anterior margin of eye, anterior nares possessing an anterior rim with a posterior fleshy flap forming a half-tube. Two pairs of barbels, rostral barbels short, not reaching the anterior edge of operculum when extended backwards, maxillary barbel slightly long compared to rostral barbel, beyond the anterior edge of operculum when extended backwards (Table 5).

Figure 4. 

Lateral view of adult holotype GZNU20230304001 of Sinocyclocheilus xiejiahuai sp. nov. in preservative A left side B right side.

Dorsal fin rays iii, 6½, pectoral fin rays i,13, pelvic fin rays i, 7, anal fin rays iii, 5, and 13 branched caudal fin rays. Dorsal fin long, 24% of SL, less than head length, distal margin truncated, origin posterior to pelvic fin insertion, situated slightly anterior to midpoint between snout tip and the caudal fin base, last unbranched ray strong, softening toward tip, with serrations along posterior margin, first branched ray longest, shorter than HL, tip reaching to the vertical of the anus. Pectoral fin developed, distal margin rounded, length slightly small than HL, 16% of SL, tips beyond 2/3 of the distance between pectoral-fin origin and pelvic-fin origin, tips not reaching to pelvic fin-origin. Pelvic fin moderately developed, distal margin rounded, length 14% of SL, and tips not reaching to anus. Anal fin short, 15% of SL, distal margin truncated, origin close to the anus, tips not reaching to caudal fin base. Caudal peduncle well developed, length 52.4 mm, depth 23.4 mm, and without adipose crests along both dorsal and ventral sides. Caudal fin slight forked, upper lobe equal in length to the lower one, tips rounded.

Body non-scale, lateral line pores 74. Complete lateral line, slightly curved, curved downward at the anus position, originating from posterior margin of operculum and extending to end of caudal peduncle.

Coloration

In 7% formalin solution, the specimen was grayish brown overall, with each fin pale yellow.

Geographical distribution and habitat

Sinocyclocheilus xiejiahuai sp. nov. is the only vertical cave found at an altitude of 2276 m in Hongguo Town, Panzhou City, Guizhou Province, China, some distance away. The discovery site is within the Beipanjiang River Basin (Fig. 1). There is no light in the cave. Individuals were distributed in a small pool ~ 25 m from the cave entrance. The pool is ~ 1.8 m wide and 80 cm deep, and the water temperature at the time of collection was ~ 16 °C and pH 7.4. Inside the cave, the species of S. xiejiahuai sp. nov. is symbiotic with S. longicornus (S. angularis group) and Triplophysa panzhouensis. The arable land outside the cave is mainly cultivated with maize, wheat, and potatoes.

Etymology

The specific name xiejiahuai is in honor of Professor Jia-Hua Xie (谢家骅), for his contribution to zoological research in China. Before retiring from Guizhou Normal University, he described S. angustiporus, the first species distributed in Guizhou within the S. tingi species group, and his work has been an important contribution to the study of zoology in Guizhou, especially the conservation of critically endangered species. We propose the common English name “Xie’s Golden-lined Fish” and the Chinese name “Xiè Shì Jīn Xiàn Bā (谢氏金线鲃).”

Discussion

Based on previous records, the genus Sinocyclocheilus (Fang, 1936) was recorded with 79 species (Zhao and Zhang 2009; Xu et al. 2023; Luo et al. 2023; Shao et al. 2024), all of which are endemic to southwestern China, and the description of the new species in this study increases it to 80 species. Up to now, there are 27 species in the tingi group, mainly distributed in eastern Yunnan and western Guizhou. Sinocyclocheilus xiejiahuai sp. nov. is the third species of the S. tingi species group discovered in Guizhou with S. angustiporus and S. robutus. Although the description of this new species is based on a single specimen and some measurements may not be sufficient, the fin characteristics (see morphological comparisons above) and genetic differences support the validity of this new species. The new species is phylogenetically close to S. lateristriatus and S. qujingensis (Fig. 2), with genetic distances of 1.9% and 3.1% (Suppl. material 3), which was greater than that between sister species of the same genus, e.g., 1.2% between S. yimenensis and S. huizeensis (Suppl. material 2). The new species co-inhabits a cave with S. longicornus, and is the first report of the sympatric distribution of the S. tingi and S. angularis groups. Furthermore, our phylogenetic tree suggests that S. longshanensis should be assigned to the S. microphthalmus species group.

Our reconstructed divergence times are similar to those of Wen et al. (2022) who discussed the origin and early diversification of the genus Sinocyclocheilus. Time-tree-based results suggest that Sinocyclocheilus originated in the late Eocene, with its most recent common ancestor/Clade I occurring at 34.83 Ma, and that divergence of the remaining clades was centered in the Oligocene to early Miocene, ca 19.39–32.92 Ma (Fig. 2). The origin and early divergence events are well-coupled with the rapid uplift of the Qinghai-Tibetan Plateau during the Oligocene-middle Miocene (Ding et al. 2022). Middle Miocene Climate Optimum (17–14 Ma), the monsoon climate brings precipitation to promote the development of caves in the karst region (Farnsworth et al. 2019), which increases the ecological opportunities for the formation of cave fishes within the Sinocyclocheilus. We also observed that Sinocyclocheilus xiejiahuai sp. nov., S. lateristriatus, and S. qujingensis are distributed in the Nanpanjiang River basin in close phylogenetic and geographic proximity, and this congruence may indicate that the formation of these species, and even of species in the S. tingi group, is related to historical drainage changes resulting from the uplift of the Yunnan-Guizhou Plateau since the Late Miocene (Zhang et al. 2020). Thus, geographic isolation from historical drainage changes has shaped the formation of species diversity in the S. tingi species group (Mao et al. 2021,2022; Zhao and Zhang 2009; Wen et al. 2022).

This new species is presently restricted to its type locality. Given that its habitat borders the urban area of Panzhou, which is experiencing rapid urbanization, there is a significant risk of habitat disturbance and destruction in the near future. In the last five years, we have conducted a total of 16 field surveys at the type locality, and no new individuals have been detected except for the first one, suggesting that the population size of this species is very small. The Chinese government listed all species of Sinocyclocheilus endemic to China as second-class of the national protected animals on 5 February 2021 (National Forestry and Grassland Administration & National Park Administration, 2021). Therefore, the new species has strict legal protection in China, and the local government should strengthen publicity about the protection of this species to avoid it being caught and trafficked. In addition to the small population size, the following threats to the habitat of the new species include declining water levels in caves, pesticide use, domestic waste, and as potential land for urban construction. Therefore, we suggest the species may be eligible for listing as Endangered (B1ab (i, ii, iii) + 2ab (i, ii, iii)) in the IUCN Red List of Threatened Species.

Acknowledgments

We thank Ya-Li Wang, Xing-Liang Wang, and other for help with specimen collection. We thank Dr. Min Rui of the Kunming Institute of Zoology, Yunnan, China, for his assistance in the examination of specimens. We thank researcher Hongfu Yang for providing tissue samples of some species for sequencing. We thank LetPub (www.letpub.com) for its linguistic assistance during the preparation 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

This research was supported by the programs of the Vertebrate Diversity in the Mountains of Southwest China (XDB31000000).

Author contributions

Jiang Zhou and Tao Luo conceived and designed the research; Cui Fan, Man Wang, Jia-Jun Zhou, and Tao Luo, conducted field surveys and collected samples; Tao Luo and Cui Fan performed molecular work; Cui Fan, Man Wang, and Ning Xiao processed the English language of the manuscript; Jiang Zhou provided financial support. All authors read and approved the final version of the manuscript.

Author ORCIDs

Cui Fan https://orcid.org/0009-0002-8039-649X

Man Wang https://orcid.org/0000-0001-6201-1811

Jia-Jia Wang https://orcid.org/0009-0007-8637-5469

Tao Luo https://orcid.org/0000-0003-4186-1192

Jia-Jun Zhou https://orcid.org/0000-0003-1038-1540

Jiang Zhou https://orcid.org/0000-0003-1560-8759

Data availability

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

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1Cui Fan and Man Wang contributed equally to this work.

Supplementary materials

Supplementary material 1 

Morphometric data for nine S. tingi group species

Cui Fan, Man Wang, Tao Luo, Jia-Jun Zhou, Ning Xiao, Jiang Zhou

Data type: docx

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.
Download file (26.58 kb)
Supplementary material 2 

The best model obtained using PartitionFinder v. 2.1.1 evaluated under the Bayesian information criterion

Cui Fan, Man Wang, Tao Luo, Jia-Jun Zhou, Ning Xiao, Jiang Zhou

Data type: docx

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.
Download file (11.96 kb)
Supplementary material 3 

Uncorrected p-distance (%) between 69 species of the genus Sinocyclocheilus based on mitochondrial genes

Cui Fan, Man Wang, Tao Luo, Jia-Jun Zhou, Ning Xiao, Jiang Zhou

Data type: xlsx

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.
Download file (32.94 kb)
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