Zoosyst. Evol. 100 (2) 2024, 583–595 | DOI 10.3897/zse.100.

120676

Tachysurus wuyueensis (Teleostei, Bagridae), a new species of

catfish from the Qiantang-Jiang basin, southeast China
Jia-Jun Zhou1, Le-Yang Yuan2, Wei-Han Shao3
1 Zhejiang Forest Resource Monitoring Center, Hangzhou, China
2 Zhejiang Museum of National History, Hangzhou, China
3 Institute of Hydrobiology, Chinese Academy of Sciences, Wuhan, China

https://zoobank.org/3D10FE75-FC05-4A93-A03A-17C77BD60999

Corresponding author: Wei-Han Shao ([email protected])

Academic editor: Nicolas Hubert  ♦  Received 10 February 2024  ♦  Accepted 28 March 2024  ♦  Published 17 May 2024

Abstract
Tachysurus wuyueensis, new species, is described from the Qiantang-Jiang Basin, situated in Suichang County, Zhejiang Province and
Xiuning County, Anhui Province, southeast China. The coastal basin drains into the East China Sea. The new species belongs to the
T. pratti-T. truncatus group within the genus Tachysurus defined by having a smooth anterior margin of the pectoral-fin spine, short max-
illary barbels not extending beyond the base of the pectoral-fin spine, short dorsal spine not exceeding two thirds of head length and an
emarginated caudal fin. This new species is distinct from all other species of this group, Tachysurus pratti (Gunther, 1892), T. truncatus
(Regan 1913), T. gracilis (Li, Chen & Chan, 2005) and T. brachyrhabdion (Cheng, Ishihara & Zhang, 2008), in having a shorter prepelvic
body (length 40.0–46.4% of SL vs. 45.8–54.8%). It further differs from T. pratti, T. truncatus and T. gracilis in having more vertebrae
(45–47 vs. 37–44) and more anal fin rays (21–25 vs. 14–20), from T. brachyrhabdion in having a more slender body (depth 10.1–13.5% of
HL vs. 13.1–17.6%). Molecular phylogeny, based on the mitochondrial cytochrome b (cyt. b) gene confirms the validity of T. wuyueensis
and the T. pratti-T. truncatus group. Furthermore, this study addresses the diagnostic traits distinguishing the T. pratti-T. truncatus group
from the T. tenuis-T. crassilabris group which have historically been treated as a single species group due to morphological similarities.

Key Words
Caudal fin shape, new taxon, stream-dewelling species, taxonomy

Introduction example, Tachysurus ondon (Shaw 1930), T. adiposa-

lis (Oshima 1919), T. albomarginatus (Rendahl 1928),
The genus Tachysurus Lacépède 1803 is a group of T. trilineatus (Zheng 1979) and T. tenuis (Günther 1873).
East Asian endemic catfish that is widely distributed Unlike its congeners, species of Tachysurus always pos-
throughout most of the East Asian continent (Ku et al. sess elongated bodies and rounded, truncated or slightly
2007; Watanabe 2010; Shao and Zhang 2023). It is also emarginated (round-tailed) caudal fins regarded herein
one of the most diversified catfish genus in China con- as key adaptions for the torrent environments (Gosline
taining more than thirty species with strong differenti- 1997; Krishnadas et al. 2018). Moreover, the absence
ation in ecological niches (Shao et al. 2021; Shao and of diagnosable characters and ambiguous descriptions
Zhang 2023). South China, which is characterised by have obscured the species boundaries of these species
high mountains, deep valleys and abundant precipita- (Ferraris 2007; Shao et al. 2021). For instance, T. albo-
tion (López-Pujol et al. 2011; Wang et al. 2018; Sun marginatus has long been synonymised under T. tenuis
et al. 2021), harbours numerous Tachysurus species re- (Zheng & Dai, 1999) until Cheng et al. (2021) resur-
stricted to montane streams with fast-flowing high-ox- rected the former species, based on morphometric and
ygen waters and predominance of rocky substrate, for osteological analyses.

Copyright Jia-Jun, Z. et al. 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.
584 Jia-Jun, Z. et al.: A new Tachysurus species from the Qiantang-Jiang basin, China

In addition, rheophilic (fast-water) fishes are always T. albomarginatus, T. adiposalis and T. tenuis, were
confined to a narrow distribution which limits their dis- morphologically examined in this study, which came
persal ability and contributes to their habitat fragmen- from the following collections: Museum of Aquatic Or-
tation (Lima et al. 2017). Previous works have high- ganisms of Institute of Hydrobiology, Chinese Academy
lighted the underestimated diversity of Tachysurus, of Sciences, Wuhan (IHB), the Natural History Muse-
v.gr., T. albomarginatus that may be an assemblage of um, London (BMNH) and French Museum of Natural
five species (Cheng et al. 2021; Shao et al. 2021). A History, Paris (FMNH).
similar scenario occurs in T. adiposalis, originally de- Measurements were taken point to point with digital
scribed by Oshima (1919) on a single specimen of 170 calipers linked to a data recording computer and data
mm SL (standard length) collected from the Tamusui were recorded to the nearest 0.1 mm and made on the
River in Taiwan. T. adiposalis has been treated as wide- left side of each individual whenever possible, following
spread species, with a distribution including Taiwan the methods for Cheng et al. (2008). The head length and
and river basins from the southern region of Chinese measurements of other parts of the body are estimated as
mainland, such as the mid-lower Yangtze, Pearl Riv- percentages of the standard length (SL). Subunits of head
er and Qiantang River (= Qiantang-Jiang in Chinese) are provided as percentages of the head length (HL). The
(Zheng and Dai 1999). The key characters useful in dif- number of rays from the dorsal- and anal fins were count-
ferentiating Tachysurus species are absent or obscured ed using the method by Watanabe (1995). Other fin rays
in the original description of T. adiposalis, which re- were counted under a binocular dissecting microscope
sulted in the taxonomic confusion. Recent studies have utilising transmitted light. Vertebral count was taken
suggested that specimens from the Xi-Jiang (the Pearl from X-ray photographs, with the anterior five vertebrae,
River Basin) and the Yuan-Jiang (the middle Yangtze namely the Weberian complex, not counted.
River Basin) belonging to T. adiposalis s.l. were de- Morphometric measurements were subject to principal
termined as Tachysurus gracilis (Li, Chen & Chan, component analysis (PCA) in order to examine external
2005) and Tachysurus brachyrhabdion (Cheng, Ishi- morphological differentiation and determine the relative
hara & Zhang, 2008). However, the taxonomic status contributions of specific variables to morphological dif-
of specimens from the Qiantang-Jiang which were also ferences in the target species. PCA was run with SPSS
recognised as Tachysurus adiposalis has not been men- 16 (SPSS, Chicago, IL, USA). Prior to the analysis, all
tioned in previous studies and it potentially represents morphometric measurements, except standard length,
an undescribed species. were normalised following the method of Reist (1985)
A fish field survey conducted by the authors in the to eliminate the influence of allometry of body parts and
Qiantang-Jiang Basin of Zhejiang and Anhui Provinces sample size.
yielded 14 specimens initially identified as T. adiposis. Phylogenetic analysis was performed on cyt. b and
Careful morphological examination revealed that these the sequences uploaded to NCBI GenBank (Table 1).
were, in fact, not conspecific with any other known spe- The thirty-eight cyt. b gene sequences, here amplified
cies of Tachysurus and represent a new species. The from twenty-five species of Tachysurus, were used for
purpose of the present paper is to provide a formal de- molecular phylogenetic analysis. Tachysurus trilineatus
scription of this unnamed species, based on multiple was used as the outgroup which is the basal lineage of
lines of evidence containing morphological and phylo- Tachysurus genus (Ku et al. 2007). The sequences were
genetic datasets. revised manually and then aligned using ClustalW in
MEGA7 (Kumar et al. 2016). Both Maximum Likeli-
hood (ML) and Bayesian Inference (BI) methods were
Materials and methods utilised to reconstruct the phylogenetic relationships. The
optimal nucleotide substitution model was selected by
Fourteen specimens of the new species were caught ModelFinder (Kalyaanamoorthy et al. 2017) according to
during two field surveys conducted, respectively, in the Akaike Information Criterion. Maximum Likelihood
September 2020 and May 2023 into the Qiantang-Ji- analysis was run by IQ-tree (Nguyen et al. 2015), with
ang flowing into the East China Sea. Amongst them, six the selected TIM3+F+I+G4 model and 1,000 non-para-
specimens were fixed in 10% formalin after removal of metric bootstrap replicates. Bayesian Inference was per-
right-side pelvic-fin clips. These fin clips were stored formed in MrBayes (Ronquist et al. 2012) under the se-
in 95% ethyl alcohol and utilised for molecular analy- lected GTR+F+I+G4 model. Two independent runs were
sis. The remaining caught specimens were directly pre- carried out with four Monte Carlo Markov chains (three
served in 10% formalin preservative for morphological hot chains and one cold chain) for 20 million generations
examination. Their voucher specimens are deposited in to calculate posterior probability. Trees were sampled ev-
the Museum of Aquatic Organisms of the Institute of ery 1000 generations. The initial 25% of sampled trees
Hydrobiology (IHB), Chinese Academy of Sciences, were discarded as burn-in. Convergence of the runs was
Wuhan. Eight species, including T. gracilis, T. trun- assessed by the average standard deviation of split fre-
catus, T. pratti, T. brachyrhabdion, T. crassilabris, quencies (< 0.01).

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Table 1. GenBank accession numbers for molecular phylogenetic analysis.

Taxon Locality Distribution Accession number

Ingroup
(1) Tachysurus albomarginatus Zhejiang, China Qiantang-Jiang PP266663
(2) Tachysurus analis Jiangxi, China Gan-Jiang of lower Yangtze River PP266668
(3) Tachysurus argentivittatus Jiangxi, China Gan-Jiang of lower Yangtze River PP266678
(4) Tachysurus brachyrhabdion 1 Guizhou, China Yuan-Jiang of middle Yangtze River PP266650
Tachysurus brachyrhabdion 2 Guizhou, China PP266651
Tachysurus brachyrhabdion 3 Hunan, China PP266652
(5) Tachysurus crassilabris Hunan, China Xiang-Jiang of middle Yangtze River PP266665
(6) Tachysurus dumerili Jiangsu, China Lower Yangtze River PP266661
(7) Tachysurus gracilis1 Guangxi, China Xiang-Jiang of middle Yangtze River PP266654
(8) Tachysurus gracilis 2 Hunan, China Lijiang River of Pearl River PP266655
(9) Tachysurus intermedius Hainan, China Nandu-Jiang PP266676
(10) Tachysurus kyphus Guangxi,China Fangcheng- Jiang PP266671
(11) Tachysurus lani Guangxi, China Gui-Jiang of the Pearl River PP266662
(12) Tachysurus longispinalis Vietnam Red River PP266672
(13) Tachysurus nitidus Hunan, China Xiang-Jiang of middle Yangtze River PP266660
(14) Tachysurus ondon Zhejiang, China Ou-Jiang PP266677
(15) Tachysurus pratti 1 Sichuan, China Jinsha-Jiang of upper Yangtze River PP266656
Tachysurus pratti 2 Yunan, China PP266657
(16) Tachysurus similis Fujian, China Min-Jiang PP266664
(17) Tachysurus sinensis Hubei, China Middle Yangtze River PP266674
(18) Tachysurus tenuis Zhejiang, China PP266666
(19) Tachysurus truncatus 1 Sichuan, China Upper Yangtze River PP266658
Tachysurus truncatus 2 Shaanxi, China Han-Jiang of middle Yangtze River PP266659
(20) Tachysurus ussuriensis Heilongjiang, China Heilong-Jiang PP266669
(21) Tachysurus vachelli Hubei, China Middle Yangtze River PP266670
(22) Tachysurus virgatus Hainan, China Jiajihe River PP266673
(23) Tachysurus wuyueensis (SUIC55690) Zhejiang, China Qiantang-Jiang PP266644
Tachysurus wuyueensis (SUIC55691) PP266645
Tachysurus wuyueensis (SUIC55692) PP266646
Tachysurus wuyueensis (XIUN65294) Anhui, China PP266647
Tachysurus wuyueensis (XIUN65295) PP266648
Tachysurus wuyueensis (XIUN65296) PP266649
(24) Tachysurus zhangfei Hunan, China Yuan-Jiang of middle Yangtze River PP266667
Outgroup
(25) Tachysurus trilineatus Guangdong, China Dong-Jiang of Pearl River PP266679

The genetic distances, based on cyt. B, were computed and the above all are sister to another species pair: T. trun-
in MEGA 7 using the Kimura-2-parameter (K2P) model catus and T. pratti. Herein, these five species are designated
(Kimura 1980). as belonging to the T. truncatus-T. pratti group given their
affinities in both morphology and molecular phylogeny.
The estimated K2P genetic distances of the cyt. b gene
Results between T. wuyueensis and congenerics range from 6.9% to
12.6% (Table 2). The distances between T. wuyueensis and
Phylogenetic analysis members of the T. pratti-T. truncatus group are in a range
of 6.9–8.3%. The paired species, T. truncatus and T. pratti,
A total of 1092 bps were included in the aligned dataset of endemic to the upper Yangtze River Basin have a 1.4% ge-
the cyt. b gene, with 668 conservative sites, 492 variable netic distance, while another paired species, T. brachyrhab-
sites, 375 Parsim-informative sites and 117 singleton sites. dion and T. gracilis, displays a 4.1% distance. The range of
The mean frequency of the four nucleotides in the sequenc- intraspecific genetic distances within T. wuyueensis is 0.1%.
es of Tachysurus wuyueensis are A = 29.1%, G = 15.6%,
C = 27.7% and T = 28.5%. The phylogenetic trees recon- Table 2. K2P distances (%) for species within the Tachysurus
structed by ML and BI methods are identical in topology pratti-Tachysurus truncatus group, based on the cyt. b gene.
(Fig. 1). A monophyly formed by samples of Tachysurus 1 2 3 4
wuyueensis was recovered with 100% posterior probabili- 1. T. wuyueensis sp. nov.
ties (pp) and 1.00 bs in ML and BI trees, respectively and be- 2. T. pratti 7.1
longed to a clade containing T. gracilis, T. brachyrhabdion, 3. T. truncatus 6.9 1.4
T. truncatus and T. pratti. Tachysurus wuyueensis is sister to 4. T. gracilis 8.3 9.1 8.6
a species pair consisting of T. gracilis and T. brachyrhabdion 5. T. brachydhrbdion 7.5 8.3 8.0 4.1

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586 Jia-Jun, Z. et al.: A new Tachysurus species from the Qiantang-Jiang basin, China

Figure 1. Phylogenetic tree of Tachysurus species inferred from cyt. b using Bayesian Inference and Maximum Likelihood methods.
Bayesian posterior probabilities (> 0.8) and Maximum Likelihood bootstrap values (> 70%) are shown, respectively.

Principal component analysis Table 3. Loadings on the first three principal components ex-
tracted from morphometric data for T. wuyueensis, T. truncatus,
The T. truncatus-T. pratti group can be differentiated from T. brachyrhabdion and T. gracilis.
all other Tachysurus species by having a smooth anterior
PC1 PC2 PC3
margin of the pectoral spine, short dorsal spine not ex-
Standard length 21.2 18.5 7.8
ceeding two-thirds of head length and short maxillary bar-
Body depth at anus 21.3 5.6 26.4
bels not extending beyond the base of the pectoral spine. Predorsal length 19.4 7.0 6.7
Amongst these five species, excluding T. pratti which Pre-anal length 21.1 9.6 9.1
has a deeply forked caudal fin (Fig. 2), the other four round- Prepelvic length 20.7 9.3 7.5
tailed species were included in the principal component Prepectoral length 18.0 3.6 14.0
analysis performed on the variance-covariance matrix of Length of dorsal-fin spine 15.8 14.2 8.5
log-transformed measurements (Table 3; Fig. 3). The re- Length of dorsal-fin base 20.3 2.2 6.8
Length of pectoral-fin spine 16.1 17.2 1.7
sults show that the combination of PC1 against PC3 and
Length of anal-fin base 19.6 41.2 29.2
PC2 against PC3 enabled the separation of T. wuyueen- Height of adipose fin 16.5 13.7 20.2
sis from T. truncatus, T. gracilis and T. brachyrhabdion. Adipose to caudal distance 23.9 19.1 13.8
Characters with main loadings in PC1 were the adipose to Length of caudal peduncle 24.7 21.9 24.2
caudal distance and body depth at anus; in PC2, they were Depth of caudal peduncle 19.7 4.2 23.5
the outer mandibular barbel length, anal-fin base length, Head length at latera 19.0 1.2 5.5
nasal-barbel length and maxillary barbel length; and, in Head depth 18.3 0.08 12.2
Head width 19.8 3.0 18.0
PC3, they were the inner mandibular barbel length, eye
Snout length 18.5 0.9 14.7
diameter and interorbital width (Table 3).
Interorbital width 20.2 8.8 30.2
Eye diameter 12.9 22.6 36.2
Mouth width 20.4 7.7 12.2
Morphological comparisons Length of nasal barbel 21.8 37.6 23.4
Length of maxillary barbel 22.6 36.3 3.8
Amongst the characters displaying main loading in PCA, Length of inner mandibular barbel 21.4 24.7 45.9
Length of outer mandibular barbel 22.7 46.2 19.5
some exhibit stable variations between species and can

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Zoosyst. Evol. 100 (2) 2024, 583–595 587

Figure 2. Lateral view of T. pratti (A) for IHB201909013758, 114.2 mm SL, Huili County, Sichuan Province, T. truncatus (B)
for IHB201812028889, 79.5 mm SL, Qianwei County, Sichuan Province, T. brachyrhabdion (C) for IHB 2017090539, 120.8 mm
SL, Songtao County, Guizhou Province, T. gracilis (D) for IHB 201803023401, 98.2 mm SL, Guanyang County, Guangxi Zhuang
Autonomous Region.

be used to distinguish between them (Fig. 4). Tachysurus ure 4B). It further differs from T. truncatus in having a
wuyueensis is distinct from the other three species in shorter pre-anal length (53.0–60.1% vs. 61.0–67.7%;Fig.
having a shorter prepelvic body (length 40.0–46.4% of 4C), longer anal-fin base (length 25.4–31.1% SL vs.
HL vs. 45.8–54.8%; see Fig. 4A), from T. truncatus and 19.1–25.5%, Fig. 4D), narrower mouth (width 8.1–10.6%
T. gracilis in having more anal-fin rays (21–25 vs. 17–20) SL vs. 10.0–13.7%; Fig. 4E), narrower interorbital space
and vertebrae (45–47 vs. 37–43) (Fig. 5), from T. trun- (width 5.0–7.9% SL vs. 8.7–10.5%; Fig. 4F) and from
catus and T. brachyrhabdion in having a more slender T. gracilis in having longer inner mandibular barbels
body (depth 10.1–13.5% of HL vs. 13.1–20.6%; see Fig- (length 4.9–6.0% of SL vs. 3.2–5.0%; Fig. 4G).

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588 Jia-Jun, Z. et al.: A new Tachysurus species from the Qiantang-Jiang basin, China

A B

Figure 3. Scatter plot of A. PC2 against PC3 and B. PC1 against PC2 extracted from morphometric data for T. wuyueensis, T. trun-
catus, T. brachyrhabdion and T. gracilis

Figure 4. Relationship between A. Prepelvic length and SL for T. wuyueensis, T. truncatus, T. brachyrhabdion and T. gracilis; rela-
tionship between; B. Body depth and SL for T. wuyueensis, T. truncatus and T. brachyrhabdion; relationship between; C. Pre-anal
length and SL; D. Anal fin base length and SL; E. Mouth width and SL; F. Interorbital width and SL for T. wuyueensis and T. trun-
catus; relationship between; G. Inner mandibular barbel length and SL for T. wuyueensis and T. gracilis.

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Figure 5. Meristic counts difference amongst T. wuyueensis, T. truncatus, T. brachyrhabdion and T. gracilis in vertebrae number
(A) and anal fin ray number (B).

Tachysurus wuyueensis Zhou, Yuan & Shao, sp. nov. Table 4. Morphomertric data for Tachysurus wuyueensis spe-
https://zoobank.org/18A16B30-C113-4285-8A76-058999C3D0E1 cies nova.
Fig. 6
Holotype Paratypes (n = 13)
Range Mean ± SD
Pseudobagrus adiposis: Mao 1991: 169 (coastal rivers in Zhejiang Standard length 78.7 61.8–115.1 69.7 ± 14.3
Province). %SL
Body depth at anus 13.2 10.1–13.5 13.1 ± 1.5
Type materials. Holotype. IHB 202009055690, 70.7 mm Predorsal length 32.6 27.2–35.3 32.5 ± 2.8
SL, South China: Zhejiang Prov.: Suichang County: the Pre-anal length 55.8 53.0–60.1 57.6 ± 1.9
Prepelvic length 44.1 40.0–46.6 44.0 ± 1.4
Qiantang-Jiang at Jiulongshan National Natural Reserve
Prepectoral length 20.9 16.1–23.1 20.8 ± 2.1
(28°23'38"N, 118°53'41"E) (Fig. 7); collected by Zhi- Length of dorsal-fin spine 59.2 46.4–63.8 59.2 ± 3.3
Gang Xie in Sept 2020. Length of dorsal-fin base 11.4 8.6–11.7 11.3 ± 1.3
Paratypes. IHB 202009055691-2, two ex., Length of pectoral-fin spine 14.7 10.7–15.5 14.7 ± 2.2
63.8–65.2 mm SL; other data same as holotype. Length of anal-fin base 26.4 25.4–30.8 27.0 ± 1.6
IHB202212165294-304, 11 ex., 61.8–123.9 mm SL, south Height of adipose fin 3.1 2.4–5.1 4.1 ± 0.9
China: Anhui Prov.: Xiuning County: the Qiantang-Jiang Adipose to caudal distance 13.9 13.2–16.8 15.3 ± 2.1
Length of caudal peduncle 16.4 16.0–18.1 17.3 ± 0.9
at Liukou Township (29°48'57"N, 117°53'11"E) (Fig. 7);
Depth of caudal peduncle 7.9 6.5–8.0 7.6 ± 0.3
collected by Wei-Han Shao in May 2023. Head length at latera 22.8 17.9–24.8 23.7 ± 2.5
Diagnosis. Distinguished from its congeners by the Head depth 11.7 9.0–13.2 12.3 ± 1.5
following combination of characters: a smooth anterior Head width 14.2 13.1–15.7 15.2 ± 1.1
margin of the pectoral-fin spine, short maxillary barbels Snout length 5.5 4.2–8.7 6.0 ± 1.9
not extending beyond the base of the pectoral-fin spine, Interorbital width 6.8 5.0–8.4 7.7 ± 1.3
short dorsal spine not exceeding two-thirds of head Eye diameter 4.5 4.3–5.6 4.9 ± 0.3
Mouth width 8.6 8.1–10.4 10.1 ± 1.0
length, prepelvic length 40.0–46.4% SL, 45–47 verte-
Length of nasal barbel 6.5 5.2–7.5 7.1 ± 1.4
brae, 21–25 anal-fin rays, body depth 10.1–13.5% SL, a
Length of maxillary barbel 14.5 8.6–13.7 13.1 ± 3.8
slightly emarginated caudal fin. Length of inner mandibular barbel 6.0 4.6–6.6 6.1 ± 0.7
Description. Morphometric measurements taken Length of outer mandibular barbel 9.5 5.9–10.7 9.6 ± 2.1
from the holotype (78.7 mm SL) and 13 paratypes (61.8–
123.9 mm SL) summarised in Table 4.
Body elongated, anteriorly cylindrical and slightly Head depressed, broad, and covered with thin skin.
compressed posteriorly. Dorsal profile rising gradually Supra-occipital process slender, with evenly converg-
from snout tip to dorsal-fin origin, then sloping evenly ing sides and pointed tip, separated from nuchal plate
from there to posterior end of adipose-fin base, and grad- by a broad interspace. Snout slightly pointed in dorsal
ually increasing to dorsal origin of procurrent caudal-fin view and obtuse or blunt in lateral view, longer than
rays. Ventral surface of head flattened; ventral profile of eye diameter. Interorbital space moderately space wide
body straight or slightly rounded from head to anal-fin and slightly flattish. Eyes moderately large, ellipti-
origin, decreasing evenly from posterior end of anal-fin cal, covered with thick membrane and anterolateral in
base to origin of ventral procurrent caudal-fin rays. Lat- head, visible when viewed dorsally, but not ventrally,
eral line complete, straight and mid-lateral in position. with slightly convex and comparatively narrow inter-
Vertebrae 5 + 45 (46, 47). orbital space.

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590 Jia-Jun, Z. et al.: A new Tachysurus species from the Qiantang-Jiang basin, China

Figure 6. Lateral (A), dorsal (B) and ventral (C) views of Tachysurus wuyueensis species nova, IHB202009055690, holotype,
70.7 mm SL.
Mouth subterminal, transverse. Upper jaw anteriorly rooted posterolateral to inner mandibular barbel, just ex-
protruded, longer than lower jaw in length; interorbit- tending to posterior margin of eye.
al space narrower than mouth opening. Teeth villiform, Dorsal fin with a spinelet, one spine and seven soft
in irregular rows on all tooth-bearing surfaces. Premax- branched rays. Dorsal-fin origin equidistant to pecto-
illary tooth plates broad, of equal width throughout. ral-fin insertion and ventrally to pelvic-fin insertion, also
Dentary tooth plates arched, broadest at symphysis and equidistant to anal-fin origin and snout tip. Spinelet flat-
narrowing laterally, of same width at symphysis as pre- tened, with long blunt distal tip. Dorsal-fin spine slender,
maxillary tooth plates. Vomerine tooth plate unpaired, with smooth anterior margin and slightly serrated distal
continuous across mid-line, slightly curved anteriorly posterior margin, equal to or slightly longer than pecto-
and much narrower than premaxillary plate. Gill open- ral-fin spine. First dorsal-fin soft ray longest, surpass-
ing wide, extending from the post-temporal region to ing tip of last ray. Distal margin of dorsal-fin rays nearly
beyond isthmus. straight. Nuchal plate triangular, with anterior pointed
Barbels in four pairs; nasal barbels small, thread-like, tip anteriorly.
not reaching beyond posterior margin of eye; maxillary Adipose fin inserted slightly behind vertical through
barbels slender, slightly exceeding posterior margin of pectoral-fin origin, with convex distal margin along entire
eye; mandibular barbels in two pairs, thick, short, inner length and deeply incised posterior part to form round-
barbels positioned in transverse row at level of posterior ed apex. Adipose fin base moderately long, equal to or
naris, extending beyond mid-point of eye, outer barbels slightly longer than anal fin base length.

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Figure 7. Map showing distributions of five species of Tachysurus in China: T. wuyueensis species nova, T. pratti, T. truncatus,
T. brachyrhabdion and T. gracilis.

Pectoral fin with one spine and seven (or eight) soft caudal peduncle) (Fig. 6) and fading to a more uniform
branched rays, inserted slightly anterior or at level of yellow or brown in larger individuals (Fig. 8). Adipose fin
posteriormost point of opercle, not reaching halfway to brownish, with a yellow anterodorsal margin and a slight-
pelvic-fin insertion. Pectoral-fin spine very stout, sharp- ly transparent posterodorsal margin. Dorsal fin trans-
ly pointed at tip, equidistant to or slightly longer than parent for anterior two-thirds, rest of fin greyish-brown.
dorsal-fin spine, with a smooth anterior margin and 8 Caudal fin greyish in posterior third, rest of fin transpar-
(9, mean 8.3) strong serrations along posterior margin. ent with a slightly white margin in smaller individuals.
Cleithral process triangular with a sharp pointed tip, ex- Pectoral, pelvic and anal fins transparent.
tending for half of pectoral fin spine length.
Pelvic fin with one unbranched and five branched soft
rays, inserted closer to tip of snout than to posterior end
of anal fin base, closer to depressed tip of dorsal fin than
to anterior end of anal fin base. Tip of depressed pelvic fin
reaching or slightly extending beyond anal fin origin. Pel-
vic fin distal margin convex. Anus and urogenital opening
nearer to anal fin origin than to posterior end of pelvic fin
base. Males with a conical genital papilla not reaching
base of first anal fin soft ray.
Anal fin long, with 21 (22, 23, 25) branched rays; ad-
ipose fin posterior margin away from caudal fin. Anal fin
origin to caudal fin than to tip of snout. Distal margin of
anal fin convex; anterior rays shortest. Figure 8. A. Habitat of Tachysurus wuyueensis; B. T. wuyueen-
Caudal fin with 9+10 principal rays, slightly emargin- sis in situ.
ated, with middle rays longer than two-thirds of longest
rays; both lobes rounded, with upper lobe slightly longer
than lower lobe; procurrent rays slightly extending from Distribution and habitat. Currently only known from
anterior to fin base. Lowest point of caudal peduncle be- the Qiantang-Jiang in Suichang County, Zhejiang Prov-
hind posterior end of anal fin base. ince and Xiuning County, Anhui Province, south China
Colouration. Body yellowish-grey with three ob- (Fig. 7). T. wuyueensis is found in montane streams of
scured and broad vertical brown blotches in smaller in- this river basin (Fig. 8), co-existing with Acrossocheilus
dividuals (the first one below the dorsal fin, the second fasciatus, Pseudogastromyzon fasciatus, Rhinogobius ni-
one closely above the anal fin and the third occupying the ger and Tachysurus albomarginatus.

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592 Jia-Jun, Z. et al.: A new Tachysurus species from the Qiantang-Jiang basin, China

Etymology. The specific epithet is based on the two ri-

val states Wu and Yue which were bordered by the Qian-
tang-Jiang in southeast China more than 2000 years ago.
The onomatopoeic Chinese sound of this species is “Wu
Yue Ni Chang”.

Discussion
Shao and Zhang (2023) erected the Tachysurus prat-
ti group using morphological characters, such as smooth
anterior margin of the pectoral spine, short maxillary-bar-
bel not extending to the base of pectoral-fin, fewer than
20 anal-fins and uniform yellow or brown body in adults.
Under this taxonomic treatment, the T. pratti group is the Figure 9. Relationship between dorsal spine length and HL for
largest group within the genus, with more than twenty spe- the T. pratti-T. truncatus group (including T. wuyueensis, T. prat-
cies. However, the monophyly of the T. pratti group was ti, T. truncatus, T. brachyrhabdion, T. gracilis) and the T. te-
not supported under the molecular scrutiny in this study nuis-T. crassilabris group (including T. tenuis, T. albomargina-
(Fig. 1), indicating these species were wrongly clustered tus, T. analis, T. lani, T. zhangfei, T. ussuriensis, T. crassilabris).
due to retention of conservative morphological features,
common in catfishes (Zhou et al. 2016). Members in the
T. pratti group formed two distinct phylogenetic entities
in our analyses: T. wuyueensis clustering with T. trunca-
tus, T. pratti, T. gracilis and T. brachyrhabdion form a
well-supported monophyletic clade, namely the T. pratti-T.
truncatus group, that is sister to T. nitidus. The remaining
species of the “T. pratti group” form another monophyly
here defined as the T. tenuis-T. crassilabris group, sister to
T. vachelli. In addition, the T. pratti-T. truncatus group plus
T. nitidus appears to be sister to a clade with T. vachelli and
T. tenuis-T. crassilabris group.
Although the morphological similarities between these
two newly-erected species groups are numerous, there are
some distinguishing features that can be used to diagnose
them. The T. pratti-T. truncatus group differs from the
T. tenuis-T. crassilabris group in having a shorter dorsal
spine (length not longer than vs. longer than two-thirds of
HL) (Figs 2, 9, 10). Differences in colour patterns were
also found to be apparent between the two clades. The
body colour of the T. pratti-T. truncatus group is yellow-
ish-brown (Fig. 2), but dark brown in the T. tenuis-T. crass-
ilabris group (Fig. 10). The distinction in colouration of
the abdomen is conspicuous with numerous dark spots in
the T. tenuis-T. crassilabris group, but is absent in the T.
pratti-T. truncatus group (Fig. 11). In addition, the cau-
dal-fin colouration of round-tailed species can be used as
another diagnostic character between these two clades.
Except for T. tenuis, the white or yellowish caudal-fin
margin is broad in the T. tenuis-T. crassilabris group Figure 10. Lateral view of T. albomarginatus (A) for IHB
(Fig. 10), but which is narrow or absent in the T. prat- 202110025885, 133.5 mm SL, Macheng County, Hubei Prov-
ti-T. truncatus group (Fig. 2). The T. pratti-T. truncatus ince; T. analis (B) for IHB201707014311, 124.2 mm SL, Yudu
group can also be distinguished from other species groups County, Jiangxi Province, in Gan-Jiang; T. adiposalis (C) for
of Tachysurus in the external morphology. It differs from FMNH 59079, holotype, 172 mm SL, Tamusui River, Taiwan
the T. aurantiacus group, the T. vachelli group and the Island; (D) Colour in life of adult of T. adiposalis, cited from
T. trilineatus group in having short maxillary barbels not Zhou and Gao (2011).
extending to (vs. extending beyond or reaching) the base
of pectoral fin, from the T. nitidus group, the T. vachelli The topotypes of T. adiposalis were not available in
group and T. virgatus group in having a dorsal fin spine this study, but our photographic examination on the type
shorter (vs. longer) than two-thirds of HL. (FMNH59079) confirmed that T. adiposalis has a long dor-

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Zoosyst. Evol. 100 (2) 2024, 583–595 593

Figure 11. Ventral view of T. zhangfei (A) and T. gracilis (B) that belong to the T. tenuis-T. crassilabris group and the
T. pratti-T. truncatus group, respectively.

sal-fin spine (longer than two-thirds of HL) and 19 anal- 2. Tachysurus albomarginatus: IHB200605264–7,
fin rays (Fig. 10C), which is distinct from T. wuyueensis. 200605269–86, 220605268 72.1–128.5 mm SL,
Based on data from Taiwan, the dark brown body and wide topotypes, China: Anhui Province: Dangtu County:
white margin in the caudal fin are present in a living speci- the lower Yangtze River.
men of T. adiposalis (Fig. 10D) (Zhou and Gao 2011), sug- 3. Tachysurus brachyrhabdion: IHB2017090535–9,
gesting this species may belong to the T. tenuis-T. crass- 107.6–121.8 mm SL, China: Guizhou Province:
ilabris group. Moreover, a phylogenetic analysis, based Songtao County, the Yuan-Jiang of middle Yangtze
on a short segment of mtDNA, showed the specimens of River Basin; IHB2017090522–31, China: Chongq-
T. adiposalis collected from Taiwan are closely related to ing City: Xiushan County, the Yuan-Jiang of the
T. ussuriensis of the T. tenuis-T. crassilabris group. These middle Yangtze River Basin; IHB2016106554,
data confirm our conclusion (Watanabe et al. 2007). 6, 7, China: Hunan Province: Jishou County, the
High levels of inter-specific variations in the shape of Yuan-Jiang of the middle Yangtze River Basin;
the caudal fin occur in the T. pratti-T. truncatus group; for IHB2017090548–58, 588, China: Guizhou Prov-
example, T. pratti has a deeply forked caudal fin, but the re- ince: Tongren City, the Yuan-Jiang of the middle
maining species of the group have round-tailed caudal fins. Yangtze River Basin; IHB 201906012028–31,
A similar scenario also occurs in the T. tenuis-T. crassi- 104.0–127.3 mm SL, China: Hunan Province:
labris group. It suggests the convergent evolution of caudal Xupu County, the Yuan-Jiang of the middle Yang-
fin morphology in Tachysurus. There is a tight link between tze River Basin; IHB2016106295–500, 104.4–
the shape of the caudal fin and habitat preference: round- 124.2 mm SL, China: Hunan Province: Mayang
tailed species present habitat affinities to montane streams, County, the Yuan-Jiang of the middle Yangtze
while fork-tailed species are restricted to rivers’ main River Basin; IHB2017090495–6, 135.3–150.1 mm
stream and lakes. The observation describes the effects of SL, China: Guizhou Province: Jiangkou County,
ecological niches by shaping the caudal fin in Tachysurus. the Yuan-Jiang of the middle Yangtze River Basin;
IHB2017090427–34, 65.4–89.4 mm SL, China:
Guizhou Province: Huangping County, the Yuan-Ji-
Comparative materials examined ang of the middle Yangtze River Basin.
4. Tachysurus crassilabris (Günther 1864): BMNH
1. T. adiposis: FMNH59079, 170 mm SL, China: Ta- 1864.7.9.9 (holotype), 155.2 mm SL, China (photo-
musui River in Taiwan Island. graph and X-ray examined); IHB 201909034566–9,

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594 Jia-Jun, Z. et al.: A new Tachysurus species from the Qiantang-Jiang basin, China

97.1–107.2 mm SL, China: Sichuan Province: Huili IHB201909034750, 57.3 mm SL, Shuifu County,
County: the Jinsha Jiang of the upper Yangtze Riv- the Jinsha-Jiang of the upper Yangtze River Basin;
er; IHB201909017347–61, 69.1–120.5 mm SL, IHB202009016033-6, 48.8–56.0 mm, SL, China:
China: Jiangsu Province: Nanjing City: the lower Yunnan Province: Binchuan County, the Jinsha-Ji-
Yangtze River. ang of upper Yangtze River Basin.
5. T. gracilis: IHB201803023394–403, 93.6–
140.1 mm SL, China: Guangxi Zhuang Autono-
mous Region: Guanyang County: the middle Yang- Conflict of interest
tze River Basin; IHB2017122195, 2017090542–6,
84.1–119.3 mm SL, China: Guangxi Zhuang Au- The authors declare that they have no conflict of interest.
tonomous Region: Yangshuo County: the Pearl
River Basin; IHB201909019374–83, 69.9–112.3
mm SL, China: Hunan Province: Shuangpai Coun- Funding
ty: the Xiang-Jiang of the middle Yangtze River Ba-
sin; IHB201906011312, 175.9 mm SL, China: Hu- This work was partially funded by a grant from the Na-
nan Province: Xinning County: the Zi-shui of the tional Science & Technology Fundamental Resourc-
middle Yangtze River Basin; IHB201906011084, es Investigation Program of Mount Dabie (Grant No.
132.0 mm SL, China: Hunan Province, Dongkou 2019FY101800).
County, the Zi-shui of the middle Yangtze Riv-
er Basin; IHB201906011497–503, 100.8–125.0
mm SL, China: Guangxi Zhuang Autonomous Authors’ contributions
Region: Ziyuan County: the Zi-shui of the mid-
dle Yangtze River Basin; IHB201809019719–22, Jia-Jun Zhou designed the study and revised the manu-
201809019850–1, 48.4–138.8 mm SL, China: Yan- script. Yuan Le-Yang extracted the genomic DNA and per-
ling County, the Xiang-Jiang of the middle Yangtze formed the molecular analysis. Wei-Han Shao examined
River Basin. the specimens and prepared the manuscript. All authors
6. Tachysurus tenuis: IHB201909113617–13629, read and approved the final version of the manuscript.
124.5–159.6 mm SL, China: Zhejiang Province: Ji-
axing City, the Qiantang-Jiang.
7. Tachysurus truncatus: IHB201909038001–10, Acknowledgements
84.9–109.5 mm SL, China: Sichuan Province:
Wanyuan County, the Jialing-Jiang of the upper We are grateful to Huan-Shan Wang (Museum of Aquat-
Yangtze River; IHB2019090112599–613, 76.0– ic Organisms at the Institute of Hydrobiology, IHB) for
111.2 mm SL, China: Sichuan Province: Nanjiang providing comparative specimens; to Zhan-sheng Tang
County, the Jialing-Jiang of the upper Yangtze (Zhejiang Jiulongshan National Natural Reserve) for as-
River; IHB201909037495–506, 20210505666–70, sistance in field surveys.
73.6–123.5 mm SL, China: Hubei Province: Yu-
an’an County, the Qing-Jiang of the middle Yang-
tze River; IHB201904028989-96, 71.5–138.8 mm References
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