Zoosyst. Evol. 100 (2) 2024, 555-563 | DOI 10.3897/zse.100.122472 > PENSUFT. Ate BERLIN A new species of Liobagrus Hilgendorf, 1878 (Teleostei, Siluriformes, Amblycipitidae) from the lower Changjiang River basin in southeast China Zhong-Guang Chen‘, Yan-Shu Guo’, Yu-Ting Dai’, Xiao-Chen Huang!, Jun-Hao Huang’, Jiao Jiang*, Shan Ouyang", An-Xiang Wen°, Xiao-Ping Wu! 1 School of Life Sciences, Nanchang University, Nanchang, Jiangxi 330031, China 2 Ecological Restoration and Conservation on Forest and Wetland Key Laboratory of Sichuan Province, Sichuan Academy of Forestry Sciences, Chengdu, Sichuan 610081, China 3 Museum of Aquatic Organisms, Institute of Hydrobiology, Chinese Academy of Sciences, Wuhan, Hubei 430072, China 4 Zhejiang Museum of Natural History, Hangzhou, Zhejiang 310012, China 5 College of Life Sciences, Sichuan Agricultural University, Yaan, Sichuan 625014, China https://zoobank. org/8SA E11 F0-FC3E-44E2-920B-273CA4370F 0F Corresponding authors: An-Xiang Wen (2960657740@qq.com); Xiao-Ping Wu (xpwu@ncu.edu.cn) Academic editor: Nicolas Hubert # Received 7 March 2024 # Accepted 2 April 2024 Published 16 May 2024 Abstract A new catfish species, Liobagrus chenhaojuni Chen, Guo & Wu, sp. nov., is described from the Tiaoxi River, a tributary of Taihu Lake, located in Zhejiang Province, China. This description is based on morphological characteristics and phylogenetic analysis. This species belongs to a group defined by the presence of a smooth posterior edge of the pectoral-fin spine and can be distinguished from other species in the group by a unique combination of characteristics, including: an upper jaw longer than the lower jaw; maxil- lary barbels reaching the middle of the pectoral fin; irregular blotches present on the lateral body; a rounded caudal-fin with a length ranging from 16.5% to 19.9% of the standard length; 39 to 41 post-Weberian vertebrae; and 15 to 17 anal-fin rays. The validity of this new species is further supported by the molecular phylogenetic analysis based on Cytb sequences. Key Words catfish, phylogeny, taxonomy, Zhejiang Province Introduction The genus Liobagrus comprises a group of small fresh- water catfish endemic to East Asia. To date, 20 species have been described, with 12 of them found in mainland China: Liobagrus marginatus (Gunther, 1892), Liobagrus nigricauda Regan, 1904, Liobagrus stvani Regan, 1908, Liobagrus anguillicauda Nichols, 1926, Liobagrus mar- ginatoides (Wu, 1930), Liobagrus kingi Tchang, 1935, Liobagrus aequilabris Wright & Ng, 2008, Liobagrus chenghaiensis Sun, Ren & Zhang, 2013; Liobagrus huai- heensis Chen, Wu & Wen, 2021; Liobagrus pseudostyani Chen & Guo, 2021; Liobagrus brevispina Xie, Cao & Zhang, 2022; and Liobagrus chengduensis Chen, Guo, Wu & Wen, 2022 (He 1999; Wright and Ng 2008; Sun et al. 2013; Chen and Guo 2021; Chen et al. 2021; Chen et al. 2022; Xie et al. 2022). Research on the taxonomy of Chinese Liobagrus has mainly focused on the upper and middle reaches of the Changjiang River (Yangtze) and the Huaihe River basin. However, the Liobagrus species in the lower Changjiang River have not been thorough- ly surveyed or studied, potentially leaving other unde- scribed species undiscovered. The Tiaoxi River is a small river situated in the western part of the Hangjiahu Plain in Zhejiang Province and is one of the main tributaries of Taihu Lake. Despite its small Copyright Chen, Z.-G. 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. 556 watershed area, the Tiaoxi River boasts a diverse fish pop- ulation, with a total of 84 recorded fish species (Li and Shimatani 2016; Zhang et al. 2022). Among these species, the presence of L. styani is highly questionable, as it is believed to be narrowly distributed in the Juanshui River basin in the middle Changjiang River (Wu et al. 2013). Moreover, the L. styani recorded in the basin is not only far from its type locality but also from the distribution regions of all other Liobagrus species in China. Based on collec- tions made between 2022 and 2024, it has been discovered that Liobagrus specimens with a smooth posterior edge of the pectoral-fin spine, distributed in the Tiaoxi River, represent an undescribed species. Here, we introduce this Species as new to science. The discovery of this new taxon contributes to our understanding of the high level of fish endemism in the lower Changjiang River basin. Materials and methods Specimens were manually collected from the Tiaox1 Riv- er between 2022 and 2024. Thirty-six type specimens were initially preserved in 10% formalin and subsequent- ly transferred to 70% ethanol for long-term storage. Ad- ditionally, seven type specimens were preserved in 99% ethanol for molecular phylogenetic analyses. Vertebrae and fin rays were detected and counted using X-ray imag- ing. Measurements were obtained using digital calipers, with values recorded to the nearest 0.1mm. Genomic DNA was extracted from the ventral fin of specimens preserved in 99% ethanol using the Baypure Magnetic Bead Method Animal Genomic DNA Ex- traction Kit (BayBio, Guangzhou, China). The quality and concentration of the DNA were checked using 1% agarose gel electrophoresis and NanoDrop 2000 (Ther- mo Scientific, USA). Cytb sequences were amplified us- ing primers L14724 (GACTTGAAAAACCACCGTTG) and H15915 (CTCCGATCTCCGGATTACAAGAC). Polymerase chain reaction (PCR) amplifications of Cytb were performed in a final 25-yL volume mixture contain- ing 1 uL of template DNA, | uL of each pair of primers, 12.5 uwL of Green Taq Mix (Vazyme, China), and 9.5 wL ddH20. Thermal cycling began with one cycle at 95 °C for 10 s, followed by 35 cycles of denaturation at 94 °C for 1 min, 55 °C for 1 min, and 72 °C for 1 min, witha final extension step at 72 °C for 10 min. PCR products were purified and sequenced using an ABI 3730XL an- alyzer by Sangon Biotech (China). Accession numbers of all newly obtained sequences are provided in Table 1. Sequences were aligned using MEGA v. 6.0 (Tamura et al. 2013) and manually checked. Genetic distance, based on the uncorrected p-distance model, was calculated us- ing MEGA vy. 6.0. Phylogenetic relationships were re- constructed using Bayesian inference (BI) and maximum likelihood (ML). Xiurenbagrus xiurenensis (Yue, 1981), X. gigas Zhao, Lan & Zhang, 2004, Akysis brachybar- batus Chen, 1981, Jctalurus furcatus (Valenciennes, zse.pensoft.net Chen, Z.-G. et al.: A new species of Liobagrus 1840), and Noturus taylori Douglas, 1972, were used as the outgroup for rooting the tree. ML analyses were per- formed in IQ-TREE v. 1.6.12 (Minh et al. 2013) using the Ultrafast Fast Bootstrap approach (Minh et al. 2013) with 10,000 reiterations. The most appropriate model of sequence evolution (GTR+I+G) was selected using Parti- tonFinder2 v. 1.1 (Robert et al. 2017). Bayesian inference (BI) was conducted in MrBayes v. 3.2.6 (Ronquist et al. 2012). The most appropriate model of sequence evolution (GTR+I+G) was selected under ModelFinder (Subha et al. 2017). Four simultaneous runs with four independent Markov Chain Monte Carlo (MCMC) algorithms were executed for 10 million generations, and trees were sam- pled every 1000 generations with a burn-in of 25%. The convergence was verified with the average standard devi- ation of split frequencies of <0.01 and the potential scale reduction factor (PSRF) of ~1. Trees were visualized using FigTree v.1.4.3 (http://tree.bio.ed.ac.uk/software/ figtree/). Institutional abbreviations used: NCU_XPWU Laboratory of Xiao-Ping Wu, Nanchang University (Nanchang, Jiangxi, China); [HB Museum of Aquatic Or- ganisms, Institute of Hydrobiology, Chinese Academy of Sciences (Wuhan, Hubei, China). Results Liobagrus chenhaojuni Chen, Guo & Wu, sp. nov. https://zoobank.org/9966A 844-20BC-4DCD-9E09-A 5FFA9F 14659 Figs 1, 2A—C, Table 2 Liobagrus styani Li & Shimatani, 2016: 165-167 (Tiaoxi River, Zhe- jiang, China). Type material. Holotype. 24 NCU _XPWU_Y01, Siling Reservoir [VUlS7k ], Tiaoxi River [2/34], Yuhang dis- trict [At], Hangzhou City [HiT], Zhejiang Prov- ince [WryL#@4], China, 30°25'42"N, 119°45'18"E, leg. Hao-Jun Chen, February 2024. September 2022. Paratypes. 24 NCU_XPWU_Y02-— 16, n=15, other information same as holotype; 22_ NCU_XPWU_ YOI-17, n=17, leg. Zhong-Guang Chen & Hao-Jun Chen, September 2022, other information same as holotype; 22 NCU XPWU_Y18-—25, n=8, IHB- T-A0000007-8, n=2, Tiaoxi River [73%], Deqing Coun- ty (78 2], Huzhou City [WIN TH], Zhejiang Province [ Wty |, China, leg. local people, September 2022. Diagnosis. Liobagrus chenhaojuni sp. nov. is a mem- ber of the group defined by the presence of a smooth posterior edge of the pectoral-fin spine (1.e., L. reinii, L. formosanus, L. styani, L. nantoensis, L. anguillicauda, L. marginatoides, and L. aequilabris). It can be distin- guished from all other species in this group by the fol- lowing characteristics: the upper jaw 1s longer than the lower jaw (vs. equal in L. aequilabris and L. formosanus, shorter in L. marginatoides), the maxillary barbels reach the middle of the pectoral fin (vs. reach the pectoral-fin Zoosyst. Evol. 100 (2) 2024, 555-563 Table 1. GenBank accession numbers of the sequences for this study. 557 Species Access number Liobagrus chenhaojuni sp. nov. 1 PP446311 Liobagrus chenhaojuni sp. nov. 2 PP446312 Liobagrus chenhaojuni sp. nov. 3 PP446313 Liobagrus chenhaojuni sp. nov. 4 PP446314 Liobagrus chenhaojuni sp. nov. 5 PP446315 Liobagrus chenhaojuni sp. nov. 6 PP446316 L. styani 1 KY653576 L. styani 2 KY653577 L. aequilabris 1 KY653673 L. aequilabris 2 KY653674 L. anguillicauda 1 KY653651 L. anguillicauda 2 KY653652 L. marginatus 1 KY653578 L. marginatus 2 KY653579 L. kingi KC193779 L. huaiheensis 1 ON638213 L. huaiheensis 2 ON638214 L. obesus DQ321752 L. andersoni DQ321753 L. pseudostyani 1 ON638209 L. pseudostyani 2 ON638210 L. brevispina 1 ON638211 L. brevispina 2 ON638212 L. chengduensis 1 ON638203 L. chengduensis 2 ON638204 L. mediadiposalis 1 KX265422 L. mediadiposalis 2 KX265423 L. hyeongsanensis MZ066608 L. geumgangensis 1 KX265431 L. geumgangensis 2 KX265433 L. somjinensis MN756661 L. reinii 1 [6333217 L. reinii 2 LC333224 Xiurenbagrus xiurenensis DQ192464 Xiurenbagrus gigas EU490936 Akysis brachybarbatus AF499603 Ictalurus furcatus KM576102 Noturus taylori KP013089 insertion in L. styani, L. reinii, and L. nantoensis); pres- ence of irregular blotches on the lateral body (vs. absence in L. formosanus, L. nantoensis, L. anguillicauda, L. mar- ginatoides, and L. aequilabris), the caudal fin is round- ed (vs. sub-truncate in L. marginatoides), the caudal fin length ranges from 16.5% to 19.9% standard length (vs. 13.1-16.2 in L. styani, 20.3—27.0 in L. anguillicauda and 20.1—26.9 in L. aequilabris); it possesses 39-41 post-We- berian vertebrae (vs. 35-37 in L. aequilabris), the anal-fin rays range from 15 to 17 (vs. 12 in L. nantoensis) (Table 3). Description. Morphometric data for type specimens are shown in Table 2. Body elongated, anteriorly de- pressed (wider than deep), and posteriorly evenly com- pressed to the tail. Lateral line short, with 6-9 pores. Head depressed and broad when viewed dorsally, with a broad- ly rounded snout in dorsal view. Anterior nostril tubular, with a rim bearing a fleshy flap forming a short tube; Locality Reference Huzhou, Zhejiang, China This study Huzhou, Zhejiang, China This study Huzhou, Zhejiang, China This study Hangzhou, Zhejiang, China This study Hangzhou, Zhejiang, China This study Hangzhou, Zhejiang, China This study Xianning, Hubei, China NCBI Xianning, Hubei, China NCBI Unknown NCBI Unknown NCBI Wuyishan, Fujian, China NCBI Wuyishan, Fujian, China NCBI Chongaing, China NCBI Chongaing, China NCBI Unknown NCBI Xinyang, Henan, China NCBI Xinyang, Henan, China NCBI Korea NCBI Korea NCBI Chengdu, Sichuan, China NCBI Chengdu, Sichuan, China NCBI Chengdu, Sichuan, China NCBI Chengdu, Sichuan, China NCBI Chengdu, Sichuan, China NCBI Chengdu, Sichuan, China NCBI Korea NCBI Korea NCBI Korea NCBI Korea NCBI Korea NCBI Korea NCBI Japan NCBI Japan NCBI Guangxi, China NCBI Guangxi, China NCBI Yunnan, China NCBI Unknown NCBI Unknown NCBI posterior nostril pore-like, with the rim posteriorly con- fluent with the base of the nasal barbel. Eyes small, dorso- lateral, and subcutaneous. Mouth terminal, with the upper jaw noticeably longer than the lower jaw, lips thickened. Premaxillary and mandibular tooth pads curved, bearing small and setiform teeth; palatine teeth absent. Four pairs of barbels: the maxillary barbel long, extending to the pectoral-fin insertion; nasal barbel short, not reaching the gill-membrane margin; inner mandibular barbel approxi- mately half the length of the outer mandibular barbel and does not extend to the pectoral-fin insertion; outer mental barbel longest, reaching the middle of the pectoral fin. Dorsal fin II, 5—6 rays, with a convex distal margin; tip of adpressed fins does not reach the pelvic-fin in- sertion. Dorsal-fin spine covered by thick, straight skin with smooth anterior and posterior margins, slightly shorter than the pectoral-fin spine. Adipose fin high, zse.pensoft.net 558 Chen, Z.-G. et al.: A new species of Liobagrus Figure 1. Liobagrus chenhaojuni sp. nov. A—C. Dorsal, lateral, and ventral view of holotype (24 NCU _XPWU_Y01):; D. Dorsal view of pectoral-fin spine of paratype (22 NCU _XPWU_Y31). Arrows show the anus. with its base longer than the anal-fin base, confluent with the caudal fin without a marked incision at the confluence. Pectoral fin I, 7-8 rays, with its origin at the vertical through the edge of the operculum, par- tially covered by the opercular membrane. Pectoral-fin spine long and sharp, with smooth anterior and posteri- or margins (Fig. 1D), reaching the dorsal-fin insertion. Pelvic fin 1, 5—6 rays, short, with the adpressed tip not reaching the anal-fin origin. Anal fin 15-17 rays with a rounded distal margin, and its tip approaches the or- igin of the ventral procurrent caudal-fin rays, longer zse.pensoft.net than the dorsal-fin base but shorter than the adipose-fin base, with a convex distal edge. Anus closer to the pel- vic-fin insertion than to the anal-fin origin. Caudal fin rounded, with 43—50 rays. Vertebral column consists of 39-41 post-Weberian elements. Body generally dark brown to brownish red, adorned with irregular yellowish blotches that fade to light yellow ventrally. All barbels grayish white to light yellow, while dorsal fins dark brown, and adipose and caudal fins gray- ish white to light brown. All fins exhibit narrow, gray- ish white to light yellowish distal margins (Fig. 2A, B). Zoosyst. Evol. 100 (2) 2024, 555-563 Figure 2. Living specimens of Liobagrus chenhaojuni sp. nov. and its similar congeneric species. A, B. Liobagrus chenhaojuni sp. nov.; C. Liobagrus chenhaojuni sp. nov. albino individual; D. L. styani; E. L. anguillicauda, F. L. brevispina, G. dorsal view of pectoral-fin spine of L. brevispina. An albino individual was found, exhibiting a generally pink body without irregular yellowish blotches (Fig. 2C). Etymology. This species is named after Mr. Hao-Jun Chen, who assisted in the field survey. Vernacular name. {711k (Pinyin: zhe jiang yang). Distribution and ecology. Liobagrus chenhaojuni sp. nov. is exclusively found within the Tiaoxi River basin (Fig. 3). Within this habitat, it typically resides at the bottom of the stream with medium pebbly substrates, together with Rhinogobius leavelli (Herre, 1935), Micro- physogobio bicolor (Nichols, 1930), Vanmanenia steno- soma (Boulenger, 1901), Acrossocheilus fasciatus (Stein- dachner, 1892), Opsariichthys bidens Ginther, 1873, and Zacco tiaoxiensis Zhang, Zhou & Yang, 2022 (Fig. 4). Molecular analyses. A dataset consisting of 33 Cytb sequences and five outgroup taxa was employed for phylogenetic analyses (Table 1). The alignment of Cytb exhibited a length of 1116 characters, with 343 variable sites and 313 sufficiently informative sites. Phylogenetic analyses generated ML and BI trees with largely congru- ent topologies (Fig. 5). Notably, species from China with a smooth posterior edge of the pectoral-fin spine formed a monophyletic group, while those with a serrated posterior edge of the pectoral-fin spine constituted a paraphyletic group. The validity of Liobagrus chenhaojuni sp. nov. is supported by the molecular-phylogenetic result. It be- longs to the group with a smooth posterior edge of the pectoral-fin spine, and the phylogenetic relationship with- zse.pensoft.net 560 Table 2. Morphometric data for type specimens of Liobagrus chenhaojuni sp. nov. Morphometrics Holotype Paratypes Standard length (mm) 70.2 38.8-79.3 % of standard length Head length 22.4 20.6-23.0 Body depth 18.8 13.6-18.4 Dorsa-fin base length 10.0 9.1-10.0 Anal-fin base length 16.5 12.3-19.7 Adipose-fin base length 28.8 27.7-37.2 Caudal peduncle length 20.4 19.3-23.1 Caudal peduncle depth Le? 14.0-17.7 Dorsal-fin spine length iS) 6.3-7.4 Pectoral-fin spine length 9.3 7.2-9.6 Caudal-fin length 18.4 16.5-19.9 Anus to pelvic-fin insertion 3.6 3.4-6.4 Anus to anal-fin origin 4.3 4.9-7.5 Predorsal length 28.8 27.0-30.5 Prepectoral length 18.4 17.7-21.4 Prepelvic length 46.0 42.9-45.5 Preanal length 58.4 57.3-60.1 Dorsal to adipose-fin origin 298 25.3-31.8 % of head length Head width 89.2 83.0-93.3 Head depth 61.8 50.0-59.4 Snout length 31.8 26.3-32.0 Mouth width 69.4 68.0-76.1 Interorbital width B57 33.1-40.0 Nasal barbel length ESi2 67.6-73.9 Maxillary barbel length 94.9 81.4-95.4 Inner mandibular barbel length 61.8 50.0-62.1 Outer mandibular barbel length 99.4 93.0-98.7 Width between anterior nares 12.4 12.5-16.1 Width between postoral nares Dour 28.0-36.6 102°E 105°E Chen, Z.-G. et al.: A new species of Liobagrus in the group is represented as Liobagrus chenhaojuni sp. nov. + (L. anguillicauda + (L. aequilabris + L. styani)). The genetic distances between Liobagrus chenhaojuni sp. nov. and other congeneric species ranged from 5.8% to 14.2% (Suppl. material 1). Discussion The placement of the new species within Liobagrus is supported by both its morphology and phylogeny. Species of Liobagrus can be divided into two groups based on the possession of a smooth or serrated posterior edge of the pectoral-fin spine. Xie et al. (2022) described L. brevispi- na as having a smooth posterior edge of the pectoral-fin spine. However, based on the examination of specimens, L. brevispina was found to have a serrated posterior edge of the pectoral-fin spine (Fig. 2G). The serrations on its pectoral-fin spine weaken but do not disappear as it grows. Currently, the group defined by the presence of a smooth posterior edge of the pectoral-fin spine comprises only seven species: L. reinii, L. formosanus, L. styani, L. nan- toensis, L. anguillicauda, L. marginatoides, and L. aequi- labris. In this group, Liobagrus chenhaojuni sp. nov. can be easily distinguished from L. aequilabris, L. formosa- nus, and L. marginatoides by having the upper jaw longer than the lower (vs. shorter or equal). Liobagrus chenhao- juni sp. nov. is similar to L. reinii, L. styani, L. nantoen- sis, and L. anguillicauda by the similar upper and lower jaw positions, but differs based on the maxillary barbels reaching the middle of the pectoral fin (vs. reaching the 123°E 30°N 27°N 24°N 117°E 120°E 123°E Figure 3. Distribution of Liobagrus with a smooth posterior edge of the pectoral-fin spine in Mainland China. Dot. Liobagrus chenhaojuni sp. nov.; square. L. anguillicauda;, rhombus. L. styani; star. L. aequilabris, triangle. L. marginatoides. Solid show the type localities. zse.pensoft.net Zoosyst. Evol. 100 (2) 2024, 555-563 561 Wi = i la eS Figure 4. Sampling locality of Liobagrus chenhaojuni sp. nov. Siling Reservoir, Tiaoxi River, Yuhang district, Hangzhou City, Zhejiang Province, China. BI smooth pectoral-fin spine ; 0.99 serrated pectoral-fin spine serrated pectoral-fin spine . Liobagrus geumgangensis 2 0.05 ~ Liobagrus anguillicauda 1 ~~ ~~~ Liobagrus anguillicauda 2 Liobagrus chenhaojuni sp. Liobagrus chenhaojuni sp. -- Liobagrus chenhaojuni sp. ~~ Liobagrus chenhaojuni sp. Liobagrus chenhaojuni sp. ~ Liobagrus chenhaojuni sp. ooo Liobagrus kingi Liobagrus huaiheensis 1 . Liobagrus andersoni -- Liobagrus pseudostyani 1 - Liobagrus pseudostyani 2 Liobagrus brevispina 1 Liobagrus brevispina 2 . Liobagrus chengduensis 1 Liobagrus chengduensis 2 .. Liobagrus mediadiposalis 1 .. Liobagrus mediadiposalis 2. -- . Liobagrus hyeongsanensis Liobagrus reinii 2 .... Xiurenbagrus xiurenensis ..... ; Xiurenbagrus gigas —...... cee Akysis brachybarbatus AG Cal GEIS TEC AES - teed ssc taeratntrvatrte dS exhbesselige: a nares abel eich cece codehish wu... Noturus taylori sec CRIS ee Sid aelaalesh Moai = Liobagrus styani 2 Liobagrus aequilabris 1 ~~ Liobagrus aequilabris 2 = MOVs at ae nov. 3 nov. “| 9 ochaipiabbls iat eecoea teers tn . nov. 4 China nov. 5 nov. 6 heen itestangehaghetsanand Liobagrus marginatus 10 - - Liobagrus marginatus 20 Liobagrus huaiheensis 2 ~~~ Korea Jeeps ee, re meme nero eth OY China 85 . Liobagrus geumgangensis 1. a4 Korea Liobagrus SOMPINENSIS . Liobagrus reinii1 ——— ~~~ 100 Japan ——————E— 100 0.05 Figure 5. Bayesian inference tree and maximum likelihood tree inferred from Cytb gene sequences. Posterior probabilities/boot- strap supports are shown on the left/right of nodes. pectoral-fin insertion in L. reinii, L. styani, and L. nan- toensis), presence of irregular blotches on the body lat- eral (vs. absence in L. nantoensis and L. anguillicauda), caudal-fin length is 16.5—19.9% of standard length (vs. 13.1-16.2% in L. styani and 20.3—27.0% in L. anguil- licauda), 15-17 anal-fin rays (vs. 12 in L. nantoensis). Furthermore, Liobagrus chenhaojuni sp. nov. has a dis- tinctive distribution, being exclusively found in the Tia- oxi River within the lower Changjiang River basin, far from any other congeners. The discovery of Liobagrus chenhaojuni sp. nov., as well as Zacco tiaoxiensis in the Tiaoxi River, shows zse.pensoft.net 562 Chen, Z.-G. et al.: A new species of Liobagrus Table 3. Comparisons of major diagnostic characters of Liobagrus with a smooth posterior edge of the pectoral-fin spine. Characters Liobagrus chenhaojuni L. nantoensis? L. reinii? L. styan® sp. nov. Upper/lower jaw in length Esl alk > >t post-Weberian vertebrae 39-41 Unknown Unknown 39-41 Anal-fin rays 15-17 12 17 17-19 Caudal-fin length as % of standard 16.5-19.9 Unknown Unknown 13.11-16.2 length Caudal-fin shape Rounded Rounded Rounded Rounded Maxillary barbels Reaching middle of Reaching pectoral-fin Reaching pectoral-fin Reaching pectoral-fin pectoral fin insertion insertion insertion Irregular blotches on body lateral Present Absent Present Present Characters L. anguillicauda L. aequilabris? L. formosanus® L. marginatoides¢ Upper/lower jaw in length >1 =] =i