Biodiversity Data Journal :
Taxonomic Paper
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Corresponding author: Chuanjiang Zhou (chuanjiang88@163.com), Guoxing Nie (niegx@htu.cn)
Academic editor: Yahui Zhao
Received: 26 Feb 2021 | Accepted: 01 Jun 2021 | Published: 16 Jun 2021
© 2021 Chuanjiang Zhou, Wenwen Ma, Xi Wang, Yongtao Tang, Xiaoling Meng, Guoxing Nie
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:
Zhou C, Ma W, Wang X, Tang Y, Meng X, Nie G (2021) Homatula guanheensis sp. nov. (Teleostei: Nemacheilidae), a new species of loach from Henan Province, China. Biodiversity Data Journal 9: e65130. https://doi.org/10.3897/BDJ.9.e65130
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The genus Homatula belongs to the order Cypriniformes and family Nemacheilidae. Nichols (1925) set up the genus as a subgenus of Barbatula by the type species of Nemacheilus potanini. Currently, it is recognised as a valid genus. Nineteen valid species have been already reported in the drainage of the Yellow, Yangtze, Pearl, Lancang, Red and Nujiang Rivers. H. variegata, H. longidorsalis, H. berezowskii and H. potanini are distributed in the Yangtze River drainage in China. H. laxiclathra is mainly distributed in the Weihe River, a tributary of the Yellow River. The remaining species are mainly distributed in the rivers of Yunnan Province.
Homatula guanheensis sp. nov., a new species, is described from the Guanhe River of the HanJiang River drainage (a tributary of the Yangtze River), Xixia County, Henan Province, China. It can be distinguished from its congeners by a combination of the following characters: the vertical brown bars on the body are wider than their interspaces, numbering 19–22; predorsal body partially scaled; the lateral line complete; adipose crest on caudal peduncle not reaching forward; the position of the anal-fin origin and the intestinal form. The new species displays distinct molecular divergence in the Cytochrome oxidase I (COI) and Cytochrome b (Cyt b) genes.
Homatula, morphology, Hanjiang River drainage, taxonomy
The Nemacheilid genus Homatula was established by Nichols in 1925 as a subgenus of Barbatula, based on the type species Nemacheilus potanini. Species from Homatula are small-size benthic fishes that are distributed in the drainage of the Yellow, Yangtze, Pearl, Lancang, Nujiang and Red Rivers. Some researchers have treated Homatula as a synonym of Paracobitis according to its adipose-crest (
The morphological variation of Homatula in the upper Yellow River drainage, the upper-middle reaches of Yangtze River drainage and the upper Pearl River drainage was investigated by Zeng et al. in 2012. However, the taxonomic status of the specimens from the Hanjiang River of the middle Yangtze River drainage were undetermined (
All specimens were examined and stored in the collection of the College of Fisheries, Henan Normal University, Xinxiang, Henan Province, China. For morphological analysis, 10 specimens were fixed in 10% formalin. All measurements and counts were made following
Species |
COI Gene Accession Numbers |
Cyt b Gene Accession Numbers |
H. longidorsalis |
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H. anguillioides |
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H. acuticephala |
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H. berezowskii |
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H. coccinocola |
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H. disparizona |
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H. potanini |
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H. pycnolepis |
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H. variegata |
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H. wuliangensis |
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H. guanheensis sp. nov |
MT771706-MT1722 |
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Schistura latifasciata |
* Sequences were retrieved from GenBank.
All the sequences in this study were retrieved from GenBank and the accession numbers of the newly determined sequences in this study are MT771689-MT771705 (COI) and MT771706-MT1722 (Cyt b).
Body elongate, anterior portion nearly cylindrical and posterior portion compressed; body depth 11.63% (10.41–13.47%) in SL (Table
Holotype |
Paratypes (n = 9) |
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range |
mean |
SD |
||||
Standard length (mm) |
99.6 |
76.9-109.26 |
91.8 |
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As%SL |
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Body depth |
11.6 |
10.4-13.4 |
11.9 |
0.8 |
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Head length |
18.5 |
17.3-20.3 |
18.7 |
0.9 |
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Dorsal-fin length |
18.5 |
17.3-21.3 |
19.3 |
1.3 |
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Pectoral-fin length |
12.9 |
11.4-14.2 |
12.7 |
0.9 |
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Pelvic-fin length |
10.5 |
10.3-12.7 |
11.4 |
0.8 |
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Anal-fin length |
13.4 |
11.6-14.1 |
13 |
0.8 |
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Predorsal length |
46.5 |
42.3-49.1 |
45.7 |
2.3 |
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Pre-anus length |
18.1 |
16.4-20.2 |
18.4 |
1.3 |
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Prepelvic length |
47.2 |
43.9-49.3 |
46.7 |
1.2 |
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Pre-anal length |
72.7 |
68.8-73.4 |
71.1 |
2 |
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Caudal peduncle depth |
13.6 |
10.1-13.6 |
11.3 |
1.2 |
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Caudal peduncle length |
20.9 |
16.4-23.6 |
20.2 |
1.8 |
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As%Caudal peduncle length |
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Caudal peduncle depth |
65.1 |
45.7-76.8 |
56.6 |
8.9 |
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As%HL |
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Head depth |
43.4 |
42.4-49.9 |
45.8 |
2.6 |
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Head width |
53.7 |
49.3-61.1 |
56.1 |
3.7 |
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Eye diameter |
14.6 |
12.2-16.9 |
14.6 |
1.2 |
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Snout length |
40.1 |
34.4-44.3 |
40.7 |
3.5 |
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Interorbital width |
29.9 |
23.8-31.1 |
28.5 |
2.2 |
Component matrix of the Principal Component Analysis from morphometric data of H. guanheensis and H. laxiclathra.
PC1 | PC2 | PC3 | |
Body depth | 0.156 | 0.126 | 0.889 |
Head length | 0.851 | 0.264 | 0.229 |
Head depth | 0.515 | -0.088 | 0.46 |
Head width | 0.136 | -0.074 | 0.099 |
Eye diameter | 0.807 | -0.041 | 0.384 |
Interorbital width | 0.137 | 0.91 | 0.045 |
Snout length | 0.701 | 0.148 | 0.036 |
Dorsal-fin length | 0.015 | 0.976 | -0.172 |
Pectoral-fin length | 0.78 | 0.094 | 0.31 |
Pelvic-fin length | 0.683 | 0.527 | 0.196 |
Anal-fin length | 0.182 | 0.961 | -0.078 |
Predorsal length | 0.346 | 0.054 | 0.724 |
Prepelvic length | 0.292 | -0.09 | 0.753 |
Prepectoral length | 0.481 | 0.095 | 0.606 |
Pre-anal length | 0.621 | -0.161 | 0.333 |
Caudal peduncle length | -0.235 | 0.817 | 0.229 |
Caudal peduncle depth | -0.681 | 0.111 | -0.447 |
Cumulative variance (%) | 59.35 | 79.65 | 85.10 |
Head and body brown; a series of 19–22 body bars, each bar at least twice as wide as the interspace. Abdomen yellowish. Dorsal fin with two dark brown marks, one at the base, the other at postmedian of the fin; posterior border of dorsal fin white. Pectoral fins with dark brown spots. Pelvic and anal fins white, dark at the base. Adipose keels white with dark brown spots. Caudal fin dark grey; brown vertical bars on caudal fin base (Fig.
Homatula guanheensis is different from its congeners in the following characters: partly scaled (vs. in the latter, scales are totally absent or only a few scales on the caudal peduncle in H. nanpanjiangensis, H. oligolepis, H. disparizona and H. wenshanensis vs. scales all over the body, except for the head in H. acuticephala, H. anguillioides, H. pycnolepis, H. wuliangensis, H. change and H. coccinocola); complete lateral line (vs. incomplete lateral line in H. potanini and H. wujiangensis); the vertical brown bars on the body are wider than their interspaces, numbering 19–22 (vs. equal to interspace or slightly wider than its interspace in H. variegate, H. berezowskii and H. longidorsalis); caudal fin micro-rounded (vs. truncated or oblique in H. variegata and H. berezowskii); adipose crest on caudal peduncle not reaching forward of the position of the anal-fin origin (vs. beyond in H. variegate vs. identical in H. longidorsalis); predorsal body partially scaled (vs. absent scales in H. berezowskii and H. longidorsalis); anterior nostril forming a valve (vs. forming a spool in H. longidorsalis). The vertical brown bars on the body of the new species are similar to H. laxiclathra. The new species can be further distinguished from H. laxiclathra in that the intestine forms a bend, not reaching the posterior surface of U-shaped stomach (vs. a loop anteriorly reaching the posterior surface of the U-shaped stomach); anterior with only a few and scattered scales (vs. scaleless).
The specific epithet Guanheensis is derived from Guanhe River (鹳河 in Chinese, type locality) with the Latin suffix "-ensis".
Keys to species of the genus Homatula in China |
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1 | Lateral line incomplete | 2 |
– | Lateral line complete | 3 |
2 | Scales present; covering whole body except head and abdomen; dorsal crest high and long, from dorsal-fin base to caudal-fin base (Jinshajiang River, Sichuan Province) | H. potanini (Günther, 1896) |
– | Body scaleless or with rudimentary scales; dorsal crest high and short, upper crest not reaching the posterior point of anal-fin base (Wujiang River, Sichuan Province) | H. wujiangensis (Ding & Deng, 1990) |
3 | Anterior ending of adipose crest does not reach posterior end of anal-fin base | 4 |
– | Anterior ending of adipose crest reach posterior end of anal-fin base | 7 |
4 | The length of dorsal-fin base longer than the longest branched dorsal-fin ray (Nujiang River, Yunnan Province) | H. cryptoclathrata (Li et al., 2019) |
– | The length of dorsal-fin base shorter than the longest branched dorsal-fin ray | 5 |
5 | Having clear pattern of marks on the flank | 6 |
– | No marks on the flank (Nujiang River, Yunnan Province) | H. nigra (Li et al., 2019) |
6 | 27-34 marks on the flank; dorsal and pelvic fins closer to snout (Nujiang River, Yunnan Province) | H. anteridorsalis (Li et al., 2019) |
– | 20-22 marks on the flank; dorsal and pelvic fins located intermediate of body (Mengyejiang River, Yunnan Province) | H. change (Endruweit, 2015) |
7 | Scales are totally absent or only a few scales on the caudal peduncle | 8 |
– | Scales clearly present, covering posterior of body at least | 11 |
8 | No median notch on lower jaw | 9 |
– | Median notch on lower jaw | 10 |
9 | The caudal fin emarginate; caudal peduncle narrow and long; vertebrae 4+39~40 (Panlong River, Yunnan Province) | H. disparizona (Min et al., 2013) |
– | The caudal fin furcate; caudal peduncle wider and shorter; vertebrae 4+47~48 (the Red River, Yunnan Province) | H. wenshanensis (Yang et al., 2017) |
10 | Body with regular vertical bars and bars in front of dorsal fin conspicuously thinner than those behind; no vermiform markings on parietal area or obscure; tip of pelvic fin closing or reaching anus (Nanpanjiang River, Yunnan Province) | H. nanpanjiangensis (Min et al., 2010) |
– | Body and head with vermiform markings; dorsal fin and pectoral fin covered by small spots on both sides; tip of pelvic fin quite far away from anus (Yangzonghai River, Yunnan Province) | H. oligolepis (Cao & Zhu, 1989) |
11 | Scales covering whole body, except head | 12 |
– | Posterior part of the body covered by scales; anterior part scaleless or with only a few and scattered scales | 16 |
12 | Having pelvic axillary lobe | 13 |
– | Absent pelvic axillary lobeAbsent pelvic axillary lobe | 15 |
13 | A pair of free protrusions present in pelvic fins (Yangbijiang River, Yunnan Province) | H. pycnolepis (Hu & Zhang, 2010) |
– | Absent free protrusions in pelvic fins | 14 |
14 | Caudal fin slightly emarginated to nearly truncate; having notch on the lower jaw; 16-19 brown bars on a beige background, bars are somewhat straight and never vertically split, notch on the lower jaw (Tengtiaojiang River, Yunnan Province) | H. coccinocola (Min et al., 2018) |
– | Caudal fin rounded; lacking notch on the lower jaw; 22-26 brown bars on body (Lancang River, Yunnan Province) | H. wuliangensis (Min et al., 2012) |
15 | Caudal fin truncated; body depth extremely decreased posterior of dorsal-fin base; head sharp (Haixihai Lake, Yunnan Province) | H. acuticephala (Zhou & He, 1993) |
– | Caudal fin oblique; body depth quite uniform from head to tail; head blunt (Mekong River, Yunnan Province) | H. anguillioides (Zhu & Wang, 1985) |
16 | Anterior nostrils pierced in front side of a tube; dorsal fin is far from the snout (Nanpangjiang River, Yunnan Province) | H. longidorsalis (Yang et al., 1989) |
– | Anterior nostrils pierced in front side of a flap, dorsal fin closer to the snout | 17 |
17 | Vertical brown bars narrower or slightly wider than their interspaces | 18 |
– | Vertical brown bars distinctly wider than interspaces | 19 |
18 | Caudal fin oblique; intestine forming a single loop; adipose crest of the caudal peduncle anteriorly extending through the anal-fin origin (Jinshajiang River, Weihe River) | H. variegata (Sanvage & Dabry, 1874) |
– | Caudal fin truncate; intestine forming a zigzag loop; adipose crest of the caudal peduncle anteriorly not extending through the anal-fin origin (Jialingjiang River, Gansu Province) | H. berezowskii (Günther, 1896) |
19 | Caudal fin oblique; intestine with a loop anteriorly reaching the posterior surface of the U-shaped stomach; anterior scaleless (Weihe River, Shaanxi Province) | H. laxiclathra (Gu & Zhang, 2012) |
– | Caudal fin micro-rounded; intestine forming a bend; anterior with only a few and scattered scales (Guan River, Henan Province) | H. guanheensis sp. nov. |
Genetic distance and phylogentic trees
Based on the COI gene, sequences of 11 studied species were used to construct a BI tree. The new species belonged to a different clade with strong high PP values (100%, Fig.
Pairwise comparisons of genetic distance of nine Homatula species, based on K2P model from sequences of the COI gene, with number of sequences per species (n), the intraspecific variation, followed by the distance between species (in %).
Species |
n |
intraspecific variation |
2 |
3 |
4 |
5 |
6 |
7 |
8 |
9 |
10 |
11 |
H. guanheensis |
17 |
0.0 |
8.2 |
7.7 |
5.1 |
7.9 |
8.7 |
4.7 |
8.1 |
8.3 |
7.6 |
10.6 |
H. coccinocola (2) |
5 |
0.0 |
8.0 |
7.1 |
7.0 |
9.7 |
6.5 |
8.9 |
6.9 |
4.9 |
11.9 |
|
H. wuliangensis (3) |
3 |
0.0 |
6.5 |
7.3 |
9.8 |
6.7 |
8.9 |
7.5 |
6.3 |
10.5 |
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H. variegata (4) |
4 |
0.5 |
7.2 |
8.1 |
3.2 |
7.4 |
6.8 |
6.2 |
10.5 |
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H. pycnolepis (5) |
4 |
0.0 |
7.2 |
6.2 |
8.4 |
2.9 |
6.4 |
10.0 |
||||
H. potanini (6) |
3 |
0.9 |
7.2 |
8.0 |
7.4 |
8.4 |
10.2 |
|||||
H. longidorsalis (7) |
2 |
0.0 |
7.1 |
6.2 |
5.9 |
9.6 |
||||||
H. disparizona (8) |
4 |
0.1 |
8.3 |
8.3 |
10.0 |
|||||||
H. anguillioides (9) |
2 |
0.0 |
6.2 |
10.3 |
||||||||
H. change (10) |
5 |
0.0 |
11.1 |
|||||||||
S. latifasciata (11) |
1 |
Pairwise comparisons of genetic distance of six Homatula species, based on K2P model from sequences of the Cyt b gene, the other annotations being the same as Table
Species |
n |
intraspecific variation |
|||||||
12 |
13 |
14 |
15 |
16 |
17 |
18 |
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H. guanheensis |
17 |
0.1 |
4.8 |
10.3 |
9.9 |
4.2 |
5.8 |
9.5 |
15.8 |
H. variegata (12) |
4 |
0.1 |
9.8 |
9.3 |
1.6 |
4.9 |
9.3 |
14.7 |
|
H. pycnolepis (13) |
3 |
0.4 |
10.3 |
9.5 |
9.2 |
4.0 |
15.4 |
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H. potanini (14) |
5 |
0.1 |
8.7 |
8.7 |
9.2 |
13.6 |
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H. berezowskii (15) |
2 |
0.2 |
4.7 |
8.5 |
14.5 |
||||
H. longidorsalis (16) |
4 |
1.5 |
8.2 |
14.3 |
|||||
H. acuticephala (17) |
5 |
0.0 |
14.3 |
||||||
S. latifasciata (18) |
1 |
New species were collected from the Guanhe River of the Hanjiang River drainage (the middle and lower reaches of the Yangtze River). In terms of geographic distribution, the known species in the Yangtze River Basin are as follows: Homatula variegata, H. wujiangensis, H. berezowskii and H. potanini. H. variegata and H. potanini are mainly distributed in the upper reaches of the Yangtze River, while H. berezowskii is mainly distributed in the Jialingjiang River and the Hanjiang River (
According to the results of the phylogenetic tree, H. variegata, H. berezowskii, H. longidorsalis and the new species gather into a branch, H. longidorsalis and H. guanheensis + (H. berezowskii + H. variegata) form a sister group (supported by the Cyt b gene, Fig.
This work was supported by the following funding: The National Natural Science Foundation of China (31872199, U2004146), Henan Province Department of Science and Technology (182102110046), The Training Plan for Young Teachers in Colleges and Universities of Henan Province (2019GGJS063). Study support was provided by The High-Performance Computing Center of Henan Normal University. Thanks to Prof. E. Zhang from Institute of Hydrobiology, Chinese Academy of Sciences, Wuhan (IHB) for providing morphological data of H. laxiclathra.
Zhou Chuanjiang designed the study, analysed data and performed manuscript review, Ma Wenwen carried out the study, collected background information and drafted the manuscript. Zhou Chuanjiang, Meng Xiaolin, Wang Xi and Nie Guoxing collected samples in the field. All authors read and approved the final manuscript.
Morphometric characters of Homatula variegata, H. berezowskii, H. longidorsalis and H. laxiclathra from Gu and Zhang (2011), comparative materials with Table 2.