Zootaxa 3841 (2): 257–270 www.mapress.com /zootaxa / Copyright © 2014 Magnolia Press
Article
ISSN 1175-5326 (print edition)
ZOOTAXA
ISSN 1175-5334 (online edition)
http://dx.doi.org/10.11646/zootaxa.3841.2.5 http://zoobank.org/urn:lsid:zoobank.org:pub:B8B9CDD0-902A-40C8-9A3C-CEA95F77B458
Paraqianlabeo lineatus, a new genus and species of labeonine fishes (Teleostei: Cyprinidae) from South China HAI-TAO ZHAO1,2, JOHN P. SULLIVAN3, YAO-GUANG ZHANG1 & ZUO-GANG PENG1 1
Key Laboratory of Freshwater Fish Reproduction and Development (Ministry of Education), Southwest University School of Life Science, Beibei, Chongqing 400715, China. E-mail: [email protected]; [email protected] 2 School of Environment and Life Science, Bijie University, Bijie, Guizhou 551700, China 3 Cornell University Museum of Vertebrates, 159 Sapsucker Woods Road, Ithaca, New York 14850 USA. E-mail: [email protected]
Abstract Paraqianlabeo, new genus, is described from the Chishui-He and Wu-Jiang rivers of the Yangtze River basin in Guizhou Province, South China. This new genus is distinguished from all other labeonine genera by a combination of morphological and molecular evidence. It is distinguished from all other Asian garrains by a unique oromandibular morphology; i.e., an arched, wide, rudimentary sucker formed by raised front and lateral margins of lower lip, postlabial groove and mental groove between middle and lateral lobes of lower lip chevron-shaped; anteroventral margin of rostral cap arched, thin and papillose; rostral cap not overlying upper lip, upper lip connected with lower lip around corners of mouth; well developed upper lip free from upper jaw; lower lip divided into two lateral fleshy lobes and one central plate, these two lateral fleshy lobes small and short, median lobe of lower lip large; anterior and anterolateral edges of upper and lower lip finely papillose; shallow, straight groove between lower lip and lower jaw; lower jaw bears thin, cornified cutting edge. Additionally, scales on mid-ventral region from pectoral fins to pelvic fins subcutaneous and half-hidden; dorsal fin with 3 simple and 7½-8 branched rays. Paraqianlabeo lineatus, new species, type species of this genus, has longitudinal dark stripe along side of body. Analyses of mitochondrial DNA data indicate that this new genus forms a highly diverged lineage within the Garraina group of Labeoninae. Key words: taxonomy, Garraina, new genus, Yangtze River drainage, Guizhou Province
Introduction The Labeonini (sensu Reid 1982; Stiassny & Getahun 2007) is composed of a large number of cyprinid genera from the freshwaters of tropical Africa and Asia (Zhu et al. 2011). Labeonins are widely distributed in rivers and streams, and most species are adapted to rapidly flowing waters. Currently, this group is represented in Southeast Asia and South China by approximately 36 genera (Zhang & Zhou 2012). Most of the generic-level diversity of the Labeonini is concentrated in South China where 23 genera are found. Recently, four new genera, Protolabeo, Cophecheilus, Sinigarra and Stenorynchoacrum, have been discovered and described, totaling 27 genera in China (An et al. 2010, Zhu et al. 2011, Zhang & Zhou 2012, Huang et al. 2014), of which 13 are endemic. These taxa are distinguished from other cyprinid fish groups by a high degree of morphological modification of their oromandibular structures (Zhang et al. 2000). Consequently, the taxonomy of the Labeonini at the generic-level rank is poorly understood. This tribe is the subject of recent molecular phylogenetic analyses that seek to better understand their relationships and to develop more accurate taxonomic classifications (Zheng et al. 2010; Yang & Mayden 2010; Yang et al. 2012). During recent fieldwork in Guizhou Province from May to September 2013, several specimens were sampled from the Wu-Jiang and Chishui-He rivers of the upper Yangtze River basin. The new specimens had distinct phenotypes and could not be assigned confidently to any genus. These observations combined with the morphological and molecular analyses of previous studies (Zhang et al. 2000; Zhang & Chen 2004; Zhang & Chen 2006; Zhang et al. 2006; Zhang et al. 2008; Zhang & Zhou 2012), convince us that a new genus is warranted for these specimens, which is described herein. Accepted by L. Page: 25 Jun. 2014; published: 25 Jul. 2014
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FIGURE 1. Map showing the distribution of Paraqianlabeo lineatus in Guizhou Province of South China.
Material and methods Taxon sampling Twenty-two specimens were collected from the Wu-Jiang and Chishui-He rivers of the upper Yangtze River basin at Wangcao, Suiyang County; Shuikan, Tongzi County; and Jingmen, Jinsha County; in Guizhou Province, China (Fig. 1). Following euthanization, tissues were taken from adult specimens and preserved in 95% ethanol. Voucher specimens were fixed in 10% buffered formalin, and later transferred to 70% ethanol. The holotype (SWU 20130710005) and paratypes (SWU 20130710006-22) are deposited in the Museum of the Key Laboratory of Freshwater Fish Reproduction and Development (Ministry of Education), Southwest University, China. The remaining four paratypes are deposited in the Cornell University Museum of Vertebrates (CUMV 98006).
Morphological examinations Measurements were taken point to point with digital calipers and recorded to the nearest 0.1 mm. Counts and measurements were made on the left side whenever possible, following Kottelat (2001). Predorsal, prepectoral, prepelvic and preanal lengths were taken, respectively, from the anteriormost tip of the snout to the dorsal-,
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pectoral-, pelvic- and anal-fin origins. Interorbital width was measured as the shortest distance between the upper margins of the orbits. Pelvic fin and caudal fin rays were counted following Roberts (1989). Description of pharyngeal teeth follow Hubbs & Lagler (1947). Subunits of the head are given as proportions of the head length (HL). Head length and measurements of other parts of the body are expressed as percentages of the standard length (SL).
DNA extraction and sequencing Total genomic DNA was extracted using the phenol-chloroform method (Sambrook et al. 1989). A 588 bp fragment of the COI [cytochrome oxidase gene subunit I] was amplified using polymerase chain reaction (PCR) with primers COI–L5956: 5'-CACAAAGACATTGGCACCCT-3' and COI–H6855: 5'AGTCAGCTGAAKACTTTTAC-3' (Peng et al. 2009). Standard polymerase chain reactions (PCR) were performed in a final volume of 30 μl with the following procedures: initial denaturation at 94°C for 4 min, 35 amplification cycles at 94°C for 1 min, annealing for 40 sec at 52°C, extension for 1 min at 72°C, and final extension at 72°C for 10 min. After amplification, a 2 μL sample of each PCR product was visualized on 0.8% lowmelting agarose gels; the remaining product was sequenced directly by a commercial sequencing corporation.
Phylogenetic analysis In total, 27 DNA sequences (see below) were used for the reconstruction of phylogenetic relationships, including three samples of the new species. Most of the known genera in China were included except for Cophecheilus, Sinigarra, Protolabeo, Sinilabeo and Longanalus (Table 1), whose tissue samples were not available. The sequences of Opsariichthys uncirostris and Catlocarpio siamensis were chosen as outgroup taxa based on the studies of Yang et al. (2012). Voucher specimens, and their localities and GenBank accession numbers are given in Table 1. CLUSTALX v1.83 (Thompson et al. 1997) was used to perform the sequence alignment, with parameters set at default. Genetic divergences were estimated with MEGA 5.0 (Tamura et al. 2011) by calculating the Kimura 2-parameter distance (K2P; Kimura 1980). Phylogenetic analyses were performed using neighbor-joining (NJ) tree. A rooted NJ analysis based on K2P distances model was run using MEGA 5.0.
Description of the new genus Paraqianlabeo, new genus Type species: Paraqianlabeo lineatus, new species (Figs. 2, 4) Diagnosis. Paraqianlabeo can be distinguished from all other genera of the Garraina (sensu Stiassny & Getahun 2007) by its uniquely modified oromandibular morphology; i.e., arched, wide, rudimentary sucker formed by raised front and lateral margins of lower lip, postlabial groove and mental groove between middle and lateral lobes of lower lip chevron-shaped (Fig. 2). The genus is further distinguished by having a unique combination of the following characters: anteroventral margin of rostral cap arched, thin and papillose; rostral cap not overlying upper lip, upper lip connected with lower lip around corners of mouth; well developed upper lip free from upper jaw; lower lip divided into two lateral fleshy lobes and one central plate, these two lateral fleshy lobes small and short, median lobe of lower lip large; anterior and anterolateral edges of upper and lower lip finely papillose; shallow, straight groove between lower lip and lower jaw; lower jaw bearing thin, cornified cutting edge; scales in ventral medial region from pectoral fins to pelvic fins half-hidden and subcutaneous; 3 scale rows below lateral line; 7½-8 branched dorsal-fin rays. Etymology. The generic name is from a combination of the Latin word Para (beside; beyond or outside; associated with) and Qian (abbreviation for Guizhou Province), where the type species was collected, and the Latin word Labeo, a generic name and commonly utilized suffix for Labeoninae genera. Gender: masculine.
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TABLE 1. Labeonine mitochondrial cytochrome oxidase subunit I (COI) sequence samples used in the present study. Specimen
Voucher
Locality
GenBank No.
Source
Akrokolioplax bicornis
KIZ05207
Bangana lippa
CTOL3846
China: Lushui, Yunnan
GU086590
Zheng et al. (2010)
China: Mengna, Yunnan
JX074198
Yang et al. (2012)
China: Hekou, Yunnan
GU086576
Zheng et al. (2010)
Ingroup
Cirrhinus molitorella KIZCXY20071011 Crossocheilus reticulatus
CBM ZF-11235
Cambodia: Market, Kampong Chhnang
HM536925
Discogobio yunnanensis
KIZ2006003377
China: Jinning, Yunnan
GU086583
Epalzeorhynchos tum
frena- CTOL01577
Zheng et al. (2010) Yang et al. (2012)
JX074170 Garra orientalis
IHBCY0403441
China: Tengchong, Yunnan
HM536884 Yang & Mayden (2010)
Labeo yunnanensis
CTOL3956
China: Ledong, Hainan
JX074205
Yang et al. (2012)
Hongshuia microstomatus KIZMR200800576 China: Lingyun, Guangxi
GU086586
Zheng et al. (2010)
Labiobarbus leptochilus
HM536885
Yang & Mayden (2010)
IHBCY0407001
China: Mengna, Yunnan
Lobocheilos melanotaenia CTOL03415
Vietnam
Yang et al. (2012)
Osteochilus salsburyi
IHBCY0308001
China: Rong’an, Guangxi
HM536883
Yang & Mayden (2010)
Parasinilabeo assimilis
CTOL3837
China: Rong’an, Guangxi
JX074191
Yang et al. (2012)
CTOL3841
China: Nanchong, Sichuan
JX074193
Yang et al. (2012)
China: Du’an, Guangxi
JX074192
Yang et al. (2012)
JX074207
Pseudogyrinocheilus prochilus Sinocrossocheilus aensis
bam- CTOL3885
Ptychidio jordani
CTOL3839
China: Tian’e, Guangxi
JX074194
Yang et al. (2012)
Rectoris posehensis
KIZMR20080065
China: Libo, Guizhou
GU086595
Zheng et al. (2010)
Semilabeo notabilis
CTOL3867
China: Du’an, Guangxi
JX074195 Yang et al. (2012)
Stenorynchoacrum xijian- KIZDLN20080001 China: Guilin, Guangxi gensis
GU086597
Zheng et al. (2010)
Sinocrossocheilus labia- KIZZLP20090021 tus
GU086596
Zheng et al. (2010)
China: Tongzi, Guizhou
‘Qianlabeo striatus’
KIZMR20080505
China: Zunyi, Guizhou
GU086588
Zheng et al. (2010)
Qianlabeo
SWU20130917002 China: An’shun, Guizhou
KJ960281
This study
Paraqianlabeo lineatus
SWU20130618002 China: Jinsha, Guizhou
KJ960279
This study
Paraqianlabeo lineatus
SWU20130711001 China: Shuiyang, Guizhou
KJ960280
striatus
This study Paraqianlabeo lineatus
SWU20130712001 China: Tongzi, Guizhou
KJ960278
This study
Outgroup Opsariichthys uncirostris
CTOL444
Mayden et al. (2007) EF452894
Catlocarpio siamensis
CBM ZF-11224
Cambodia: Landing port, Kampong Chh- HM536911 nang
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TABLE 2. Morphometric data for Paraqianlabeo lineatus. Characters
Holotype SWU 20130710005
Paratypes (21) Range
Mean
SD
Standard length (mm)
65.1
62.8-80.8
67.3
4.16
21.6
19.8-24.4
21.6
1.12
Percentage of standard length Body depth Head length
24.0
22.4-24.9
23.5
0.80
Caudal-peduncle length
17.2
16.8-19.9
18.7
0.87
Caudal-peduncle depth
12.2
10.9-12.3
11.7
0.41
Dorsal length
20.6
18.5-21.2
19.6
0.86
Pectoral length
17.4
15.8-18.1
17.2
0.61
Ventral length
17.0
15.2-17.1
15.9
0.57
Anal length
14.8
13.9-16.8
14.9
0.93
Predorsal length
51.7
49.5-52.6
50.7
0.96
Prepectoral length
23.3
21.2-23.6
22.4
0.71
Preventral length
53.2
49.9-53.9
52.0
1.06
Preanal length
74.6
71.8-75.4
73.5
1.26
Percentage of head length Head depth
61.9
60.4-72.0
63.3
2.98
Head width
72.0
64.7-72.0
68.3
2.35
Snout length
41.2
33.7-42.2
37.5
1.97
Eye diameter
22.0
21.1-24.3
22.6
0.96
Postorbial length
25.1
20.0-30.4
24.7
2.24
Paraqianlabeo lineatus, new species (Fig. 2; Table 2) Holotype. SWU 20130710005, 65.1mm SL, a tributary flowing into Furong-Jiang of Wu-Jiang (Yangtze River) drainage at Wangcao, Suiyang County, Guizhou, South China, (28°13' N 107°13' E); coll. H.T. Zhao, July 2013. Paratypes (21). CUMV 98006 (4), SWU 20130710006–22 (17), 62.8–80.8 mm SL, same data as holotype. Diagnosis. A species of Paraqianlabeo with longitudinal dark stripe along side of body (Fig. 3); for additional features see the generic diagnosis. Description. Whole body and morphology of oromandibular structures are illustrated in Figs. 2 and 4, respectively. Morphometric data for 22 type specimens are given in Table 2. A small-sized fish attaining length of up to 81 mm, body elongate and compressed, greatest depth at dorsal-fin origin. Caudal peduncle slender with smallest depth close to base of caudal fin. Dorsal profile of body gradually ascending from snout tip to dorsal-fin origin; from there to origin of dorsal procurrent caudal-fin rays profile slightly concave. Ventral profile from tip of snout to pectoral-fin insertion nearly straight; convex from there to anal-fin origin. Anal-fin base straight; slightly concave from posterior end of anal-fin base to origin of ventral procurrent caudal-fin rays. In size and shape of head, body and fins, it is most similar to another diminutive labeoninae Qianlabeo striatus Zhang & Chen, 2004, endemic to the Beipan-Jiang tributary (Fig. 4). Head moderately large, longer than wide; interorbital space slightly concave. Eye medium-sized, dorsolaterally placed in anterior half of head. Snout blunt in lateral view and pointed in dorsal view, with shallow sublachrymal groove extending obliquely to corner of mouth; no tubercles on tip. Two pairs of barbels; rostral pair at anterior end of sublachrymal groove and well-developed maxillary pair at corner of mouth, longer than eye diameter but seven-tenths as long as rostral pair. Three rows of pharyngeal teeth, tooth pattern 5,4,2–2,4,5, with compressed and pointed distal tips (Fig. 5a). Air bladder bipartite, anterior chamber oval, wider than posterior one; posterior chamber stick-like or elongate with an enlarged, pointed distal end, about twice as long as anterior chamber (Fig. 5b). Gill rakers on outer side of first gill arch 14–16, short and small. Mouth inferior and arched.
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Peritoneum black.
FIGURE 2. Oromanibular structures of Paraqianlabeo lineatus, SWU 20130710005, holotype, 65.1 mm SL. lj = lower jaw; lll = lateral lobe of lower lip; mb = maxillary barbel; mg = mental groove; ml l = median lobe of lower lip; pg = postlabial groove; rb = rostral barbel; rc = rostral cap; uj = upper jaw; ul = upper lip.
Body scales moderate in sized; ventral scales between pectoral fins and pelvic fins half-hidden and subcutaneous. Lateral line complete, horizontal, 39–42 scales; 5 scale rows above lateral line and 3 below. Predorsal midline scales very much smaller than ones on flank, irregularly arranged and embedded under skin anteriorly, difficult to count. Circumpeduncular scale rows 15–17. Axillary scale present at pelvic-fin base. Two scales between vent and anal-fin origin. Dorsal-fin with 3 unbranched and 7½–8 branched rays, origin slightly nearer to tip of snout than to caudal-fin base; last simple ray soft, equal to or somewhat longer than HL and last branched ray split; distal margin slightly concave. Dorsal-fin origin roughly at midpoint of body, while pelvic-fin origin slightly posterior. Pectoral fin falcate, no longer than HL, extending slightly beyond midway to pelvic-fin insertion and as far as sixth or seventh scale anterior to pelvic fin insertion; with one unbranched and 13 or 14 branched rays. Pelvic fin with one unbranched and 8 or 9 branched rays, inserted vertically posterior to second branched dorsal-fin ray base, and extending to vent, but not to anal-fin origin. Anal fin with 3 unbranched and 5 branched rays, last one split to base; distal margin slightly concave; origin equidistant between pelvic-fin insertion and caudal-fin base. Caudal deeply forked; upper and lower lobes equal in length and similarly shaped, with tapering, rounded tips.
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FIGURE 3. Ventral view of oromandibular structures in: (a) Qianlabeo striatus, SWU 20130917005, 68.1mm SL, BeipanJiang of Pearl River drainage at An’shun County, Guizhou Province; (b) Sinocrossocheilus labiatus, SWU 20121120001, 60.1 mm SL, Chishui-He River of Yangtze River basin at Bijie City, Guizhou Province; (c) Akrokolioplax bicornis, SWU 20071115020, 102 mm SL, Nujiang River drainage at Baoshan City, Yunnan Province; (d) Crossocheilus burmanicus SWU 201305110001, 79.4 mm SL, Binlang-Jiang of Irrawaddy River drainage at Tengchong County, Yunnan Province; (e) Sinocrossocheilus bamaensis, SWU 20131124001, 103.5mm SL, Hongshui-He of Pearl River drainage at Wangmo County, Guizhou Province; (f) Rectoris posehensis, SWU 20051114006, 107.4mm SL, Yangtze River drainage at Kaixian County, Chongqing.
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FIGURE 4. Lateral view of (a) Paraqianlabeo lineatus, SWU 20130710007, paratype, 65.1mm SL and (b) Qianlabeo striatus, SWU 20130917005, 68.1mm SL, Beipan-Jiang drainage at An’shun County, Guizhou Province.
Color pattern in formalin In formalin-preserved specimens, top of head grayish brown, snout and lip grayish; cheek with sparse brownish pigments; opercle and subopercle brownish. Body gray and yellowish dorsally and laterally, slightly pale ventrally. A longitudinal dark stripe originating near upper extremity of gill opening and terminating medially at caudal-fin base, extends anteriorly along first longitudinal scale row above lateral line and posteriorly along lateral line, slightly wider and more prominent posteriorly, especially at the end of caudal base. All fins yellowish with black speckles on it. Dorsal, caudal, pectoral and pelvic-fins with dark chromatophores on rays, conferring a dusky appearance. Distribution. Known from tributaries of the Wu-Jiang and Chishui-He rivers within the upper Yangtze River basin at Wangcao (28°13' N 107°13' E), Suiyang County; Shuikan (28°4' N 106°46' E), Tongzi County; and Jingmen (27°32' N 105°59' E), Jinsha County, Guizhou Province, South China (Fig. 1).
Habitat and ecology Paraqianlabeo lineatus was collected in clearwater hill streams with sluggish flow over gravel substrate (Fig. 6). The catchment of this tributary is endowed with numerous hill streams with substrates of stone. The width of the river channel was about 5–6 meters and water depth approximately 60 cm. Individuals of Paraqianlabeo lineatus swim slowly and hide among the rocks. Only two other species were collected with them: Carassius auratus (Cyprinidae) and Pseudorasbora parva (Cyprinidae). Etymology. The specific epithet is from the Latin lineatus, meaning lined, referring to the longitudinal dark stripe along the side of the body.
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0.145 0.182
0.186
0.192
0.202
0.166
0.159
0.148
0.160
0.169
0.144
0.143
0.139
0.157
0.164
0.148
0.148
0.158
0.144
0.146
0.144
0.131
0.155
0.146
0.180
0.183
0.159
0.159
0.154
0.129
0.151
0.163
0.133
0.159
0.156
0.175
0.144
0.167
0.151
0.156
0.151
0.147
0.147
0.133
2
0.161 0.170
0.153
1
0.147
0.133
0.149
0.142
0.150
0.155
0.140
0.147
0.136
0.143
0.144
0.135
0.157
0.139
0.128
0.152
0.174
0.141
0.178
0.157
0.138
0.152
3
0.183
0.165
0.176
0.164
0.166
0.152
0.163
0.173
0.161
0.190
0.151
0.163
0.173
0.161
0.152
0.166
0.166
0.150
0.183
0.173
0.151
4
0.133
0.135
0.141
0.126
0.150
0.135
0.132
0.150
0.125
0.152
0.142
0.126
0.157
0.153
0.132
0.144
0.137
0.141
0.174
0.154
5
0.142
0.151
0.162
0.151
0.146
0.144
0.169
0.169
0.141
0.163
0.144
0.144
0.151
0.133
0.151
0.142
0.167
0.159
0.190
6
0.183
0.164
0.178
0.180
0.207
0.183
0.184
0.189
0.184
0.197
0.200
0.183
0.212
0.197
0.187
0.185
0.162
0.184
7
0.135
0.127
0.148
0.146
0.160
0.139
0.137
0.155
0.134
0.150
0.141
0.139
0.162
0.137
0.131
0.135
0.125
8
0.128
0.136
0.123
0.140
0.147
0.145
0.142
0.151
0.111
0.136
0.129
0.120
0.150
0.132
0.123
0.134
9
0.084
0.121
0.088
0.070
0.094
0.076
0.096
0.088
0.144
0.139
0.092
0.078
0.092
0.090
0.072
10
0.074
0.132
0.082
0.082
0.094
0.072
0.094
0.092
0.135
0.104
0.108
0.084
0.096
0.078
11
0.097
0.160
0.115
0.092
0.096
0.107
0.096
0.096
0.142
0.148
0.104
0.092
0.102
12
0.072
0.141
0.126
0.109
0.005
0.098
0.107
0.096
0.155
0.128
0.108
0.106
13
0.070
0.128
0.086
0.084
0.100
0.088
0.087
0.081
0.146
0.121
0.096
14
0.098
0.148
0.110
0.088
0.110
0.108
0.104
0.106
0.138
0.125
15
0.119
0.155
0.121
0.126
0.121
0.114
0.130
0.128
0.152
16
0.149
0.123
0.146
0.153
0.153
0.146
0.141
0.164
17
0.088
0.143
0.090
0.094
0.090
0.083
0.047
18
0.094
0.125
0.096
0.092
0.100
0.091
19
0.074
0.136
0.096
0.104
0.096
20
0.070
0.139
0.120
0.103
21
0.086
0.121
0.082
22
0.088
0.121
23
0.120
24
Note. 1. Labeo yunnanensis, 2. Osteochilus salsburyi, 3. Labiobarbus leptochilus, 4. Catlocarpio siamensis, 5. Lobocheilos melanotaenia, 6. Epalzeorhynchos frenatum, 7. Opsariichthys uncirostris, 8. Garra orientalis, 9. Akrokolioplax bicornis, 10. Sinocrossocheilus bamaensis, 11. Ptychidio jordani, 12. Semilabeo notabilis, 13. Paraqianlabeo, 14. Parasinilabeo assimilis, 15 Bangana lippa, 16. Qianlabeo, 17. Crossocheilus reticulatus, 18. Hongshuia microstomatus, 19. Rectoris posehensis, 20. Stenorynchoacrum xijiangensis, 21. ‘Qianlabeo’ (sensu Zheng et al., 2010) [=Paraqianlabeo lineatus], 22. Discogobio yunnanensis, 23. Sinocrossocheilus labiatus, 24. Cirrhinus molitorella, 25. Pseudogyrinocheilus prochilus.
25
1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24
TABLE 3. K2P distances of the mitochondrial COI sequences between species of labeonines.
FIGURE 5. (a) Left side pharyngeal tooth plate in dorsomedial view, and (b) gas bladder in ventral view of Paraqianlabeo lineatus.
Discussion There are three basic patterns of oromandibular structures among Southeast and East Asian genera of the Labeonini (Zhang & Chen 2004; Zhang et al. 2006; Zhang et al. 2008). Paraqianlabeo belongs to the third group, the subtribe Garraina (= Garrae sensu Rainboth 1996), that have a well-developed or reduced upper lip that is separated from the upper jaw by a deep groove. This group comprises the following genera: Garra, Crossocheilus, Epalzeorhynchos, Parasinilabeo, Sinocrossocheilus, Rectoris, Paracrossochilus, Pseudocrossocheilus, Discogobio, Discocheilus, Placocheilus, Pseudogyrinocheilus, Akrokolioplax, Semilabeo, Stenorynchoacrum and Hongshuia. Paraqianlabeo can be easily distinguished from Hongshuia, Discocheilus, Placocheilus and Discogobio by the presence of three (vs. two) rows of pharyngeal teeth. It also can be separated from Rectoris, Qianlabeo and Stenorynchoacrum by having mental grooves present (vs. absent). Six other garrain genera, Crossocheilus, Pseudocrossocheilus, Sinocrossocheilus, Epalzeorhynchos, Akrokolioplax and Rectoris, have rostral caps with fimbriate (grooved) ventral margins, while that of the new genus is not fimbriate, with no obvious grooves (Fig. 3). Further, the rostral fold and cap of the new genus differs from those of Semilabeo, Pseudogyrinocheilus and Parasinilabeo in being smooth vs. densely covered with papillae (Zhu et al. 2011; Fig.
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2). These three genera have rostral caps with a densely papillated ventral margin and a densely papillated lower lip; these characters all easily distinguish them from Paraqianlabeo. The detailed characters utilized to distinguish Paraqianlabeo from other related genera are given in diagnosis.
FIGURE 6. Type locality of Paraqianlabeo lineatus, Jingmen (Chishui-He) Village, Jinsha County, Guizhou Province, China. Photograph by H.T. Zhao.
Qianlabeo is very similar to Paraqianlabeo in having a longitudinal dark stripe along the side of the body and in body size, but Paraqianlabeo is conspicuously distinct from Qianlabeo in having a suctorial lower lip (vs. absent), a short postlabial groove (vs. long), a mental groove present (vs. absent), and development of papillae on the rostral fold and a complete upper lip (vs. incomplete and restricted to the side of upper jaw). Four sequences were newly determined and deposited in GenBank (Table 1). All the 27 aligned sequences of the COI contain 588 nucleotide sites, of which 215 were variable and 176 potentially parsimony-informative. No indels were observed, and no stop codons were found in protein coding regions, suggesting that nuclear pseudogenes were not sequenced. The genetic distance (K2P) between Paraqianlabeo and other labeonine genera ranges from 6.9% to 19.0% (Table 3); the lowest pairwise genetic distance is between Pseudogyrinocheilus and Paraqianlabeo. The average genetic distance of three geographic populations of Paraqianlabeo is less than 2% and within the range typical for intra-specific variation in other cyprinids (Zhao et al., unpublished data). The NJ tree is shown in Fig. 7. The monophyly of labeonine was recovered, consistent with findings in previous studies (e.g., Zheng et al. 2010; Yang et al. 2012). Among Paraqianlabeo, individuals from the three locations in northern Guizhou Province formed a monophyletic group with strong support (BPP = 100; Fig. 7). A specimen identified as ‘Qianlabeo striatus’ (sequence data from Zheng et al. 2010) that falls within the Paraqianlabeo group in the tree was not originally collected from its type locality Beipan-Jiang of Pearl River drainage, Guizhou Province, but from the Yangtze River drainage in Zunyi City, Guizhou Province. This ‘Qianlabeo striatus’ specimen was most probably misidentified by the authors (Zheng et al. 2010) and should be
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assigned to Paraqianlabeo. The genus Qianlabeo (sensu Zhang & Chen 2004) is situated between the Labeonini (= Labeonina) and the Garrini (= Garraina) based on a Qianlabeo sequenced from its type locality (Beipan-Jiang of the Pearl River drainage in the central region of Guizhou Province), consistent with the position suggested by Zhang & Chen (2004). Taken together, our data suggest that Qianlabeo is not distributed in the Yangtze River drainage, but is restricted to its type locality, the Beipan-Jiang tributary of the Pearl River drainage in the central region of Guizhou Province.
FIGURE 7. Neighbor-joining (NJ) tree of labeonine species and two outgroup species, Opsariichthys uncirostris and Catlocarpio siamensis, based on mitochondrial COI sequences. Node values = bootstrap test (1,000 pseudoreplicates), low supports (< 50%) were not shown.
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Comparative material Qianlabeo striatus: SWU 20130917001–012, 54.7–80.8mm SL, Beipan-Jiang of Pearl River drainage at An’shun County, Guizhou Province; Sinocrossocheilus labiatus: SWU 20121120001–010, 53.2–60.0 mm SL, Chishui-He River of Yangtze River basin at Bijie City, Guizhou Province; Akrokolioplax bicornis: SWU 20071115020, 102 mm SL, Nujiang River drainage at Baoshan City, Yunnan Province; Crossocheilus burmanicus: SWU 201305110001, 79.4 mm SL, Binlang-Jiang of Irrawaddy River drainage at Tengchong County, Yunnan Province; Sinocrossocheilus bamaensis: SWU 20131124001–011, 54.7–123.5 mm SL, Hongshui-He of Pearl River drainage at Wangmo County, Guizhou Province; Rectoris luxiensis: SWU 20051114006, 107.4 mm SL, Jialing-Jiang of upper Yangtze River drainage at Kaixian County, Chongqing; Bangana rendahli: SWU 20130804001, 123.5 mm SL, Wu-Jiang of upper Yangtze River drainage at Wuchuan County, Zunyi City, Guizhou Province; Discogobio yunnanensis: SWU 20130806002–008, 70.3–95.1 mm SL, Wu-Jiang of upper Yangtze River drainage at Tongzi County, Zunyi City, Guizhou Province; Pseudogyrinocheilus prochilus: SWU 20130804003, 134.6 mm SL, Wu-Jiang of upper Yangtze River drainage at Wuchuan County, Zunyi City, Guizhou Province; Sinilabeo hummeli: SWU 20080410001, 161.1 mm SL, Jialing-Jiang of upper Yangtze River drainage at Hechuan County, Chongqing; Ptychidio jordani: SWU 020121201089, 84.8 mm SL; China: Guangxi Province, Hongshui-He of Pearl River drainage at Du’an County.
Acknowledgments We are indebted to Drs. Z. J. Wang (Southwest University) and H. P. Zhao (Henan University) for their help with specimen collection. We are also grateful to L. Y Wang for his help with preparing photographs. This work was supported by grants from the Fundamental Research Funds for the Central Universities of China (XDJK2014c003) and the National Natural Science Foundation of China (31272283).
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Natural History, 16, 497–510. http://dx.doi.org/10.1080/00222938200770401 Roberts, T.R. (1989) The freshwater fishes of western Borneo (Kalimantan Barat, Indonesia). Memoirs of the California Academy Sciences, 14, 1–210. Sambrook, J., Fritsch, E.F. & Maniatis, T. (1989) Molecular Cloning: A Laboratory Manual, 2nd edition. Cold Spring Harbor Laboratory Press, New York, 1626 pp. Stiassny, M.L.J. & Getahun, A. (2007) An overview of labeonin relationships and the phylogenetic placement of the AfroAsian genus Garra Hamilton, 1922 (Teleostei: Cyprinidae), with the description of five new species of Garra from Ethiopia, and a key to all African species. Zoological Journal of the Linnean Society, 150, 41–83. http://dx.doi.org/10.1111/j.1096-3642.2007.00281.x Tamura, K., Peterson, D., Peterson, N., Stecher, G., Nei, M. & Kumar, S. (2011) MEGA5: Molecular evolutionary genetics analysis using maximum likelihood, evolutionary distance, and maximum parsimony methods. Molecular Biology and Evolution, 28, 2731–2739. http://dx.doi.org/10.1093/molbev/msr121 Thompson, J.D., Gibson, T.J., Plewniak, F., Jeanmougin, F. & Higgins, D.G. (1997) The CLUSTAL_X windows interface: Flexible strategies for multiple sequence alignment aided by quality analysis tools. Nucleic Acids Research, 25, 4876–4882. http://dx.doi.org/10.1093/nar/25.24.4876 Yang, L. & Mayden, R.L. (2010) Phylogenetic relationships, subdivision, and biogeography of the cyprinid tribe Labeonini (sensu Rainboth, 1991) (Teleostei: Cypriniformes), with comments on the implications of lips and associated structures in the labeonin classification. Molecular Phylogenetics and Evolution, 54, 254–265. http://dx.doi.org/10.1016/j.ympev.2009.09.027 Yang, L., Arunachalam, M., Sado, T., Levin, B.A., Golubtsov, A.S., Freyhof, J., Friel, J.P., Chen, W.-J., Hirt, M.V., Manickam, R., Agnew, M.K., Simons, A.M., Saitoh, K., Miya, M., Mayden, R.L. & He, S. (2012) Molecular phylogeny of the cyprinid tribe Labeonini (Teleostei: Cypriniformes). Molecular Phylogenetics and Evolution, 65, 362–379. http://dx.doi.org/10.1016/j.ympev.2012.06.007 Zhang, E. & Zhou, W. (2012) Sinigarra napoense, a new genus and species of labeonin fishes (Teleostei: Cyprinidae) from Guangxi Province, South China. Zootaxa, 3586, 17–25. Zhang, E. & Chen, Y.Y. (2004) Qianlabeo striatus, a new genus and species of Labeoninae from Guizhou Province, China (Teleostei: Cyprinidae). Hydrobiologia, 527, 25–33. http://dx.doi.org/10.1023/b:hydr.0000043315.64357.da Zhang, E. & Chen, Y.Y. (2006) Revised diagnosis of the genus Bangana Hamilton, 1822 (Pisces: Cyprinidae), with taxonomic and nomenclatural notes on its Chinese species. Zootaxa, 1281, 42–54. Zhang, E., Kullander, S.O. & Chen, Y.Y. (2006) Fixation of the type species of the genus Sinilabeo and description of a new species from the upper Yangtze River basin, China (Pisces: Cyprinidae). Copeia, 2006, 96–102. http://dx.doi.org/10.1643/0045-8511(2006)006[0096:fottso]2.0.co;2 Zhang, E., Xin, Q. & Lan, J.H. (2008) Description of a new genus and two new species of labeonine fishes from South China (Teleostei: Cyprinidae). Zootaxa, 1682, 33–44. Zhang, E., Yue, P.Q. & Chen, J.X. (2000) Labeoninae. In: Yue, P.Q. (Ed.), Fauna Sinica (Osteichthyes: Cypriniformes III). Science Press, Beijing, pp. 172–272. [in Chinese] Zheng, L.P., Yang, J.X., Chen, X.Y. & Wang, W.Y. (2010) Phylogenetic relationships of the Chinese Labeoninae (Teleostei, Cypriniformes) derived from two nuclear and three mitochondrial genes. Zoologica Scripta, 39, 559–571. http://dx.doi.org/10.1111/j.1463-6409.2010.00441.x Zhu, Y., Zhang, E., Zhang, M. & Han, Y.Q. (2011) Cophecheilus bamen, a new genus and species of labeonine fishes (Teleostei: Cyprinidae) from South China. Zootaxa, 2881, 39–50.
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Article
ISSN 1175-5326 (print edition)
ZOOTAXA
ISSN 1175-5334 (online edition)
http://dx.doi.org/10.11646/zootaxa.3841.2.5 http://zoobank.org/urn:lsid:zoobank.org:pub:B8B9CDD0-902A-40C8-9A3C-CEA95F77B458
Paraqianlabeo lineatus, a new genus and species of labeonine fishes (Teleostei: Cyprinidae) from South China HAI-TAO ZHAO1,2, JOHN P. SULLIVAN3, YAO-GUANG ZHANG1 & ZUO-GANG PENG1 1
Key Laboratory of Freshwater Fish Reproduction and Development (Ministry of Education), Southwest University School of Life Science, Beibei, Chongqing 400715, China. E-mail: [email protected]; [email protected] 2 School of Environment and Life Science, Bijie University, Bijie, Guizhou 551700, China 3 Cornell University Museum of Vertebrates, 159 Sapsucker Woods Road, Ithaca, New York 14850 USA. E-mail: [email protected]
Abstract Paraqianlabeo, new genus, is described from the Chishui-He and Wu-Jiang rivers of the Yangtze River basin in Guizhou Province, South China. This new genus is distinguished from all other labeonine genera by a combination of morphological and molecular evidence. It is distinguished from all other Asian garrains by a unique oromandibular morphology; i.e., an arched, wide, rudimentary sucker formed by raised front and lateral margins of lower lip, postlabial groove and mental groove between middle and lateral lobes of lower lip chevron-shaped; anteroventral margin of rostral cap arched, thin and papillose; rostral cap not overlying upper lip, upper lip connected with lower lip around corners of mouth; well developed upper lip free from upper jaw; lower lip divided into two lateral fleshy lobes and one central plate, these two lateral fleshy lobes small and short, median lobe of lower lip large; anterior and anterolateral edges of upper and lower lip finely papillose; shallow, straight groove between lower lip and lower jaw; lower jaw bears thin, cornified cutting edge. Additionally, scales on mid-ventral region from pectoral fins to pelvic fins subcutaneous and half-hidden; dorsal fin with 3 simple and 7½-8 branched rays. Paraqianlabeo lineatus, new species, type species of this genus, has longitudinal dark stripe along side of body. Analyses of mitochondrial DNA data indicate that this new genus forms a highly diverged lineage within the Garraina group of Labeoninae. Key words: taxonomy, Garraina, new genus, Yangtze River drainage, Guizhou Province
Introduction The Labeonini (sensu Reid 1982; Stiassny & Getahun 2007) is composed of a large number of cyprinid genera from the freshwaters of tropical Africa and Asia (Zhu et al. 2011). Labeonins are widely distributed in rivers and streams, and most species are adapted to rapidly flowing waters. Currently, this group is represented in Southeast Asia and South China by approximately 36 genera (Zhang & Zhou 2012). Most of the generic-level diversity of the Labeonini is concentrated in South China where 23 genera are found. Recently, four new genera, Protolabeo, Cophecheilus, Sinigarra and Stenorynchoacrum, have been discovered and described, totaling 27 genera in China (An et al. 2010, Zhu et al. 2011, Zhang & Zhou 2012, Huang et al. 2014), of which 13 are endemic. These taxa are distinguished from other cyprinid fish groups by a high degree of morphological modification of their oromandibular structures (Zhang et al. 2000). Consequently, the taxonomy of the Labeonini at the generic-level rank is poorly understood. This tribe is the subject of recent molecular phylogenetic analyses that seek to better understand their relationships and to develop more accurate taxonomic classifications (Zheng et al. 2010; Yang & Mayden 2010; Yang et al. 2012). During recent fieldwork in Guizhou Province from May to September 2013, several specimens were sampled from the Wu-Jiang and Chishui-He rivers of the upper Yangtze River basin. The new specimens had distinct phenotypes and could not be assigned confidently to any genus. These observations combined with the morphological and molecular analyses of previous studies (Zhang et al. 2000; Zhang & Chen 2004; Zhang & Chen 2006; Zhang et al. 2006; Zhang et al. 2008; Zhang & Zhou 2012), convince us that a new genus is warranted for these specimens, which is described herein. Accepted by L. Page: 25 Jun. 2014; published: 25 Jul. 2014
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FIGURE 1. Map showing the distribution of Paraqianlabeo lineatus in Guizhou Province of South China.
Material and methods Taxon sampling Twenty-two specimens were collected from the Wu-Jiang and Chishui-He rivers of the upper Yangtze River basin at Wangcao, Suiyang County; Shuikan, Tongzi County; and Jingmen, Jinsha County; in Guizhou Province, China (Fig. 1). Following euthanization, tissues were taken from adult specimens and preserved in 95% ethanol. Voucher specimens were fixed in 10% buffered formalin, and later transferred to 70% ethanol. The holotype (SWU 20130710005) and paratypes (SWU 20130710006-22) are deposited in the Museum of the Key Laboratory of Freshwater Fish Reproduction and Development (Ministry of Education), Southwest University, China. The remaining four paratypes are deposited in the Cornell University Museum of Vertebrates (CUMV 98006).
Morphological examinations Measurements were taken point to point with digital calipers and recorded to the nearest 0.1 mm. Counts and measurements were made on the left side whenever possible, following Kottelat (2001). Predorsal, prepectoral, prepelvic and preanal lengths were taken, respectively, from the anteriormost tip of the snout to the dorsal-,
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pectoral-, pelvic- and anal-fin origins. Interorbital width was measured as the shortest distance between the upper margins of the orbits. Pelvic fin and caudal fin rays were counted following Roberts (1989). Description of pharyngeal teeth follow Hubbs & Lagler (1947). Subunits of the head are given as proportions of the head length (HL). Head length and measurements of other parts of the body are expressed as percentages of the standard length (SL).
DNA extraction and sequencing Total genomic DNA was extracted using the phenol-chloroform method (Sambrook et al. 1989). A 588 bp fragment of the COI [cytochrome oxidase gene subunit I] was amplified using polymerase chain reaction (PCR) with primers COI–L5956: 5'-CACAAAGACATTGGCACCCT-3' and COI–H6855: 5'AGTCAGCTGAAKACTTTTAC-3' (Peng et al. 2009). Standard polymerase chain reactions (PCR) were performed in a final volume of 30 μl with the following procedures: initial denaturation at 94°C for 4 min, 35 amplification cycles at 94°C for 1 min, annealing for 40 sec at 52°C, extension for 1 min at 72°C, and final extension at 72°C for 10 min. After amplification, a 2 μL sample of each PCR product was visualized on 0.8% lowmelting agarose gels; the remaining product was sequenced directly by a commercial sequencing corporation.
Phylogenetic analysis In total, 27 DNA sequences (see below) were used for the reconstruction of phylogenetic relationships, including three samples of the new species. Most of the known genera in China were included except for Cophecheilus, Sinigarra, Protolabeo, Sinilabeo and Longanalus (Table 1), whose tissue samples were not available. The sequences of Opsariichthys uncirostris and Catlocarpio siamensis were chosen as outgroup taxa based on the studies of Yang et al. (2012). Voucher specimens, and their localities and GenBank accession numbers are given in Table 1. CLUSTALX v1.83 (Thompson et al. 1997) was used to perform the sequence alignment, with parameters set at default. Genetic divergences were estimated with MEGA 5.0 (Tamura et al. 2011) by calculating the Kimura 2-parameter distance (K2P; Kimura 1980). Phylogenetic analyses were performed using neighbor-joining (NJ) tree. A rooted NJ analysis based on K2P distances model was run using MEGA 5.0.
Description of the new genus Paraqianlabeo, new genus Type species: Paraqianlabeo lineatus, new species (Figs. 2, 4) Diagnosis. Paraqianlabeo can be distinguished from all other genera of the Garraina (sensu Stiassny & Getahun 2007) by its uniquely modified oromandibular morphology; i.e., arched, wide, rudimentary sucker formed by raised front and lateral margins of lower lip, postlabial groove and mental groove between middle and lateral lobes of lower lip chevron-shaped (Fig. 2). The genus is further distinguished by having a unique combination of the following characters: anteroventral margin of rostral cap arched, thin and papillose; rostral cap not overlying upper lip, upper lip connected with lower lip around corners of mouth; well developed upper lip free from upper jaw; lower lip divided into two lateral fleshy lobes and one central plate, these two lateral fleshy lobes small and short, median lobe of lower lip large; anterior and anterolateral edges of upper and lower lip finely papillose; shallow, straight groove between lower lip and lower jaw; lower jaw bearing thin, cornified cutting edge; scales in ventral medial region from pectoral fins to pelvic fins half-hidden and subcutaneous; 3 scale rows below lateral line; 7½-8 branched dorsal-fin rays. Etymology. The generic name is from a combination of the Latin word Para (beside; beyond or outside; associated with) and Qian (abbreviation for Guizhou Province), where the type species was collected, and the Latin word Labeo, a generic name and commonly utilized suffix for Labeoninae genera. Gender: masculine.
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TABLE 1. Labeonine mitochondrial cytochrome oxidase subunit I (COI) sequence samples used in the present study. Specimen
Voucher
Locality
GenBank No.
Source
Akrokolioplax bicornis
KIZ05207
Bangana lippa
CTOL3846
China: Lushui, Yunnan
GU086590
Zheng et al. (2010)
China: Mengna, Yunnan
JX074198
Yang et al. (2012)
China: Hekou, Yunnan
GU086576
Zheng et al. (2010)
Ingroup
Cirrhinus molitorella KIZCXY20071011 Crossocheilus reticulatus
CBM ZF-11235
Cambodia: Market, Kampong Chhnang
HM536925
Discogobio yunnanensis
KIZ2006003377
China: Jinning, Yunnan
GU086583
Epalzeorhynchos tum
frena- CTOL01577
Zheng et al. (2010) Yang et al. (2012)
JX074170 Garra orientalis
IHBCY0403441
China: Tengchong, Yunnan
HM536884 Yang & Mayden (2010)
Labeo yunnanensis
CTOL3956
China: Ledong, Hainan
JX074205
Yang et al. (2012)
Hongshuia microstomatus KIZMR200800576 China: Lingyun, Guangxi
GU086586
Zheng et al. (2010)
Labiobarbus leptochilus
HM536885
Yang & Mayden (2010)
IHBCY0407001
China: Mengna, Yunnan
Lobocheilos melanotaenia CTOL03415
Vietnam
Yang et al. (2012)
Osteochilus salsburyi
IHBCY0308001
China: Rong’an, Guangxi
HM536883
Yang & Mayden (2010)
Parasinilabeo assimilis
CTOL3837
China: Rong’an, Guangxi
JX074191
Yang et al. (2012)
CTOL3841
China: Nanchong, Sichuan
JX074193
Yang et al. (2012)
China: Du’an, Guangxi
JX074192
Yang et al. (2012)
JX074207
Pseudogyrinocheilus prochilus Sinocrossocheilus aensis
bam- CTOL3885
Ptychidio jordani
CTOL3839
China: Tian’e, Guangxi
JX074194
Yang et al. (2012)
Rectoris posehensis
KIZMR20080065
China: Libo, Guizhou
GU086595
Zheng et al. (2010)
Semilabeo notabilis
CTOL3867
China: Du’an, Guangxi
JX074195 Yang et al. (2012)
Stenorynchoacrum xijian- KIZDLN20080001 China: Guilin, Guangxi gensis
GU086597
Zheng et al. (2010)
Sinocrossocheilus labia- KIZZLP20090021 tus
GU086596
Zheng et al. (2010)
China: Tongzi, Guizhou
‘Qianlabeo striatus’
KIZMR20080505
China: Zunyi, Guizhou
GU086588
Zheng et al. (2010)
Qianlabeo
SWU20130917002 China: An’shun, Guizhou
KJ960281
This study
Paraqianlabeo lineatus
SWU20130618002 China: Jinsha, Guizhou
KJ960279
This study
Paraqianlabeo lineatus
SWU20130711001 China: Shuiyang, Guizhou
KJ960280
striatus
This study Paraqianlabeo lineatus
SWU20130712001 China: Tongzi, Guizhou
KJ960278
This study
Outgroup Opsariichthys uncirostris
CTOL444
Mayden et al. (2007) EF452894
Catlocarpio siamensis
CBM ZF-11224
Cambodia: Landing port, Kampong Chh- HM536911 nang
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TABLE 2. Morphometric data for Paraqianlabeo lineatus. Characters
Holotype SWU 20130710005
Paratypes (21) Range
Mean
SD
Standard length (mm)
65.1
62.8-80.8
67.3
4.16
21.6
19.8-24.4
21.6
1.12
Percentage of standard length Body depth Head length
24.0
22.4-24.9
23.5
0.80
Caudal-peduncle length
17.2
16.8-19.9
18.7
0.87
Caudal-peduncle depth
12.2
10.9-12.3
11.7
0.41
Dorsal length
20.6
18.5-21.2
19.6
0.86
Pectoral length
17.4
15.8-18.1
17.2
0.61
Ventral length
17.0
15.2-17.1
15.9
0.57
Anal length
14.8
13.9-16.8
14.9
0.93
Predorsal length
51.7
49.5-52.6
50.7
0.96
Prepectoral length
23.3
21.2-23.6
22.4
0.71
Preventral length
53.2
49.9-53.9
52.0
1.06
Preanal length
74.6
71.8-75.4
73.5
1.26
Percentage of head length Head depth
61.9
60.4-72.0
63.3
2.98
Head width
72.0
64.7-72.0
68.3
2.35
Snout length
41.2
33.7-42.2
37.5
1.97
Eye diameter
22.0
21.1-24.3
22.6
0.96
Postorbial length
25.1
20.0-30.4
24.7
2.24
Paraqianlabeo lineatus, new species (Fig. 2; Table 2) Holotype. SWU 20130710005, 65.1mm SL, a tributary flowing into Furong-Jiang of Wu-Jiang (Yangtze River) drainage at Wangcao, Suiyang County, Guizhou, South China, (28°13' N 107°13' E); coll. H.T. Zhao, July 2013. Paratypes (21). CUMV 98006 (4), SWU 20130710006–22 (17), 62.8–80.8 mm SL, same data as holotype. Diagnosis. A species of Paraqianlabeo with longitudinal dark stripe along side of body (Fig. 3); for additional features see the generic diagnosis. Description. Whole body and morphology of oromandibular structures are illustrated in Figs. 2 and 4, respectively. Morphometric data for 22 type specimens are given in Table 2. A small-sized fish attaining length of up to 81 mm, body elongate and compressed, greatest depth at dorsal-fin origin. Caudal peduncle slender with smallest depth close to base of caudal fin. Dorsal profile of body gradually ascending from snout tip to dorsal-fin origin; from there to origin of dorsal procurrent caudal-fin rays profile slightly concave. Ventral profile from tip of snout to pectoral-fin insertion nearly straight; convex from there to anal-fin origin. Anal-fin base straight; slightly concave from posterior end of anal-fin base to origin of ventral procurrent caudal-fin rays. In size and shape of head, body and fins, it is most similar to another diminutive labeoninae Qianlabeo striatus Zhang & Chen, 2004, endemic to the Beipan-Jiang tributary (Fig. 4). Head moderately large, longer than wide; interorbital space slightly concave. Eye medium-sized, dorsolaterally placed in anterior half of head. Snout blunt in lateral view and pointed in dorsal view, with shallow sublachrymal groove extending obliquely to corner of mouth; no tubercles on tip. Two pairs of barbels; rostral pair at anterior end of sublachrymal groove and well-developed maxillary pair at corner of mouth, longer than eye diameter but seven-tenths as long as rostral pair. Three rows of pharyngeal teeth, tooth pattern 5,4,2–2,4,5, with compressed and pointed distal tips (Fig. 5a). Air bladder bipartite, anterior chamber oval, wider than posterior one; posterior chamber stick-like or elongate with an enlarged, pointed distal end, about twice as long as anterior chamber (Fig. 5b). Gill rakers on outer side of first gill arch 14–16, short and small. Mouth inferior and arched.
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Peritoneum black.
FIGURE 2. Oromanibular structures of Paraqianlabeo lineatus, SWU 20130710005, holotype, 65.1 mm SL. lj = lower jaw; lll = lateral lobe of lower lip; mb = maxillary barbel; mg = mental groove; ml l = median lobe of lower lip; pg = postlabial groove; rb = rostral barbel; rc = rostral cap; uj = upper jaw; ul = upper lip.
Body scales moderate in sized; ventral scales between pectoral fins and pelvic fins half-hidden and subcutaneous. Lateral line complete, horizontal, 39–42 scales; 5 scale rows above lateral line and 3 below. Predorsal midline scales very much smaller than ones on flank, irregularly arranged and embedded under skin anteriorly, difficult to count. Circumpeduncular scale rows 15–17. Axillary scale present at pelvic-fin base. Two scales between vent and anal-fin origin. Dorsal-fin with 3 unbranched and 7½–8 branched rays, origin slightly nearer to tip of snout than to caudal-fin base; last simple ray soft, equal to or somewhat longer than HL and last branched ray split; distal margin slightly concave. Dorsal-fin origin roughly at midpoint of body, while pelvic-fin origin slightly posterior. Pectoral fin falcate, no longer than HL, extending slightly beyond midway to pelvic-fin insertion and as far as sixth or seventh scale anterior to pelvic fin insertion; with one unbranched and 13 or 14 branched rays. Pelvic fin with one unbranched and 8 or 9 branched rays, inserted vertically posterior to second branched dorsal-fin ray base, and extending to vent, but not to anal-fin origin. Anal fin with 3 unbranched and 5 branched rays, last one split to base; distal margin slightly concave; origin equidistant between pelvic-fin insertion and caudal-fin base. Caudal deeply forked; upper and lower lobes equal in length and similarly shaped, with tapering, rounded tips.
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FIGURE 3. Ventral view of oromandibular structures in: (a) Qianlabeo striatus, SWU 20130917005, 68.1mm SL, BeipanJiang of Pearl River drainage at An’shun County, Guizhou Province; (b) Sinocrossocheilus labiatus, SWU 20121120001, 60.1 mm SL, Chishui-He River of Yangtze River basin at Bijie City, Guizhou Province; (c) Akrokolioplax bicornis, SWU 20071115020, 102 mm SL, Nujiang River drainage at Baoshan City, Yunnan Province; (d) Crossocheilus burmanicus SWU 201305110001, 79.4 mm SL, Binlang-Jiang of Irrawaddy River drainage at Tengchong County, Yunnan Province; (e) Sinocrossocheilus bamaensis, SWU 20131124001, 103.5mm SL, Hongshui-He of Pearl River drainage at Wangmo County, Guizhou Province; (f) Rectoris posehensis, SWU 20051114006, 107.4mm SL, Yangtze River drainage at Kaixian County, Chongqing.
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FIGURE 4. Lateral view of (a) Paraqianlabeo lineatus, SWU 20130710007, paratype, 65.1mm SL and (b) Qianlabeo striatus, SWU 20130917005, 68.1mm SL, Beipan-Jiang drainage at An’shun County, Guizhou Province.
Color pattern in formalin In formalin-preserved specimens, top of head grayish brown, snout and lip grayish; cheek with sparse brownish pigments; opercle and subopercle brownish. Body gray and yellowish dorsally and laterally, slightly pale ventrally. A longitudinal dark stripe originating near upper extremity of gill opening and terminating medially at caudal-fin base, extends anteriorly along first longitudinal scale row above lateral line and posteriorly along lateral line, slightly wider and more prominent posteriorly, especially at the end of caudal base. All fins yellowish with black speckles on it. Dorsal, caudal, pectoral and pelvic-fins with dark chromatophores on rays, conferring a dusky appearance. Distribution. Known from tributaries of the Wu-Jiang and Chishui-He rivers within the upper Yangtze River basin at Wangcao (28°13' N 107°13' E), Suiyang County; Shuikan (28°4' N 106°46' E), Tongzi County; and Jingmen (27°32' N 105°59' E), Jinsha County, Guizhou Province, South China (Fig. 1).
Habitat and ecology Paraqianlabeo lineatus was collected in clearwater hill streams with sluggish flow over gravel substrate (Fig. 6). The catchment of this tributary is endowed with numerous hill streams with substrates of stone. The width of the river channel was about 5–6 meters and water depth approximately 60 cm. Individuals of Paraqianlabeo lineatus swim slowly and hide among the rocks. Only two other species were collected with them: Carassius auratus (Cyprinidae) and Pseudorasbora parva (Cyprinidae). Etymology. The specific epithet is from the Latin lineatus, meaning lined, referring to the longitudinal dark stripe along the side of the body.
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0.145 0.182
0.186
0.192
0.202
0.166
0.159
0.148
0.160
0.169
0.144
0.143
0.139
0.157
0.164
0.148
0.148
0.158
0.144
0.146
0.144
0.131
0.155
0.146
0.180
0.183
0.159
0.159
0.154
0.129
0.151
0.163
0.133
0.159
0.156
0.175
0.144
0.167
0.151
0.156
0.151
0.147
0.147
0.133
2
0.161 0.170
0.153
1
0.147
0.133
0.149
0.142
0.150
0.155
0.140
0.147
0.136
0.143
0.144
0.135
0.157
0.139
0.128
0.152
0.174
0.141
0.178
0.157
0.138
0.152
3
0.183
0.165
0.176
0.164
0.166
0.152
0.163
0.173
0.161
0.190
0.151
0.163
0.173
0.161
0.152
0.166
0.166
0.150
0.183
0.173
0.151
4
0.133
0.135
0.141
0.126
0.150
0.135
0.132
0.150
0.125
0.152
0.142
0.126
0.157
0.153
0.132
0.144
0.137
0.141
0.174
0.154
5
0.142
0.151
0.162
0.151
0.146
0.144
0.169
0.169
0.141
0.163
0.144
0.144
0.151
0.133
0.151
0.142
0.167
0.159
0.190
6
0.183
0.164
0.178
0.180
0.207
0.183
0.184
0.189
0.184
0.197
0.200
0.183
0.212
0.197
0.187
0.185
0.162
0.184
7
0.135
0.127
0.148
0.146
0.160
0.139
0.137
0.155
0.134
0.150
0.141
0.139
0.162
0.137
0.131
0.135
0.125
8
0.128
0.136
0.123
0.140
0.147
0.145
0.142
0.151
0.111
0.136
0.129
0.120
0.150
0.132
0.123
0.134
9
0.084
0.121
0.088
0.070
0.094
0.076
0.096
0.088
0.144
0.139
0.092
0.078
0.092
0.090
0.072
10
0.074
0.132
0.082
0.082
0.094
0.072
0.094
0.092
0.135
0.104
0.108
0.084
0.096
0.078
11
0.097
0.160
0.115
0.092
0.096
0.107
0.096
0.096
0.142
0.148
0.104
0.092
0.102
12
0.072
0.141
0.126
0.109
0.005
0.098
0.107
0.096
0.155
0.128
0.108
0.106
13
0.070
0.128
0.086
0.084
0.100
0.088
0.087
0.081
0.146
0.121
0.096
14
0.098
0.148
0.110
0.088
0.110
0.108
0.104
0.106
0.138
0.125
15
0.119
0.155
0.121
0.126
0.121
0.114
0.130
0.128
0.152
16
0.149
0.123
0.146
0.153
0.153
0.146
0.141
0.164
17
0.088
0.143
0.090
0.094
0.090
0.083
0.047
18
0.094
0.125
0.096
0.092
0.100
0.091
19
0.074
0.136
0.096
0.104
0.096
20
0.070
0.139
0.120
0.103
21
0.086
0.121
0.082
22
0.088
0.121
23
0.120
24
Note. 1. Labeo yunnanensis, 2. Osteochilus salsburyi, 3. Labiobarbus leptochilus, 4. Catlocarpio siamensis, 5. Lobocheilos melanotaenia, 6. Epalzeorhynchos frenatum, 7. Opsariichthys uncirostris, 8. Garra orientalis, 9. Akrokolioplax bicornis, 10. Sinocrossocheilus bamaensis, 11. Ptychidio jordani, 12. Semilabeo notabilis, 13. Paraqianlabeo, 14. Parasinilabeo assimilis, 15 Bangana lippa, 16. Qianlabeo, 17. Crossocheilus reticulatus, 18. Hongshuia microstomatus, 19. Rectoris posehensis, 20. Stenorynchoacrum xijiangensis, 21. ‘Qianlabeo’ (sensu Zheng et al., 2010) [=Paraqianlabeo lineatus], 22. Discogobio yunnanensis, 23. Sinocrossocheilus labiatus, 24. Cirrhinus molitorella, 25. Pseudogyrinocheilus prochilus.
25
1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24
TABLE 3. K2P distances of the mitochondrial COI sequences between species of labeonines.
FIGURE 5. (a) Left side pharyngeal tooth plate in dorsomedial view, and (b) gas bladder in ventral view of Paraqianlabeo lineatus.
Discussion There are three basic patterns of oromandibular structures among Southeast and East Asian genera of the Labeonini (Zhang & Chen 2004; Zhang et al. 2006; Zhang et al. 2008). Paraqianlabeo belongs to the third group, the subtribe Garraina (= Garrae sensu Rainboth 1996), that have a well-developed or reduced upper lip that is separated from the upper jaw by a deep groove. This group comprises the following genera: Garra, Crossocheilus, Epalzeorhynchos, Parasinilabeo, Sinocrossocheilus, Rectoris, Paracrossochilus, Pseudocrossocheilus, Discogobio, Discocheilus, Placocheilus, Pseudogyrinocheilus, Akrokolioplax, Semilabeo, Stenorynchoacrum and Hongshuia. Paraqianlabeo can be easily distinguished from Hongshuia, Discocheilus, Placocheilus and Discogobio by the presence of three (vs. two) rows of pharyngeal teeth. It also can be separated from Rectoris, Qianlabeo and Stenorynchoacrum by having mental grooves present (vs. absent). Six other garrain genera, Crossocheilus, Pseudocrossocheilus, Sinocrossocheilus, Epalzeorhynchos, Akrokolioplax and Rectoris, have rostral caps with fimbriate (grooved) ventral margins, while that of the new genus is not fimbriate, with no obvious grooves (Fig. 3). Further, the rostral fold and cap of the new genus differs from those of Semilabeo, Pseudogyrinocheilus and Parasinilabeo in being smooth vs. densely covered with papillae (Zhu et al. 2011; Fig.
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2). These three genera have rostral caps with a densely papillated ventral margin and a densely papillated lower lip; these characters all easily distinguish them from Paraqianlabeo. The detailed characters utilized to distinguish Paraqianlabeo from other related genera are given in diagnosis.
FIGURE 6. Type locality of Paraqianlabeo lineatus, Jingmen (Chishui-He) Village, Jinsha County, Guizhou Province, China. Photograph by H.T. Zhao.
Qianlabeo is very similar to Paraqianlabeo in having a longitudinal dark stripe along the side of the body and in body size, but Paraqianlabeo is conspicuously distinct from Qianlabeo in having a suctorial lower lip (vs. absent), a short postlabial groove (vs. long), a mental groove present (vs. absent), and development of papillae on the rostral fold and a complete upper lip (vs. incomplete and restricted to the side of upper jaw). Four sequences were newly determined and deposited in GenBank (Table 1). All the 27 aligned sequences of the COI contain 588 nucleotide sites, of which 215 were variable and 176 potentially parsimony-informative. No indels were observed, and no stop codons were found in protein coding regions, suggesting that nuclear pseudogenes were not sequenced. The genetic distance (K2P) between Paraqianlabeo and other labeonine genera ranges from 6.9% to 19.0% (Table 3); the lowest pairwise genetic distance is between Pseudogyrinocheilus and Paraqianlabeo. The average genetic distance of three geographic populations of Paraqianlabeo is less than 2% and within the range typical for intra-specific variation in other cyprinids (Zhao et al., unpublished data). The NJ tree is shown in Fig. 7. The monophyly of labeonine was recovered, consistent with findings in previous studies (e.g., Zheng et al. 2010; Yang et al. 2012). Among Paraqianlabeo, individuals from the three locations in northern Guizhou Province formed a monophyletic group with strong support (BPP = 100; Fig. 7). A specimen identified as ‘Qianlabeo striatus’ (sequence data from Zheng et al. 2010) that falls within the Paraqianlabeo group in the tree was not originally collected from its type locality Beipan-Jiang of Pearl River drainage, Guizhou Province, but from the Yangtze River drainage in Zunyi City, Guizhou Province. This ‘Qianlabeo striatus’ specimen was most probably misidentified by the authors (Zheng et al. 2010) and should be
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assigned to Paraqianlabeo. The genus Qianlabeo (sensu Zhang & Chen 2004) is situated between the Labeonini (= Labeonina) and the Garrini (= Garraina) based on a Qianlabeo sequenced from its type locality (Beipan-Jiang of the Pearl River drainage in the central region of Guizhou Province), consistent with the position suggested by Zhang & Chen (2004). Taken together, our data suggest that Qianlabeo is not distributed in the Yangtze River drainage, but is restricted to its type locality, the Beipan-Jiang tributary of the Pearl River drainage in the central region of Guizhou Province.
FIGURE 7. Neighbor-joining (NJ) tree of labeonine species and two outgroup species, Opsariichthys uncirostris and Catlocarpio siamensis, based on mitochondrial COI sequences. Node values = bootstrap test (1,000 pseudoreplicates), low supports (< 50%) were not shown.
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Comparative material Qianlabeo striatus: SWU 20130917001–012, 54.7–80.8mm SL, Beipan-Jiang of Pearl River drainage at An’shun County, Guizhou Province; Sinocrossocheilus labiatus: SWU 20121120001–010, 53.2–60.0 mm SL, Chishui-He River of Yangtze River basin at Bijie City, Guizhou Province; Akrokolioplax bicornis: SWU 20071115020, 102 mm SL, Nujiang River drainage at Baoshan City, Yunnan Province; Crossocheilus burmanicus: SWU 201305110001, 79.4 mm SL, Binlang-Jiang of Irrawaddy River drainage at Tengchong County, Yunnan Province; Sinocrossocheilus bamaensis: SWU 20131124001–011, 54.7–123.5 mm SL, Hongshui-He of Pearl River drainage at Wangmo County, Guizhou Province; Rectoris luxiensis: SWU 20051114006, 107.4 mm SL, Jialing-Jiang of upper Yangtze River drainage at Kaixian County, Chongqing; Bangana rendahli: SWU 20130804001, 123.5 mm SL, Wu-Jiang of upper Yangtze River drainage at Wuchuan County, Zunyi City, Guizhou Province; Discogobio yunnanensis: SWU 20130806002–008, 70.3–95.1 mm SL, Wu-Jiang of upper Yangtze River drainage at Tongzi County, Zunyi City, Guizhou Province; Pseudogyrinocheilus prochilus: SWU 20130804003, 134.6 mm SL, Wu-Jiang of upper Yangtze River drainage at Wuchuan County, Zunyi City, Guizhou Province; Sinilabeo hummeli: SWU 20080410001, 161.1 mm SL, Jialing-Jiang of upper Yangtze River drainage at Hechuan County, Chongqing; Ptychidio jordani: SWU 020121201089, 84.8 mm SL; China: Guangxi Province, Hongshui-He of Pearl River drainage at Du’an County.
Acknowledgments We are indebted to Drs. Z. J. Wang (Southwest University) and H. P. Zhao (Henan University) for their help with specimen collection. We are also grateful to L. Y Wang for his help with preparing photographs. This work was supported by grants from the Fundamental Research Funds for the Central Universities of China (XDJK2014c003) and the National Natural Science Foundation of China (31272283).
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