A new species of Rhinogobius (Teleostei: Gobiidae) from Province,

Shih-Pin Huang, I-Shiung Chen & Kwang-Tsao Shao

Ichthyological Research The Ichthyological Society of Japan

ISSN 1341-8998 Volume 63 Number 4

Ichthyol Res (2016) 63:470-479 DOI 10.1007/s10228-016-0516-9

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Ichthyol Res (2016) 63:470–479 DOI 10.1007/s10228-016-0516-9

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A new species of Rhinogobius (Teleostei: Gobiidae) from Zhejiang Province, China

1 2 1 Shih-Pin Huang • I-Shiung Chen • Kwang-Tsao Shao

Received: 14 January 2016 / Revised: 22 February 2016 / Accepted: 23 February 2016 / Published online: 22 March 2016 Ó The Ichthyological Society of Japan 2016

Abstract A new freshwater goby, Rhinogobius niger, Introduction was collected from Zhejiang Province, Southern China. This species can be distinguished from all congeners by a The freshwater goby genus Rhinogobius Gill 1859 is combination of the following features: second dorsal fin widely distributed in East and Southeast Asia, including rays modally I/9, anal fin rays I/8, pectoral fin rays 16–18, Japan, Taiwan, Philippines, Russia, Korea, China, Viet- longitudinal scale series 35–37, transverse scale series nam, Laos, and Thailand (Herre 1927; Wu and Ni 1986; 10–12, predorsal scale series modally 0, vertebral counts Chen and Shao 1996; Chen et al. 1999a; Chen and Kottelat modally 10 ? 17 = 27, head and body blackish in male, 2000, 2003, 2005; Akihito et al. 2013). The life histories of cheek without any stripe or spot, the upper margin of the species of Rhinogobius indicate that the genus includes preoperculum with two horizontally aligned reddish brown both amphidromous and landlocked species (Mizuno 1960; stripes running backwardly, branchiostegal membrane with Mizuno and Goto 1987; Iguchi and Mizuno 1991; Akihito 15–25 small reddish orange round spots in adult male, first et al. 1993; Akihito et al. 2002; Huang and Chen 2007; dorsal fin with a large black blotch anteriorly. A diagnostic Yang et al. 2008). More than 80 species of Rhinogobius are key to all nominal species of Rhinogobius with high ver- known in East and Southeast Asia, and some of these still tebral count (27–29) from mainland southeastern Asia, need formal description (Yang et al. 2008). China, Japan and Taiwan is also provided. In southern and southeastern mainland China, most species of Rhinogobius belong to nondiadromous, land- Keywords Gobiidae Á Rhinogobius Á New species Á locked species (Chen and Miller 1998; Chen et al. 2008). Freshwater goby Á Southern China Most of these have a high vertebral count of 27–29. So far, except for mainland southeastern Asia and China, only two nondiadromous, fluvial species with high vertebral count have been found in the inland waters of Japan and Taiwan S.-P. Huang and I-S. Chen contributed equally to this work. as Rhinogobius flumineus (Mizuno 1960) and Rhinogobius This article was registered in the Official Register of Zoological rubromaculatus Lee and Chang 1996, respectively. Nomenclature (ZooBank) as 5CD974A5-F8A9-4F30-B268- There are 22 valid species of Rhinogobius with high 41BA4DEABBFF. vertebral counts (27–29) found in mainland China, as fol- This article was published as an Online First article on the online publication date shown on this page. The article should be cited by lows: Rhinogobius davidi (Sauvage and Dabry de Thiersant using the doi number. 1874), Rhinogobius cheni (Nichols 1931), Rhinogobius lindbergi Berg 1933, Rhinogobius duospilus (Herre 1935), & Kwang-Tsao Shao Rhinogobius henryi (Herre 1938), Rhinogobius filamento- [email protected] sus (Wu 1939), Rhinogobius szechuanensis (Tchang 1939), 1 Biodiversity Research Center, Academia Sinica, Nankang, Rhinogobius multimaculatus (Wu and Zheng in Zheng and Taipei, Taiwan Wu 1985), Rhinogobius lentiginis (Wu and Zheng in Zheng 2 Institute of Marine Biology, National Taiwan Ocean and Wu 1985), Rhinogobius yaoshanensis (Luo 1989a), University, Keelung, Taiwan Rhinogobius parvus (Luo 1989b), Rhinogobius 123 Author's personal copy

A new species of Rhinogobius from China 471 genanematus Zhong and Tzeng 1998, Rhinogobius xian- 26, was described from the Zhejiang Province. Here, we shuiensis Chen, Wu, and Shao 1999b, Rhinogobius describe a new species of Rhinogobius with high vertebral honghensis Chen, Yang, and Chen 1999c, Rhinogobius count collected from several rivers in the Zhejiang Pro- linshuiensis Chen, Miller, Wu, and Fang 2002, Rhinogob- vince, China. A diagnostic key to all nominal species of ius wangchuangensis Chen, Miller, Wu, and Fang 2002, Rhinogobius with high vertebral count from mainland Rhinogobius wangi Chen and Fang 2006, Rhinogobius southeastern Asia, China, Japan, and Taiwan is also changtinensis Huang and Chen 2007, Rhinogobius lung- provided. woensis Huang and Chen 2007, Rhinogobius ponkouensis Huang and Chen 2007, and Rhinogobius longyanensis Chen, Cheng, and Shao 2008, and Rhinogobius wuyanlin- Materials and methods gensis Yang, Wu, and Chen 2008. Moreover, the following 11 nominal species of Rhinogobius with high vertebral All examined specimens were collected by hand-net. The counts (27–28) can be found in mainland southeastern Asia sampling localities are given in Fig. 1. Specimens were (including Vietnam, Laos, and Thailand): Rhinogobius fixed in 10 % formalin for three days, followed by 70 % chiengmaiensis Fowler 1934, Rhinogobius mekongianus ethanol for long-term preservation. The morphological Pellegrin and Fang 1940, Rhinogobius albimaculatus Chen, measurements follow Miller (1988) and meristic counts Kottelat, and Miller 1999a, Rhinogobius lineatus Chen, follow Chen and Shao (1996). The terminology of cephalic Kottelat, and Miller 1999a, Rhinogobius taenigena Chen, sensory canals and free neuromast organ (sensory papillae) Kottelat, and Miller 1999a, Rhinogobius maculicervix is based on Sanzo (1911) and Huang and Chen (2007). Chen and Kottelat 2000, Rhinogobius milleri Chen and The meristic abbreviations are as follows: A, anal fin; Kottelat 2003, Rhinogobius nammaensis Chen and Kottelat D1, first dorsal fin; D2, second dorsal fin; LR, longitudinal 2003, Rhinogobius vermiculatus Chen and Kottelat 2003, scale series; P, pectoral fin; PreD, predorsal scales; SDP, Rhinogobius boa Chen and Kottelat 2005, and Rhinogobius scale series from origin of the first dorsal fin to the upper sulcatus Chen and Kottelat 2005. pectoral origin; TR, transverse scale series; VC, vertebral The number of vertebrae is frequently used for specific counts. All lengths are standard length (SL). Examined identification in genus Rhinogobius (Chen and Fang 2006; specimens are deposited in the National Taiwan Ocean Chen et al. 2008; Yang et al. 2008). In the Zhejiang Pro- University, Keelung, Taiwan (NTOUP) and Biodiversity vince, there are five nominal endemic species with a high Research Museum, Biodiversity Research Center, Acade- vertebral count (27–29), including R. davidi, R. lentiginis, mia Sinica, Taipei, Taiwan (ASIZP). Comparative speci- R. multimaculatus, R. genanematus, and R. wuyanlingensis mens of other nominal Rhinogobius species from mainland (Yang et al. 2008), which are all landlocked species. southeastern Asia, China, Japan and Taiwan are listed in Recently, an endemic species of Rhinogobius, Rhinogobius Chen and Miller (1998), Huang and Chen (2007), Yang wuyiensis Li and Zhong 2007, with normal vertebral count et al. (2008), and Chen et al. (2008).

Fig. 1 Sampling localities of Rhinogobius niger sp. nov. from Zhejiang Province, China. m, sampling localities; *, type locality

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Rhinogobius niger sp. nov. City, Zhejiang Province, S. P. Huang, 9 January 2008. ASIZP 78399, 5 specimens, 29.3–37.3 mm SL, Caoe (Tables 1–3; Figs. 1, 2, 3). River, Shengzhou City, Zhejiang Province, China, I-S. Holotype. NTOUP 2013-10-107, male, 43.9 mm SL; Chen, 17 October 2002. Yong’an River, a tributary of Ling River, Pan’an County, Diagnosis. Rhinogobius niger is distinguished from all Zhejiang Province, China, I-S. Chen, 17 October 2002. congeners by a combination of the following features: Paratypes. NTOUP 2013-10-108, 8 specimens, second dorsal fin rays modally I/9, anal fin rays I/8, pec- 28.3–42.3 mm SL, all collected with holotype. NTOUP toral fin rays 16–18, longitudinal scale series 35–37, 2013-10-109, 8 specimens, 26.3–38.6 mm SL, Ling River, transverse scale series 10–12, predorsal scale series mod- Xianju County, Zhejiang Province, China, I-S. Chen, 17 ally 0 (rarely 1), vertebral counts 27 (rarely 28); head and October 2002. NTOUP 2013-10-110, 18 specimens, body blackish in male, cheek without any stripe or spot, the 28.7–42.7 mm SL, Caoe River, Shengzhou City, Zhejiang upper margin of preoperculum with two horizontally Province, China, I-S. Chen, 17 October 2002. NTOUP aligned reddish brown stripes running backwardly, bran- 2013-10-111, 2 specimens, 31.5–43.3 mm SL, , chiostegal membrane gray background and with 15–25 Yongjia County, Zhejiang Province, China, I-S. Chen, 16 small reddish orange round spots in adult male, first dorsal October 2002. NTOUP 2013-10-112, 6 specimens, fin with a large black mark. 30.5–35.5 mm SL, Yong River, Xikou Town, Fenghua

Table 1 Morphometric Rhinogobius niger sp. nov. measurements of Rhinogobius niger sp. nov. from Zhejiang Male Female Province, China n = 6 n = 5

Percent standard length (%) Head length 30.6–31.1 (30.8) 28.1–28.6 (28.4) Predorsal length 39.0–41.1 (39.9) 39.4–40.5 (39.9) Snout to 2nd dorsal origin 57.7–60.3 (58.8) 57.3–59.4 (58.4) Snout to anus 58.9–61.1 (60.2) 58.5–60.1 (59.4) Snout to anal fin origin 63.0–65.6 (63.8) 62.7–65.1 (63.8) Prepelvic length 28.5–30.8 (29.7) 28.3–29.4 (28.9) Caudal peduncle length 25.7–28.1 (26.8) 25.1–26.4 (25.6) Caudal peduncle depth 12.4–14.1 (13.0) 11.8–12.6 (12.2) 1st dorsal fin base 16.3–18.2 (17.1) 15.0–16.1 (15.4) 2nd dorsal fin base 17.2–20.6 (19.4) 18.4–20.3 (19.1) Anal fin base 12.4–14.7 (13.7) 13.1–15.3 (14.2) Caudal fin length 24.9–26.9 (26.0) 21.1–23.6 (22.3) Pectoral fin length 24.3–27.8 (25.8) 23.9–24.9 (24.4) Pelvic fin length 13.4–16.6 (14.7) 13.0–14.6 (14.0) Body depth at pelvic fin origin 17.0–18.7 (17.8) 15.8–16.8 (16.4) Body depth at anal fin origin 14.4–16.6 (15.3) 15.6–15.8 (15.7) Body width at anal fin origin 9.9–11.3 (10.8) 9.7–10.6 (10.2) Pelvic fin origin to anus 26.1–29.9 (28.4) 30.5–32.0 (31.2) Percent head length (%) Snout length 30.6–32.1 (31.6) 29.2–30.4 (29.8) Eye diameter 20.4–20.4 (21.0) 21.9–24.6 (23.3) Cheek depth 25.5–27.8 (26.9) 23.8–25.6 (25.0) Postorbital length 45.5–48.0 (46.7) 47.6–51.5 (49.3) Head width in maximum 73.6–76.9 (75.0) 67.0–70.9 (69.4) Head width in upper gill 45.9–46.9 (46.3) 46.5–47.9 (47.0) Bony interorbital width 9.2–11.1 (10.2) 8.5–9.5 (9.1) Fleshy interorbital width 22.2–24.5 (23.5) 20.2–22.1 (21.3) Lower jaw length 47.6–49.2 (48.2) 37.1–39.5 (38.7)

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Table 2 Comparison of frequency distribution of meristic counts for species of Rhinogobius with high vertebral count from Zhejiang Province, China D1 D2 A P V VI Ave I/7 I/8 I/9 I/10 I/11 Ave I/7 I/8 Ave 14 15 16 17 18 Ave

R. niger sp. nov. 1 22 6.0 – 1 22 – – 9.0 – 23 8.0 – – 7 25 14 17.2 R. davidi – 5 6.0 – 1 4 – – 8.8 5 – 7.0 – 2 2 1 – 15.8 R. genanematus – 3 6.0 – 3 – – – 8.0 3 – 7.0 – 6 – – – 15.0 R. lentiginis – 12 6.0 1 10 1 – – 8.0 12 – 7.0 – 6 6 – – 15.5 R. multimaculatus – 15 6.0 – – 4 10 1 9.8 4 11 7.7 1 12 2 – – 15.1 R. wuyanlingensis 1 4 5.8 – 4 1 – – 8.2 – 5 8.0 – – – 1 9 17.9 LR TR 27 28 29 30 31 32 33 34 35 36 37 Ave 8 9 10 11 12 Ave

R. niger sp. nov. ––––––––1281736.3 – – 1 20 2 11.0 R. davidi –––32––––––30.4 – – – 4 1 11.2 R. genanematus 141––––––––28.0 1 2 – – – 8.7 R. lentiginis –––363–––––30.0 – – 12 – – 10.0 R. ––––––2255135.1 – 6 9 – – 9.6 multimaculatus R. –––163–––––31.2 – 1 4 – – 9.8 wuyanlingensis

PreD SDP VC 0 1 2 3 4 5 6 7 8 9 Ave 6 7 8 9 10 11 12 13 14 Ave 27 28 29 Ave

R. niger 22 1 – – – – – – – – 0.0 – – 5 12 6 – – – – 9.0 5 1 – 27.2 sp. nov. R. davidi 5 – – – – – – – – – 0.0 – – 2 2 1 – – – – 8.8 – 5 – 28.0 R. genanematus – – 1 1 – 1 – – – – 3.3 2 1 – – – – – – – 6.3 3 – – 27.0 R. lentiginis 12 – – – – – – – – – 0.0 5 7 – – – – – – – 6.6 12 – – 27.0 R. 15 – – – – – – – – – 0.0 – – – – – 1 4 9 1 12.7 – – 15 29.0 multimaculatus R. – – – – – – – 3 1 1 7.6 – 1 4 – – – – – – 7.8 5 – – 27.0 wuyanlingensis

D1 first dorsal fin; D2 second dorsal fin; A anal fin; P pectoral fin; LR longitudinal scale series; TR transverse scale series; PreD predorsal scales; SDP scale series from origin of first dorsal fin to upper pectoral origin; VC vertebral counts

Description. Morphometric and meristic data are given inserted below second branched soft ray of second dorsal fin. in Tables 1 and 2, respectively. Body rather slender, sub- Pectoral fin broad, the rear tip extending to vertical of anus cylindrical anteriorly and compressed posteriorly. Head when depressed in male, but not extending to vertical of anus moderately large, eye high and large, lip thick. Upper lip when depressed in female, merely reaching mid-ventral more prominent than lower lip. Snout length in male longer region with two scales away from vertical of anus. Pelvic fin than in female. Mouth oblique, corner of mouth extending to disc rounded, spinous ray with pointed membranous lobe. vertical of anterior margin of orbit in male, but just reaching Caudal fin rounded, the rear edge rounded. vertical of head pore r in female. Anterior nostril a short tube Scales. Body covered with moderately large ctenoid scales, and posterior nostril is a round hole. Gill opening, extending predorsal area usually naked, only one individual with one ventrally reaching middle vertical line of operculum. Ver- predorsal scale in all examined specimens. Belly covered with tebral counts 10 ? 17 = 27 (5) -10 ? 18 = 28 (1). cycloid scales. Longitudinal scale series 35–37 (modally 36), Fins. First dorsal fin rays V–VI (modally VI), second transverse scale series 10–12 (modally 11), predorsal scale dorsal fin rays I/8–9 (modally 9), anal fin rays I/8, pectoral fin series 0–1 (modally 0), scale series between first dorsal and rays 16–18 (modally 17). First dorsal fin spines III and IV upper pectoral fin origin 8–10 (modally 9). Opercle, preop- longest, rear tip extending to base of third soft ray of second ercle, prepelvic areas and pectoral fin base always naked. dorsal fin in male when depressed, but only extending to Head canals (Fig. 3). Nasal extension of anterior ocu- anterior margin of second dorsal fin in female. Anal fin origin loscapular canal with terminal pore r at vertical between

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Table 3 Comparison of vertebral count and cephalic pore patterns of 36 species of Rhinogobius with high vertebral count (27–29) from mainland southeastern Asia, China, Japan, and Taiwan Species Number of Cephalic canal pores Sources vertebrae rkjxx1 aqhscde

R. niger sp. nov. 27–28 2212– 2222222Present study R. albimaculatus 28 2212– 2222222Chen et al. (1999a) R. boa 27 2212– 2222222Chen and Kottelat (2005) R. changtinensis 27 2212– 2222222Huang and Chen (2007) R. cheni 27 2212– 2222222Chen et al. (2008) R. chiengmaiensis 28 2212– 2222222Chen et al. (2008) R. davidi 28 2–1–– 2––––––Chen and Miller (1998) R. duospilus 27 2212– 2222222Huang and Chen (2007) R. filamentosus 27 2212– 2222222Chen et al. (2008) R. flumineus 27 2212– 2222222Chen et al. (1999b) R. genanematus 27 2212– 2222222Chen and Fang (2006) R. henryi 28 2212– 2222222Chen et al. (2008) R. honghensis 28 2212– 2222222Chen et al. (1999c) R. lentiginis 27 ––––– –––––––Chen and Miller (1998) R. lindbergi 27 2212– 22222–2Chen et al. (2008) R. lineatus 28 2212– 2222222Chen et al. (1999a) R. linshuiensis 28 2212– 2222222Chen et al. (2002) R. longyanensis 27 2212– 2222222Chen et al. (2008) R. lungwoensis 28 2212– 2222222Huang and Chen (2007) R. maculicervix 28 2212– 2222222Chen and Kottelat (2000) R. mekongianus 28 2212– 2222222Chen et al. (1999a) R. milleri 27 2212– 2222222Chen and Kottelat (2003) R. multimaculatus 29 2–1–– 2––––––Chen and Miller (1998) R. nammaensis 27 2212– 2222222Chen et al. (2008) R. parvus 27 2212– 2222222Chen et al. (2008) R. ponkouensis 28 2212– 2222222Huang and Chen (2007) R. rubromaculatus 27 2–1–– 2––––––Chen and Shao (1996) R. szechuanensis 27 ––––– –––––––Chen et al. (2008) R. sulcatus 27 2212– 2222222Chen and Kottelat (2005) R. taenigena 27 2212– 2222222Chen et al. (1999a) R. vermiculatus 28 2212– 2222222Chen and Kottelat (2003) R. 27 2212– 2222222Chen et al. (2002) wangchuangensis R. wangi 27 2212– 2222222Chen and Fang (2006) R. wuyanlingensis 27 2212– 22222–2Yang et al. (2008) R. xianshuiensis 27 22122 2222222Chen et al. (1999b) R. yaoshanensis 28 2212– 2222222Chen et al. (2008) anterior and posterior nostrils. Anterior interorbital section with pair of pores a. Posterior margin of preoperculum with of oculoscapular canal separated, with pair of pores k. pair of pores c, d, and e. Upper margin of operculum with Single pore j present in posterior interorbital region. Pos- pair of pores q, h, and s. torbital region with pair of pores x. Lateral termination

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Fig. 2 The fresh specimen photographs of Rhinogobius niger sp. nov. from Zhejiang Province, China. a, NTOUP 2013-10-107, holotype, male, 43.9 mm SL; b, NTOUP 2013-10-110, paratype, female, 30.0 mm SL

Sensory papillae (Fig. 3). Row a short, about half of male and grayish white in female. Anal fin membrane orbit diameter. Row b approximately equivalent to orbit usually gray in male and grayish white in female. Caudal diameter, with densely set papillae, starting from vertical of fin membrane always grayish black with no vertical lines in center line of orbit. Row c approximately equivalent to adult male, but female individuals usually with one or two orbit diameter. Single cp papilla. Row d short, about two- brown vertical lines. third of orbit diameter. Opercular papillae with rows ot, oi, Distribution and habitat. This species known from the and os, rows ot and oi well separated. Row f consisting of a upper and middle reaches of Caoe River, Yong River, Ling pair of single papilla. River, and Ou River in Zhejiang Province, southern China Coloration in fresh specimen (Fig. 2). Head and body (Fig. 1). blackish in male and yellowish brown in female; side of Etymology. The Latin specific name, niger, is derived body with seven indistinct grayish black blotches. Most from Latin niger (meaning black), referring to the diag- body scales with black margin. Cheek without any stripe or nostic character of the dusky to blackish head and body in spot. Snout with a pair of reddish brown stripes united at the adult male. snout tip. The upper margin of preoperculum with two Remarks. The present new species, Rhinogobius niger, horizontally aligned reddish brown stripes running back- belongs to a group of 36 nominal species of Rhinogobius wardly. Branchiostegal membrane gray with 15–25 small having 27 or more vertebrae known from mainland reddish orange round spots in adult male; spots usually southeastern Asia, China, Japan, and Taiwan. Compared disappear after long-term preservation. First dorsal fin with to all nominal species of Rhinogobius having 27 or more large black blotch anteriorly in both sexes. Second dorsal vertebrae, Rhinogobius niger can be easily distinguished fin membrane dark to translucent and with 2–3 rows of from R. lentiginis, R. lindbergi, R. multimaculatus, R. horizontally aligned reddish brown spots in both sexes. rubromaculatus, R. szechuanensis, R. wuyanlingensis and Pectoral fin base with one round mark; pectoral fin mem- R. xianshuiensis by having different head pore patterns brane always grayish black in adult male and grayish white (Table 3). Rhinogobius niger can be easily distinguished in female. Pelvic fin membrane always grayish black in from R. albimaculatus, R. boa, R. changtinensis, R. cheni,

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color patterns (pectoral fin base with one round mark vs. pectoral fin base with a vertical dark brown line for R. parvus, and two crescent-shaped dark marks for R. flumineus). Rhinogobius niger is most similar to Rhinogobius davidi, also known from Zhejiang Province, China, in having same second dorsal fin ray count (modally I/9) and usually lacking predorsal scale. However, Rhinogobius niger can be easily distinguished from R. davidi by the following features: anal fin ray I/8 (vs. I/7), longitudinal scale series 35–37 (vs. 30–31), opercular canal pores c, d, e, q, h, and s present (vs. opercular canal pores c, d, e, q, h, and s absent), and no stripe present on the cheek (vs. a reddish brown stripe turning vertically from the eye to the corner of upper lip in R. davidi). Rhinogobius davidi is a rare species and endemic to China (Wu and Zhong 2008). So far, this species is known only from the western Zhe- jiang Province. Except for type specimens, only a few individuals of R. davidi have also been recorded from the , Zhejiang Province, during our freshwater fish fauna expedition in 2002. Some Chinese landlocked Rhinogobius species were reported as known to be only distributed in limited habitats (Huang and Chen 2007). In recent years, seven Chinese endemic Rhinogobius species were listed in China species red list as endangered species due to habitat destruction, overfishing and water pollution (Wang and Xie 2009). An overall investigation for pro- viding detailed conservation information for this genus may be necessary.

A diagnostic key to nominal species of Rhinogobius with high vertebral count (27–29) from mainland southeastern Asia (Thailand to Vietnam), China, Japan, and Taiwan

Fig. 3 The cephalic lateral-line system of Rhinogobius niger sp. 1a. Head canals and pores absent………………………2 nov., NTOUP 2013-10-107, holotype, 43.9 mm SL. a, b, c, cp, d, e, i, 1b. Head at least with an anterior oculoscapular canal…3 f, ot, oi, and os denote sensory papillae rows. r, k, j, x, a, q, h, s, c, d, and e denote cephalic canal pores. The arrow denotes ventral edge of 2a. Second dorsal fin rays I/8, anal fin rays I/7, lateral gill slit. Bar = 1mm side with 6–7 dark brown blotches, pectoral fin base with a horizontal dark brown mark…R. lentiginis (Zhejiang Province, China) R. chiengmaiensis, R. duospilus, R. filamentosus, R. 2b. Second dorsal fin rays I/9, anal fin rays I/8, lateral genanematus, R. henryi, R. honghensis, R. lineatus, R. body as dark brown net-like pattern, pectoral fin base linshuiensis, R. longyanensis, R. lungwoensis, R. mac- with a vertical blackish brown bar…R. szechuanen- ulicervix, R. mekongianus, R. milleri, R. nammaensis, R. sis (Szechuan Province, China) ponkouensis, R. sulcatus, R. taenigena, R. vermiculatus, 3a. Preopercular canal and posterior oculoscapular canal R. wangchuangensis, R. wangi,andR. yaoshanensis by absent………………………………………………4 having different cheek color patterns in male (cheek 3b. Preopercular canal and posterior oculoscapular without stripe or spot vs. cheek always with stripe or canal present………………………………………6 spot). Rhinogobius niger can be distinguished from R. 4a. Predorsal midline naked……………………………5 parvus and R. flumineus (including blotch dorsal form and 4b. Predorsal scales 9–12…R. rubromaculatus (Taiwan) pale dorsal form) by having different pectoral fin base

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5a. Longitudinal scale rows 35–36, second dorsal fin 17a. Longitudinal scale rows 29–31, pectoral fin rays rays I, 10, cheek with many tiny black spots…R. modally 15, 60–75 tiny round spots on cheek and multimaculatus (Zhejiang Province, China) opercle in male…R. albimaculatus (Laos) 5b. Longitudinal scale rows 30–31, second dorsal fin 17b. Longitudinal scale rows 32–34, pectoral fin rays rays I, 9, cheek spotless…R. davidi (Zhejiang modally 16, 7–10 large round spots on opercle in Province, China) male…R. vermiculatus (Laos) 6a. Postorbital region usually with paired pores x and 18a. Cheek with very conspicuous median longitudinal x1, opercle with red reticulation…R. xianshuiensis dark stripe in male…………………………………19 (Fujian Province, China) 18b. Cheek without median longitudinal dark stripe in 6b. Postorbital region merely with paired pore x,pores male………………………………………………22 x1absent……………………………………………7 19a. Lower half of lateral cheek with 8–10 brownish red 7a. Preopercular canal merely with two terminal pores…8 spots in male…R. linshuiensis (Hainan Province, 7b. Preopercular canal always with three terminal pores…9 China) 8a. Predorsal scale series 0…R. lindbergi ( River 19b. Cheek has less than five round dark spots in male…20 basin, China) 20a. Pectoral fin base with two conspicuous dark brown 8b. Predorsal scale series 7–9…R. wuyanlingensis (Zhe- to black spots……………………………………...21 jiang Province, China) 20b. Pectoral fin base with only one black spot…R. 9a. Scales between first dorsal and upper pectoral fin taenigena (Laos) origins 9–11, caudal fin with basal large vertical ‘‘\’’- 21a. Predorsal scales 11–12, cheek with two black spots shaped brownish black mark at least in female which on lower margin in male…R. boa (Vietnam) occupied more than 2/3 depth of caudal fin base…10 21b. Predorsal scales 8–9, cheek with four black spots on 9b. Scales between first dorsal and upper pectoral fin lower margin and some irregularly short brown bars origins 5–8, caudal fin without such large-size on ventral region in male…R. milleri (Laos) vertical dark mark in both sexes…11 22a. Cheek and opercle with over 10 round conspicuous 10a. First dorsal fin with 4th ray longest and large, square red to black round spots in male…………………23 bluish-black mark on anterior 2/3 of first dorsal fin in 22b. Cheek and opercle without such numerous dark marks male…R. flumineus (blotch dorsal form) (Honshu in male……………………………………………27 and Kyushu, Japan) 23a. Cheek and opercle with 11–14 brownish black spots 10b. First dorsal fin with 2nd and 3rd rays longest and in male, lateral side with 6–7 dark brown longitu- pale lacks this large dark mark in male…R. dinal lines…R. lineatus (Laos) flumineus (pale dorsal form) (Honshu, Japan) 23b. Cheek and opercle with over 30 rounded dark spots 11a. Predorsal scales 14–15, prepelvic region with small in male…………………………………………….24 cycloid scales…R. yaoshanensis ( Province, 24a. Pectoral fin rays modally 17, predorsal scales 0–3, China) almost round dark spots on cheek as large as pupil 11b. Predorsal scales 0–11, prepelvic region naked…12 size in male…R. honghensis (Yunnan Province, 12a. Rear tip of first dorsal fin extending to the insertion China) of 6–7 branched rays of second dorsal fin when 24b. Pectoral fin rays modally 16, predorsal scales 5–7, depressed in male…R. filamentosus (Guangxi and all or some round spots on cheek smaller than pupil Guangdong Provinces, China) in male…………………………………………….25 12b. Rear tip of first dorsal fin not extending to vertical 25a. Branchiostegal membrane with large spots about 2/3 midline of second dorsal fin base when depressed in pupil size, cheek and opercle with about 40 round male………………………………………………13 dark spots in male…R. mekongianus (Thailand and 13a. Nape with one (sometimes split into two) conspic- Laos) uous brownish black blotch…R. maculicervix (Laos) 25b. Branchiostegal membrane with tiny yellow to orange 13b. Nape without such a mark…………………………14 spots less than 1/2 pupil size, cheek and opercle with 14a. Longitudinal scale rows 34–36…R. cheni (Jiangxi 55–80 round dark spots in male………………….26 Province, China) 26a. Longitudinal scale rows modally 32, predorsal scales 14b. Longitudinal scale rows less than 33………………15 6–7, nape without dark cross-bars…R. nammaensis 15a. Pectoral fin base with a vertical dark brown line… R. (Laos) parvus (Guangxi Province, China) 26b. Longitudinal scale rows modally 30, predorsal scales 15b. Pectoral fin base lacks such mark…………………16 3–4, nape with a pair of short deep brown cross- 16a. Cheek or opercle with white round spots in male…17 bars…R. chiengmaiensis (Thailand) 16b. Cheek and opercle without such mark in male…18 123 Author's personal copy

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27a. Cheek without stripe or spot in male…R. niger sp. References nov. (Zhejiang Province, China) 27b. Cheek with stripe or spot in male…………………28 Akihito, Iwata A, Sakamoto K, Ikeda Y (1993) Suborder Gobioidei. 28a. Cheek with one or more oblique dark line in male…29 In: Nakabo T (ed) Fishes of Japan with pictorial key in the species. Tokai University Press, Tokyo, pp 997–1392 28b. Cheek with two ventral dark spots in male…R. Akihito, Sakamoto K, Ikeda Y, Sugiyama K (2002) Suborder wangchuangensis (Hainan Province, China) Gobioidei. In: Nakabo T (ed) Fishes of Japan with pictorial 29a. Cheek with three stripes from ventral side backward key in the species. Second edition. Tokai University Press, and upward to upper area in male… R. genanematus Tokyo, pp 1139–1310 Akihito, Sakamoto K, Ikeda Y, Aizawa M (2013) Gobioidei. In: (Zhejiang Province, China) Nakabo, T (ed) Fishes of Japan with pictorial keys to the species, 29b. Cheek with stripe(s) from dorsal side backward and 3rd edition. Tokai University Press, Hadano, pp 1347–1608, downward to lower region in male………………30 2109–2211 30a. Cheek with 6–7 oblique dark brown stripes in Berg LS (1933) Les poissons des eaux douces de l’U.R.S.S et des pays limitrophes. 3-e e´dition, revue et augmente´e. Leningrad male…R. wangi (Guangdong Province, China) Poiss Eaux Douces URSS Part 2:545–903 30b. Cheek with 1–3 oblique, parallel dark stripes in Chen IS, Fang LS (2006) A new species of Rhinogobius (Teleostei: male………………………………………………31 Gobiidae) from the Hanjiang Basin in Guangdong Province, 31a. A horizontal brownish black bar or spot on pectoral fin China. Ichthyol Res 53:247–253 Chen IS, Kottelat M (2000) Rhinogobius maculicervix, a new species base………………………………………………...32 of goby from the basin in northern Laos. Ichthyol 31b. A pair of black spots on pectoral fin base………33 Explor Freshw 11:81–87 32a. Pectoral fin rays modally 16, predorsal scales 0–2, Chen IS, Kottelat M (2003) Three new freshwater gobies of the genus, infraorbital mark as a horizontal brown bar below Rhinogobius (Teleostei: Gobiidae) from northeastern Laos. Raffles Bull Zool 51:87–95 orbit turning into a vertical brown line…R. changti- Chen IS, Kottelat M (2005) Four new freshwater gobies of the genus nensis (Fujian Province, China) Rhinogobius (Teleostei: Gobiidae) from northern Vietnam. J Nat 32b. Pectoral fin rays modally 17, predorsal scales 7–8, Hist 39:1407–1429 infraorbital mark as a small, square dark brown Chen IS, Miller PJ (1998) Redescription of Gobius davidi (Teleostei: Gobiidae) and comparison with Rhinogobius lentiginis. Cybium mark…R. longyanensis (Fujian Province, China) 22:211–221 33a. Predorsal midline naked…R. henryi (Guangdong Chen IS, Shao KT (1996) A taxonomic review of the gobiid fish Province, China) genus Rhinogobius Gill, 1859, from Taiwan with descriptions of 33b. Predorsal scales 4–12………………………………34 three new species. Zool Stud 35:200–214 Chen IS, Cheng YH, Shao KT (2008) A new species of Rhinogobius 34a. Branchiostegal membrane with several parallel (Teleostei: Gobiidae) from the Julongjiang Basin in Fujian orange-red stripes in male…R. duospilus (Hong Province, China. Ichthyol Res 55:335–343 Kong and Guangdong Province, China) Chen IS, Miller PJ, Wu HL, Fang LS (2002) Taxonomy and 34b. Branchiostegal membrane with round orange or red mitochondrial sequence evolution in non-diadromous species of Rhinogobius (Teleostei: Gobiidae) of Hainan Island, southern spots in male………………………………………35 China. Mar Freshw Res 53:259–273 35a. Predorsal scales 8–11, opercle with oblique dark Chen IS, Kottelat M, Miller PJ (1999a) Freshwater gobies the genus stripes in male, vertebral count 27…R. sulcatus Rhinogobius from the Mekong basin in Thailand and Laos, with (Vietnam) descriptions of three new species. Zool Stud 38:19–32 Chen IS, Wu HL, Shao KT (1999b) A new species of Rhinogobius 35b. Predorsal scales 4–6, opercle without such mark, (Teleostei: Gobiidae) from Fujian Province, China. Ichthyol Res vertebral count 28…………………………………36 46:171–178 36a. Cheek with modally two oblique dark stripes in Chen IS, Yang JX, Chen YR (1999c) A new species of Rhinogobius male, branchiostegal membrane with 15–18 small (Teleostei: Gobiidae) from the Honghe basin, Yunnan Province. Acta Zool Taiwan 10:45–52 orange-red spots…R. ponkouensis (Fujian Province, Fowler HW (1934) Zoological results of third de Schauensee Siamese China) Expedition. Part I. Fishes. Proc Acad Nat Sci Philadelphia 36b. Cheek with modally one oblique dark stripe in male, 86:67–163 branchiostegal membrane with 20–28 orange-red Gill TN (1859) Notes on a collection of Japanese fishes by Dr. J. Morrow. Proc Acad Nat Sci Philadelphia 11:144–159 spots or bars…R. lungwoensis (Guangdong Pro- Herre AWCT (1927) Gobies of Philippines and China Sea. Monogr vince, China) Bur Sci Manila 23:1–352 Herre AWCT (1935) Note on the fishes in the Zoological Museum of Stanford University, new and rare Hongkong fishes obtained in Acknowledgments We are very grateful to Mr. Y. C. Chiu for 1934. Nat Hist Soc 6:285–293 assisting in take the X-ray photographs.

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Herre AWCT (1938) Notes on a small collection of fishes from Pellegrin J, Fang PW (1940) Poissons du Laos recueillis par MM. Kwangtung Province including Hainan, China. Lingnan Sci J Delacour, Greenway, Ed. Blanc. Description d’un genre, de cinq Canton 17:425–437 espe´ces et d’une varie´te´. Bull Soc Zool Fr 65:111–213 Huang SP, Chen IS (2007) Three new species of Rhinogobius Gill, Sanzo L (1911) Distribuzione delle papille cutanee (organi ciati- 1859 (Teleostei: Gobiidae) from the Hanjiang basin, southern forme) e suo valore sistematico nei gobi. Mitt Zool Stn Neapel China. Raffles Bull Zool 14:101–110 20:249–328 Iguchi K, Mizuno N (1991) Mechanism of embryonic drift in the Sauvage HE, Dabry de Thiersant P (1874) Notes sur les poissons des amphidromous goby, Rhinogobius brunneus. Environ Biol eaux douces de la Chine. Ann Sci Nat Paris (Zool) (Se´r 6) 1:1–18 Fishes 31:295–300 Tchang TL (1939) Studies on Chinese Glossogobius. Bull Fan Lee SC, Chang LT (1996) A new goby, Rhinogobius rubromaculatus Memorial Inst Biol Zool Ser 9:67–70 (Teleostei: Gobiidae) from Taiwan. Zool Stud 35:30–35 Wang S, Xie Y (2009) China species red list. Vol. II. Vertebrates Li F, Zhong JS (2007) A new Rhinogobius species from Zhejiang (Part 1). Higher Education Press, Beijing Province, China (Teleostei: Gobiidae). Zool Res 28:539–544 Wu HL, Ni Y (1986) Gobiidae. In: The freshwater and estuarine Luo YL (1989a) Ctenogobius yaoshanensis. In: Zheng CY (ed) Zu fishes of Hainan Island. Guangdong Sci Tech Press, , Jiang Yu Rei Chi (Fishes of the Zujiang River). Ke Shuei Chu pp 259–314 Ban She (Science Press), Beijing, pp 352–353, 412–413 Wu HL, Zhong JS (2008) Fauna Sinica. Osteichthys. Perciformes Luo YL (1989b) Ctenogobius parvus. In: Zheng CY (ed) Zu Jiang Yu (V) Gobioidei. Science Press, Beijing Rei Chi (Fishes of the Zujiang River). Ke Shuei Chu Ban She Wu HW (1939) On the fishes of Li-Kiang. Sinensia 10:92–142 (Science Press), Beijing, pp 354–355, 413 Yang JQ, Wu HL, Chen IS (2008) A new species of Rhinogobius Miller PJ (1988) New species of Corcyrogobius, Thorogobius, and (Teleostei: Gobiidae) from the Feiyunjiang Basin in Zhejiang Wheelerigobius from west Africa (Teleostei: Gobiidae). J Nat Province, China. Ichthyol Res 55:379–385 Hist 22:1245–1262 Zheng ML, Wu HL (1985) A study of freshwater gobiid fishes of Mizuno N (1960) Description of a new freshwater goby from Japan. Zhejiang Province, China, with descriptions of two new species. Mem Coll Sci Univ Kyoto Ser B 27:117–119 Acta Zootax Sini 10:328–333 Mizuno N, Goto A (1987) Nihon no tansuigyorui, sono bunpu, heni, Zhong JS, Tzeng CS (1998) A new species of Rhinogobius from shubunka wo megutte (Japanese freshwater fishes, and their China (Perciformes: Gobiidae). Zool Res 19:237–241 distribution, variation and speciation). Tokai University Press, Tokyo Nichols JT (1931) Some Chinese fresh-water fishes. XXIX. A new goby from Hokou, Kiangshi. Am Mus Navit 499:1–3

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