NING Ping, SHA Zhongli, Paul D. N. HEBERT, and Barry RUSSELL



The Taxonomic Status of Japanese Threadfin Bream(Bloch, 1791) (Perciformes: Nemipteridae) with a Redescription of this Species from the South China Sea Based on Morphology and DNA Barcodes

NING Ping1), *, SHA Zhongli1), Paul D. N. HEBERT2), and Barry RUSSELL3)

1),,,266071,..2),,ON N1G 2WI,3),NT 0801,

Because of its importance as a food source,(Bloch, 1791) (Nemipteridae) or Japanese threadfin bream is the best studied of these taxa, and numerous investigations have examined its fisheries, its biology and biochemistry. Despite such intensive work, the taxonomic status of.has never been seriously questioned and itis regarded as a common species, widely distributed throughout the Indo-Western Pacific Ocean. In fact, Bloch’s description of the type specimen of.has ambiguous collection data and lacks a designation for the type locality, though it is probably Java. In this paper, DNA barcode results based on COI gene support the existence of two geographically separated lineages of the Japanese threadfin bream, both being an Indian Ocean and western Pacific lineage, with 2.7% sequence divergence, and the results indicate a possible existing of some cryptic species. The two lineages also possess a diagnostic difference in their belly color, with specimens in the South China Sea having a silver belly, while those from the Indian Ocean isolate specimen have a yellow coloration. Based upon new collections from the South China Sea, this species from the western Pacific is morphologically redescribed and its details of DNA barcode diversity are shown for the future investigations.

South China Sea; Indian Sea; Indo-Pacific Barrier; COI gene

1 Introduction

Swainson, 1839, the largest genus in the family Nemipteridae, includes about 26 species (Russell, 1990) which are distributed in the Indo-Western Pacific Ocean. Because of its importance as a food source,(Bloch, 1791) is the best studied of these taxa and numerous investigations have examined its fisheries, its biology and biochemistry (Krishnamoorthi, 1971; Russell, 1990; Chawla., 1996; Joshi, 2010; Naqasha and Nazeera, 2010). Despite this intensive work, the taxonomic status of.has never been seriously questioned and itis regarded as a common species, widely distributed throughout the Indo-Western Pacific Ocean (Froese and Pauly, 2013).

While carrying out morphological and DNA barcode studies, we discovered evidence for genetic divergence within.that is linked to collection locality. This paper provides a morphological redescription and details DNA barcode diversity in.on the basis of specimens from the South China Sea.

2 Materials and Methods

2.1 Sample Collection and Examination

There are 15 examined specimens collected from the southern coastal waters of China from 1950 to 2011 and deposited at the Marine Biological Museum (MBM), Institute of Oceanology, Chinese Academy of Sciences (IO- CAS). The other 22 specimens examined are in: Academy of Natural Sciences (ANSP) Philadelphia, USA; Academica Sinica (ASIZ), Taipei, Taiwan, China; the Natural History Museum (BMNH), London, UK; California Aca- demy of Sciences (CAS), San Francisco, USA; Museum and Art Gallery of the Northern Territory (NTM), Darwin, Australia; Senckenberg Museum (SMF), Frankfurt, Germany; Smithsonian Institution (USNM), Washington DC, USA; Zoological Museum (ZMB), Berlin, Germany.

2.2 Morphological Description

Color and pigmentation patterns were examined in digital images of freshly collected specimens, and de- tailed measurements were made on specimens preserved in 95% alcohol or 10% formalin. The technical terms and measurements followed Russell (1990): SL means stan- dard length. HL means head length. Distributional data derived from Russell (1990), FishBase (Froese and Pauly, 2013) and the Taiwan Fish Database (Shao, 2013).

2.3 DNA Extraction, PCR, and Sequencing

Genomic DNA was extracted from three individuals (IOCAS 135FB00144-1, IOCAS 135FB00144-2, IOCAS 135FB00144-3) using Tiangen® Marine Animal Tissue DNA Extraction Kit and following the manufacturer’s instructions. Polymerase chain amplification was conducted using standard fish primers (Ward., 2005) for the barcode region of the COI gene:

COIF15’-TCAACCAACCACAAAGACATTGGCAC-3’,

COIR15’-TAGACTTCTGGGTGGCCAAAGAATCA-3’.

Each reaction had a total volume of 25μL containing 1μL of template, 2.5μL of 10×PCR buffer, 1.5μL of Mg2+, 1 μL of dNTP mix, 1μL of each primer, 0.3 μL of Taq DNA polymerase, and 17.7μL of dH2O. The thermocycler regime was 3min at 95℃, followed by 35cycles, each at 94℃ for 30s, at 55℃ for 30s, at 72℃ for 1min, and final extension at 72℃ for 10min. The reaction products were separated by electrophoresis on 1.0% agarose gel, stained with ethidium bromide, and visualized under ultraviolet light. Sequencing reactions were performed at the Key Laboratory of Experimental Marine Biology (KLEMB) of IOCAS. Three COI sequences had been deposited into the NCBI database with the access numbers from KF134003 to KF134005, corresponding to voucher ID from 135FB00144-1 to 135FB00144-3.

2.4 COI Sequences Downloaded from GenBank and Checked by BOLD

COI sequences for additional specimens were obtained from GenBank:.from the Indian Ocean (GenBank accession numbers: EF609553-EF609555) (Lakra., 2011);.(GenBank accession numbers: EF609557-EF609559);.(GenBank accession numbers: FJ237833-FJ237835);.(GenBank accession number: EF609414);.(GenBank accession number: EF609415);.(GenBank accession number: EF609413). A sequence from(GenBank accession number: HQ945943) was used as an outgroup. The COI sequences from GenBank and those obtained in this study were validated on BOLD (Ratnasingham and Hebert, 2007) to ensure that specimens were properly identified.

2.5 Sequence Alignment and Phylogenetic Analysis

All sequences were aligned using the CLUSTAL W, and then truncated to a uniform length by MEGA 5.0. The aligned sequences were subjected to both maximum- likelihood (ML), and neighbour-joining (NJ) analyses using MEGA 5.0 (Tamura., 2011). The model of ‘HKY+G+I’ for ML was chosen based on results from the option of ‘Find Best DNA Models’: heuristic method is Nearest-Neighbor-Interchange (NNI), bootstrap values are computed with 1000 replications. NJ bootstrap values were estimated using 1000 replicates with Kimura’s two-parameter model of substitution (K2P distance) evolution model. The genetic distances were estimated based on Kimura’s two-parameter model using MEGA 5.0 software.

3 Results

3.1 Synonym

(Bloch, 1791)

Fig.1 Nemipterus japonicus (IOCAS 135FB00144-1), 151.5mm SL, Sanya City, Hainan Island, China. Scale bar=1.0 cm.

-Bloch, 1791: 110 (original type locality: lacks a designation for the type locality; the type specimen has ambiguous collection data, but probably Java)

?-Schneider in Bloch and Schneider, 1801: 297, fig. 58 (Tranquebar, India)

– Rüppell, 1828–30 (1829): 50, pl. 12, fig. 3 (Massaua, Red Sea)

-Valenciennes in Cuvier and Valen- ciennes, 1830: 249, 558 ('Rade de Pondicherry', India)

-Valenciennes in Cuvier and Valenciennes, 1830: 250 ( Pondicherry, India)

?-Valenciennes in Cuvier and Valen- ciennes, 1830: 252 (Tranquebar, India)

-(non Valenciennes)Cantor, 1849: 1032 (Penang)

-Bleeker, 1851: 176 (Batavia =Jakarta)

-Günther, 1859: 378 (Batavia =Jakarta)

-Günther, 1859: 378 (Red Sea; coast of Pondicherry, India)

-Bleeker, 1865: 173 (Siam)

-Day, 1865: 14 (Cochin, Malabar coast of India)

-Steindachner, 1866: 778, p1.13, fig. 6 (Zanzibar)

()-Steindachner, 1868: 976 (Mauritius)

-Jordan and Seale, 1907: 21 (Cavite, Philippines); Akazaki, 1962: 91 (Tonkin Bay, Vietnam; Iran); Cheng, 1962: 505–510 (South China Sea); Wu, 1985: 187–193 (East China Sea); Lee, 1986: 168, pl. 3, fig. 13 (Kaohsiung, Taiwan); Russell, 1986: 19–35 (species catalog, Indian Sea); Li, 1987: 336–337 (China sea); Russell, 1990: 40, pl. II, b, fig. 72(Indian Ocean and West Pacific);Lee, 1993: 369–371 (Taiwan); Russell, 1993: 295–310 (eastern Pacific)

-Fowler, 1931: 299, fig. 8 (Saukiwan, Hong Kong, China)

-Fowler, 1933: 94 (Philippines; Hong Kong, China)

-Herre, 1934: 55 (Manilla, Philippines)

3.2 Specimens Examined

Holotype – 1 ex. (146.0mm SL) (ZMB 8147), lacks a designation for the type locality.

Material examined from the South China Sea – 1 ex. (191.5mm SL) (IOCAS 55-1758), Changhua [=Town], Hainan Island, China; 1 ex. (228.5mm SL) (IOCAS 56-9188), Beibu Gulf (Gulf of Tonkin); 1 ex. (171.5mm SL) (IOCAS 76-018), Sanya [=City], Hainan Island, China; 1 ex. (147.5mm SL) (IOCAS 76-333), Baimajing [=Town], Hainan Island, China; 1 ex. (191.5mm SL) (IO- CAS 83-370), Guangzhou [=City], Guangdong Province, China; 1 ex. (142.5mm SL) (IOCAS JXY-10-0038), Beihai [=City], Guangxi Zhuang Autonomous Region [=Province], China; 1 ex. (144.5mm SL) (IOCAS JXY- 10-0039), Beihai, Guangxi Zhuang Autonomous Region, China; 1 ex. (140.5mm SL) (IOCAS JXY-10-0040), Beihai, Guangxi Zhuang Autonomous Region, China; 1 ex. (111.5mm SL) (IOCAS 20101023001), Zhanjiang [=City], Guangdong Province, China; 1 ex. (126.5mm SL) (IOCAS 201010221614), Zhanjiang [=City], Guangdong Province, China; 1 ex. (97.5mm SL) (IOCAS 20101023 1005), Beihai, Guangxi Zhuang Autonomous Region, China; 1 ex. (151.5mm) (IOCAS 135FB00144-1), Sanya, Hainan Island, China; 1 ex. (159.5mm) (IO-CAS 135FB 00144-2), Sanya, Hainan Island, China; 1 ex. (169.5mm) (IOCAS 135FB00144-3), Sanya, Hainan Island, China; 1 ex. (142.8mm) (135FB00144-4), Sanya, Hainan Island, China.

Other materials examined – 1 ex. (171.0mm SL) (NTM S. 11003-001), Sunda Strait, Indonesia; 2 ex. (89.5–90.8 mm SL) (BMNH 1880.4.21.42-43), Batavia (=Jakarta), Java, Indonesia; 2 ex. (121.8–127.0 mm SL) (CAS SU 61463), Surubaya, Indonesia; 1 ex. (161.0 mm SL) (BMNH 1984.8.20.10), SE-Java, Indonesia; 1 ex. (197.0 mm SL) (NTM S. 10735-005), Bali Strait, Indonesia; 1 ex. (150.7 mm SL) (NTM S. 10678-006), Kuching, Sarawak, Malaysia; 2 ex. (153.0–181.7 mm SL) (NTM S. 10677-014), Kota Kinabulu, Sabah, Malaysia; 1 ex. (195.0 mm SL) (SMF 19975), Bangkok, Thailand; 1 ex. (109.7 mm SL) (USNM 112811), Iloilo, Philippines; 1 ex. (142.6 mm SL) (USNM 122158), Corregidor I., Philippines; 1 ex. (170.0 mm SL) (USNM 122148), San Fernando, Philippines; 2 ex. (102.0–132.0 mm SL) (CAS SU 6100), Tolo Channel, Hong Kong, China; 3 ex. (109.0–174.0 mm) (NTM S. 10676-002), Aberdeen market, Hong Kong, China; 1 ex. (107.0 mm SL, holotype of) (ANSP 53454), Saukiwan, Hong Kong, China; 1 ex. (110.0mm SL, paratype of) (ANSP 53455), Saukiwan, Hong Kong, China; 1 ex. (156.0mm SL) (ASIZ P055432), Kao-hsiung, Taiwan, China .

3.3 Diagnosis

Pectoral fins very long, reaching to or just beyond level of origin of anal fin; pelvic fins moderately long, reaching to or just beyond anus; upper lobe of the caudal fin is produced into a short or moderately long filament (this character is not obvious in small specimens). Upper part of body pinkish, becoming silvery below; belly silvery from lower jaw to base of pelvic fin, rest yellow; top of head behind eye with a golden sheen; 11 to 12 pale golden-yellow stripes along body from behind head to base of caudal fin; a red blotch below origin of lateral line; a pale lemon stripe near base of dorsal fin, this stripe narrows anteriorly and widens on posterior part of fin; anal fin whitish with pale lemon broken lines or scribbles over most of fin; caudal fin pink, upper tip and filament yellow.

3.4 Description

Dorsal fin X, 9; anal fin III, 7; pectoral fin 17; pelvic fin i, 5; caudal fin 17; lateral-line scales 43–47; transverse scale rows 3/10; gill rakers 14–17.

Body depth 2.7–3.4 in SL, head length 3.1–3.5 in SL, caudal peduncle length 6.1–7.3 in SL, pectoral length 2.8–3.3 in SL, pelvic length 3.3–4.1 in SL, dorsal fin base length 1.8–2.0 in SL, anal fin base length 4.9–5.8 in SL, predorsal length 2.5–2.8 in SL, length from the origin of pelvic fin to the origin of the anal fin 3.0–3.6 in SL.

Snout length 3.1–3.9 in HL, eye diameter 3.1–4.0 in HL, length of upper jaw 2.7–3.2 in HL, depth of subor- bital bone 3.7–5.7 in HL.

3.5 Coloration

Upper part of body pinkish, lower flanks silvery; belly silvery from lower jaw to base of pelvic fin, rest yellow; top of head behind eye with a golden sheen; 11 to 12 pale golden-yellow stripes along body from behind head to base of caudal fin; an obvious red blotch below origin of lateral line; margin of dorsal fin yellow, edged with red; a yellow stripe near base of dorsal fin, this stripe narrow anteriorly and widening on posterior part of fin; anal fin whitish with yellow broken lines or scribbles over most of fin; pectoral fin translucent pinkish; pelvic fins whitish with yellow axillary scale; caudal fin pink, upper tip and filament yellow.

3.6 Distribution

Widespread throughout the Indian Ocean and the West Pacific ranging from East Africa, including the Persian (Arabian) Gulf and Red Sea, to the Indo-Malay Archi- pelago. In China, occurs in the Beibu Gulf (Gulf of Tokin), South China Sea (northern part) and the Taiwan Strait.

3.7 Sequence Characters and Gene Tree for

Clustal W was used to align the three new 655bp sequences for.. Once aligned with the data from GenBank, the records produced a 652bp alignment. Of these sites 476 were conserved, 176 were variable, 149 were parsimony informative and 27 were singletons. Nucleotide frequencies were A=23.4%, T=31.8%, G= 18.1% and C=26.7%. As expected, transitions (si=54) were more frequent than transversions (sv=19) with an average ratio (si/sv) of 2.79 (ingroup).

The gene trees constructed using the two methods showed a similar topology. The species ofwere grouped into three major clades by both NJ (Fig.2) and ML (Fig.3) with a bootstrap value higher than 50%: (1)and; (2); (3) the other three species. Although this analysis strongly supported the monophyly of all specimens of, two geographical isolates were apparent.

Fig.2 NJ tree based on the analysis of COI sequences from six species of Nemipterus. Bootstrap values are shown near the nodes.

Fig.3 ML tree based on the analysis of COI sequences from six species of Nemipterus. Bootstrap values are shown near the nodes.

3.8 Genetic Distances Between DifferentSpecies

Table 1 summarizes the genetic distances between species of. Interspecific distances ranged from 8.6%–20.0%, while intraspecific distances ranged from 0.0–0.3%, with the exception ofwhere the Indian seas specimens showed 2.7% divergence from those collected in the South China Sea. This divergence value is considerably higher than the normal intraspecific genetic distance in marine fish species (Ward., 2005).

3.9 Comparison

The specimens of.from the South China Sea and the Indian Ocean are morphologically indistin- guishable based on counts and proportional measure- ments (Table 2). However, there appears be some small but consistent differences in colour pattern, with speci- mens in the South China Sea having a silver belly, while those from the Indian Ocean isolate specimen have a yellow coloration (Fig.4).

Table 1 Genetic distances (K2P model) among six taxa belonging to the genus Nemipterus

Table 2 Comparison of morphological characters of N. japonicus from the Pacific and Indian Ocean

Notes:†Some measurements can not be made because of the uncomplete fish body. The count data are from the origin description by Bloch (1791).††Data from Russell (1986).

4 Discussion

was described by Bloch (1791) based on a single dried and mounted specimen (Fig.5) deposited in the Berlin Museum (ZMB 8147). Bloch’s original description is brief and his illustration (Bloch, 1791: plate 277, Fig.1. shown here as Fig.6) is inaccurate in some respects (.. the lateral line scale count is incorrect, it lacks a filamentous extension to the upper caudal lobe, and the greenish body color is not seen in life). However, the length of the pectoral and pelvic fins and the many yellow stripes along body from behind the head to the base of caudal fin are characteristic ofand together distinguish it from all other species of.

Bloch (1791) failed to mention where the type specimen was collected, although the namesuggests its origin from Japanese waters. However, in the more than two centuries since its description, very few specimens have been reported from Japan. Russell (1986, 1993) noted the distribution of the species in Japan based on five specimens from the Ryukyu Islands (USNM 6368), but the presence of.in Japan is questionable, and it is likely that the Ryukyu specimens might have been caught in Taiwan waters for sale in Japan (H. Motomura pers. com. 2013) .

As is the case with many of Bloch’s fishes, it seems likely thatcame from East Indies (Java). As pointed out by Paepke (1999: 27), many of Bloch’s species represent tropical species unknown in Japanese waters. Indeed, the vernacular ‘Japanese’ names of several fishes mentioned by Bloch are based on the Malay name ‘ikan’ for fish (Pethiyagoda and Gill, 2012). Cuvier (1828: 100) observed that though Bloch believed his specimens to be of Japanese origin, he might have been misled by the Dutch traders who sold him the specimens.

The 5’ region of the cytochromeoxidase subunit I (COI) has been adopted as the standard DNA barcode region for members of the animal kingdom. It has a strong track record in revealing cryptic species (Hebert., 2004; Ward., 2005). We used thisgene region to examine genetic diversity in. The NJ and ML trees both strongly support the conclusion that individuals of this species form two geographical isolates (South China Sea, Indian Ocean) with 2.7% divergence.

Fig.4 Nemipterus japonicus: A, NPPF1159, 90.4mm SL, Nayband National Park Coast, Bushehr, Iran, one of specimens from the Indian Ocean examined by Asgharian et al. 2011. Scale bar=2.0cm; B, NPPF 1160, 88.5mm SL, Nayband National Park Coast, Bushehr, Iran, one of specimens from the Indian Ocean examined by Asgharian et al. 2011. Scale bar=2.0cm; C, IOCAS 135FB00144-4, 142.8mm SL, Sanya City, Hainan Island, China. Scale bar=1.0cm. The belly’s color from lower jaw to base of pelvic fin (1, 2) is silver in the South China Sea specimens, while yellow in the Indian Ocean specimens; The belly’s color from base of pelvic fin to caudal peduncle (3) is yellow in both two form of specimens. The pictures of A and B are deposited in BOLD.

Fig.5 Type specimen of Nemipterus japonicus (ZMB 8147), in the Zoological Museum, Berlin. Scale bar=1.0cm.

Fig.6 Nemipterus japonicus, original figure from Bloch, 1791, plate 277, Fig.1.

Although the timing of diversification inremains uncertain, the congruent phylogeographic pat- terns in other Indo-Pacific marine fishes clearly indicate the role of shifting sea levels linked to climatic oscillations during the Pleistocene as an important cause of population isolation and differentiation (Gaither., 2011). In this study, the level of divergence between regional populations ofsuggests that gene flow between populations ofon the continental shelves of the Pacific Ocean and the Indian Ocean was interrupted by the Indo-Pacific Barrier (Fleminger, 1986) during Pleistocene sea level fluctuations. Further effort is needed to carry out an extensive collaborative study using a multi-genic analysis to examine the phylogeny of a wide range ofspecies and geographical isolates. Work also is needed to ascertain the respective geographical distributions of the West Pacific and Indian Ocean populations of.

Acknowledgements

This investigation was supported by IOCAS funding (2012IO060102). We are very grateful to Dr. Peter Bartsch, Curator of Fishes at the Museum für Naturkunde, Berlin, and to Professor Michael Tuerkay, Senckenberg Forschungsinstitut, Frankfurt, for providing a photograph of the holotype ZMB 8147, a copy of Bloch’s original description and other help with this paper. Dr. Hiromitsu Motomura, Kagoshima University Museum, Japan, provided information and photographs of specimens from the Philippines. We also thank Lian-Mei Shuai of the Marine Biological Museum, Chinese Academy of Sciences for her help in sorting the fish specimens. Prof. Jin-Xian Liu provided helpful suggestions on this paper, while Dr. Wei Jiang provided the photographs of the specimens from Hainan Island. Finally, we thank Drs. Chuan-Yan Yang, and Da-Hai Gao for their aid on various aspects of this study.

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(Edited by Ji Dechun)

DOI 10.1007/s11802-015-2609-x

ISSN 1672-5182, 2015 14 (1): 178-184

© Ocean University of China, Science Press and Springer-Verlag Berlin Heidelberg 2015

(March 1, 2014; revised April 20, 2014; accepted May 4, 2014)

* Corresponding author. E-mail:ningping@qdio.ac.cn