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Zoogeography and Systematics of Shallow Water Marine East Asian Fishes

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Zoogeography and Systematics of Shallow Water Marine East Asian Fishes KOREAN JOURNAL OF ICHTHYOLOGY, Vol. 21 Supplement, 38-43, July 2009 Received : April 21, 2009 ISSN: 1225-8598 Revised : May 29, 2009 Accepted : July 3, 2009 Zoogeography and Systematics of Shallow Water Marine East Asian Fishes By Tetsuji Nakabo The Kyoto University Museum, Kyoto University, Kyoto 606-8501, Japan 1. INTRODUCTION Anarhichas orientalis and Pholidae, which are distributed in the Okhotsk and Bering Seas across to the northern Examples of marine fishes from East Asia were first Pacific coast of North America. studied by colleagues of Carl von Linnaeus in the 18th Century. Carl P. Thunberg, one of the best-known stu- Area 2.2-Pohai and Yellow Seas-rocky-reef fishes dents of Linnaeus, visited Japan to collect plant and ani- These include Hemilepidotus villosus, Sebastes schle- mal specimens whilst a naturalist employed by Vereeni- gelii and Sebastes koreanus, the last-mentioned being ghde Oost Indische Compagnie. Through the agency of endemic (Kim and Lee, 1994). All species have cold- the latter, Martin Houttyun received and later described water origins, the population of Hemilepidotus villosus (Houttyun, 1782) 42 fish species from Japan. In the mid- being well separated from that of northern Japan. dle of the 19th Century, many other Japanese fishes were described by Temminck and Schlegel (1843 1850) and ~ Area 2.3-Temperate area-rocky-reef fishes Richardson (1846). However, the culmination of these early studies were the comprehensive works of Jordan, This area extends from southern Hokkaido southward Tanaka and Snyder (1913) and Jordan and Metz (1913), to northern and western Kyushu along the Sea of Japan whereby almost all of the fishes of Japan and Korea coast, and southward to eastern Kii Peninsula along the became known to the western world. Pacific coast, the Seto Inland Sea, Tsushima Islands, The marine fish fauna in the shallow waters of East Cheju Island and the southern coast of the Korean Penin- Asia is basically comprised of warm and cold water fish- sula. Resident fishes include Sebastes, Hexagrammos es. This was initially recognized by Tanaka (1931), who otakii, H. agrammus, Dictyosoma burgeri, D. rubrima- demonstrated biogeographical boundaries between north- culatum, Pholis nebulosa, P. crassispina, Pseudoblennius, ern and southern fishes along the Sea of Japan and Paci- Embiotocidae (Ditrema and Neoditrema) and Pterogob- fic coasts. Later, Nakabo (2002) divided the marine fish ius, all being endemic. fauna of East Asian waters into 17 areas, according to Most of the fishes listed here have North Pacific cold habitat characteristics and water temperatures. In this water origins. For example, shallow water Sebastes have paper, relationships between the zoogeographic areas and their origins in the North Pacific, having subsequently systematics of shallow water fishes are discussed. speciated in East Asian waters separately from those off the Pacific coast of North America (Kai et al., 2003). Among the former, the Sebaste inermis complex have 2. SHALLOW WATER REEFS been recently recognized as comprising S. inermis, S. ventricosus and S. cheni, endemic temperate-water rocky- Area 2.1-Subarctic-rocky-reef fishes reef fishes (Kai and Nakabo, 2002, 2008; Kai et al., 2002). This area includes the Pacific coast from Tohoku Dis- Some of the temperate-water fishes have related spec- trict northward to the Kurile Islands, Sea of Japan coast ies or genera distributed along the Pacific coast of North northward from Tsugaru Strait, Okhotsk Sea coast of America, viz. Embiotocidae and Hexagrammos, as point- Hokkaido, and the coast of Primorsky Kray northward ed out by Ekman (1953). Embiotocidae has warm-water from Chongin. Resident fishes include Sebastes, Hexag- origins and is believed to be related to Chichlidae and rammos, Cyclopteridae, Liparis, Chirolophis, Stichaeus, Pomacanthidae (Mabuchi et al., 2007). In addition, Ditre- *Corresponding author: Tetsuji Nakabo Tel: 81-75-753-3279, ma temminckii has recently been recognized as compris- Fax: 81-75-753-3277, E-mail: [email protected] ing three distinct taxa, D. temminckii temminckii, D. — 38 — http://www.fishkorea.or.kr Zoogeography and Systematics of Shallow Water Marine East Asian Fishes 39 temminckii pacificum, and D. jordani by Katafuchi and microphthalmus, Hapalogenys sennin, Oplegnathus Nakabo (2007). Pterogobius (Gobiidae) clearly has Indo- punctatus, Calotomus japonicus, Prionurus scalprum, West Pacific origins. Sphyraena iburiensis and Ostracion immaculatus. Fishes Furthermore, Pacific Ocean and Sea of Japan populat- endemic to the overall area include Lutjanus ophuysenii, ions have been found for D. temminckii, Pterogobius Chaetodon nippon, Girella leonina and Pseudolabrus zonoleucus and P. elapoides, corresponding to coasts eoethinus. Ophisurus macrorhynchus, Seriola dumerili, adjacent to the Kuroshio and Tsushima Currents (Katafu- Pseudocalanx dentex, Upeneus tragula, Parupeneus chi and Nakabo, 2007; Akihito et al., 2008). chrysopleuron, Pempheris and Sphyraena obtusata are widely distributed in the Indo-West Pacific or to the tro- Area 2.4-Temperate area - rocky-reef fishes with pical Atlantic. warm water origins Among the species endemic to the Pacific coast of sou- thern Japan, Sphyraena iburiensis forms a clade with S. This area includes the Sea of Japan coast of Niigata pinguis, widely distributed in the Indo-West Pacific (Do- Prefecture southward to northern and western Kyushu, iuchi and Nakabo, 2005, 2007). Accordingly, S. iburien- the Pacific coast of Sendai Bay southward to southern sis may have speciated in East Asian waters from a Kyushu, the Seto Inland Sea, Tsushima Islands, Cheju common ancestor with S. pinguis. Gerres equulus and Island, southern coast of the Korean Peninsula, and the G. microphthalmus both have relatives in Southeast coast of southern China from Taiwan-Fujian southward Asian waters (Iwatsuki et al., 1999, 2002), along with to Tonking Bay, the overall area closely approximating Hapalogenys sennin (Iwatsuki and Nakabo, 2005). to the “Japan Region” of Briggs (1974). Fishes of the Some fishes of Area (s) C and/or D, viz. Girella and region, with warm-water Indo-Pacific origins, include Psedolabrus, show anti-equatorial distribution, having Myliobatis tobijei, Conger japonicus, Syngnathus schle- temperate region relatives in the Southern Hemisphere. geli, Hippocampus coronatus, Pterois lunulata, Sebastis- Furthermore, East Asian Girella and Pseudolabrus may cus, Lateolabrax, Epinephelus septemfasciatus, E. aka- have evolved from Southern Hemisphere stock (Yagishi- ara, Apogon kinesis, A. semilineatus, Parapristipoma ta and Nakabo, 2000, 2003; Mabuchi and Nakabo, 1997; trilineatum, Hapalogenys, Sparus sarba, Acanthopagrus Mabuchi et al., 2004). Hapalogenys and Oplegnathus schlegelii, Chaetodon modestus, Goniistius, Chromis also have anti-equatorial distribution (Hoese et al., 2006). notata, Rhyncopelatus oxyrhynchus, Girella punctata, Oplegnathus fasciatus, Pseudolabrus sieboldi, Semicos- Area 2.6-Subtropical area - coral and rocky reef syphus reticulatus, Choerodon azurio, Sphyraena ping- fishes uis, Sphyraena japonica, Petroscirtes breviceps, Siganus fuscescens and Rudarius ercodes. Among them, Conger This area includes the Ryukyu and Ogasawara Islands, japonicus, Hippocampus coronatus, Acanthopagrus sch- the fishes being both tropical or subtropical and widely legelii, Goniistius, Chromis notata, Oplegnathus fascia- distributed in the Indo-West Pacific, viz. Lamnidae, My- tus, Rudarius ercodes and Stephanolepis cirrhifer are not liobatidae, Muraenidae, Serranidae, Apogonidae, Lutja- distributed off the South China Sea Chinese coast, nidae, Lethrinidae, Haemulidae, Pomacentridae, Chaeto- whereas Nemipterus virgtus, Chaetodon modestus, Siga- dontidae, Pomacanthidae, Labridae, Scaridae, Neosynchi- nus fuscescens, Pterois lunulata, Epinephelus septemfas- ropus, Pterosynchiropus, Gobiidae, Acanthuridae, Siga- ciatus, Sparus sarba, Sphyraena pinguis and Petroscirtes nidae and Balistidae. breviceps. are widely distributed in the West Pacific or Indo-West Pacific. 3. SANDY-MUDDY BOTTOMS ON Area 2.5-Temperate to subtropical area - rocky-reef CONTINENTAL SHELVES fishes Area 3.1-Subarctic area - demersal fishes This area includes the Pacific coast of southern Japan from Boso Peninsula southward to southern Kyushu and This area extends from the Pacific coast northward the coast of southern China from Taiwan-Fujian south- from Miyagi Prefecture to Hokkaido, and includes the ward to Tonking Bay. The fishes typically have warm- northern Sea of Japan northward from both Tsugaru Str- water Indo-West Pacific or Indo-Pacific origins. Many ait and Ch’ongin.˘ The fishes include Gymnocanthus, are endemic to the Pacific coast of southern Japan from Hemilepidotus, Artediellus, Agonidae (some species en- Boso Peninsula southward to southern Kyushu, viz. Pla- demic to the region) and Pleuronectes (except P. yokoha- tyrhina sinensis, Narke japonica, Dasyatis izuensis, mae; some other species endemic). Fishes in the area are Myripristis kochiensis, Scorpaenopsis cirrosa, Lates ja- widely distributed throughout the Okhotsk and Bering ponicus, Lateolabrax latus, Decapterus maruadsi, D. Seas, through waters off Alaska to the northern Pacific muroadsi, Leiognathus rivulatus, Gerres equulus, G. coast of North America. Cottiusculus species and Alcich- 40 Tetsuji Nakabo thys elongates are endemic. tives. For example, R. koreanus is endemic to the Yellow Sea, but has relatives (R. valenciennei and R. luntus) in Area 3.2-Temperate
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