A preliminary deep-sea fish fauna list with new and northernmost records, in the continental shelf to the upper continental slope in the east ...

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Plankton Benthos Res 17(1): 95–99, 2022
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A preliminary deep-sea fish fauna list with new and
northernmost records, in the continental shelf to the upper
continental slope in the east Enshu-nada Sea, central Japan
Akinori Teramura1,*, Hiroshi Kohno2, Hiroshi Senou3, Kiyoshi Kikuchi1, Shotaro Hirase1 &
Taiga Kunishima4,*
1
  Fisheries Laboratory, Graduate School of Agricultural and Life Sciences, The University of Tokyo, 2971–4 Bentenjima, Nihi, Maisaka,
Hamamatsu, Shizuoka 431–0214, Japan
2
  Laboratory of Ichthyology, Tokyo University of Marine Science and Technology, 4–5–7 Konan, Minato, Tokyo 108–8477, Japan
3
  Kanagawa Prefectural Museum of Natural History, 499 Iryuda, Odawara City, Kanagawa 250–0031, Japan
4
  Wakayama Prefectural Museum of Natural History, 370–1 Funoo, Kainan, Wakayama 642–0001, Japan
Received 22 February 2021; Accepted 10 September 2021 Responsible Editor: Shigeaki Kojima
doi: 10.3800/pbr.17.95

           Abstract: We focus on the ichthyofauna on continental shelves and slopes of the Enshu-nada Sea̶an area of the northwest Pacific
           Ocean off the coast of Southern Japan as one of the essential trawl fishery grounds in Japan. We conducted market samplings of the
           trawl catches from the Enshu-nada Sea, and trawl surveys of bycatch species by commercial trawlers. As a result, 208 deep-sea fish
           species (92 families) were recorded, with 46 species first recorded from the Enshu-nada Sea, of which five species were the northern-
           most record for these species. The results reveal the ichthyofaunal diversity in the continental shelf and upper continental slope of the
           Enshu-nada Sea, providing biogeographic information on various species based on the vouchers.

           Key words: Checklist, Deep-sea fish, Japanese waters, Kuroshio Current, Northwest Pacific

   Deep-sea ecosystems are generally vulnerable to human                           slopes occur in the Enshu-nada Sea (Sato & Hoshino 1962;
impacts (Koslow et al. 2000). Above all, deep-sea fishes are                        Fig. 1) which allows the trawler fisherman to catch the various
facing heavy exploitation by human activities and have been                        fisheries across seasons. In addition, the Kuroshio Current
threatened globally (Tanner et al. 2018, Miyazaki et al. 2019).                    (hereafter, the Kuroshio) flowing close to the Enshu-nada Sea
While documentation of the species diversity is the first step                      maintains the warmer environment in this area, which result-
for considering effective fishery management procedures                              ing a highly productive fishing ground is formed. The trawl
and conservation measures (Frankhan et al. 2007, Moto-                             catches from the continental shelf and upper continental slope
mura 2018), basic information such as species list is limited in                   in the Enshu-nada Sea are landed in various fishing ports in
many deep-sea fishing grounds.                                                      the Aichi and Shizuoka prefectures (Achiha 2006).
   We focused on the Enshu-nada Sea (Toto-umi Sea), which                             Ichthyofaunal diversity of Enshu-nada Sea is also a prom-
is defined as the coastal area of the Pacific Ocean along the                        ising subject for biogeographic and taxonomic study. Sev-
central part of the Japanese mainland, from off the Irago Cape                      eral species were first described from the Enshu-nada Sea
to the Omaezaki Cape, as one of the most important fishing                          (e.g., Pseudoblennius totomius Jordan & Starks, 1904; Ow-
grounds in the Japanese seas (Fig. 1). The Enshu-nada Sea                          stonia totomiensis Tanaka, 1908). Moreover, many ichthyo-
is notable as a fishing ground for the bottom seine trawlers                        faunal studies have been conducted in the Suruga Bay and
probably due to its topographic and oceanographic characters.                      Kumano-nada Sea adjacent to the Enshu-nada Sea, provid-
The combination of the continental shelf, the shelf break̶                         ing important biogeographic knowledges for understanding
the location of the boundary zone between the continental                          ichthyofaunal diversity along the Pacific coast of Japan (e.g.,
shelf and slope̶with sandy-mud bottom, and the continental                         northernmost records and first records in Japan, Shinohara
                                                                                   & Matsuura 1997, Kobayashi et al. 1999, Kubo et al. 2012).
* Corresponding author: Akinori Teramura; E-mail, akifishes@yahoo.
                                                                                   While some checklists of marine fishes have reported in the
  co.jp, Taiga Kunishima; E-mail, taigakunishima@gmail.com                         Enshu-nada Sea off the Aichi Prefecture and the Mikawa Bay
  Supplementary materials may be found in the online version of this article.      (Nakajima 1975, 2003, 2007, Arao & Tamai 2011, Tamai et al.
96                                                          A. Teramura et al.

2012, Tamai & Arao 2013, 2021), only some deep-sea fishes                Enshu-nada Sea (Table S1; Fig. 1). In most cases, specimens
on the continental shelf and slope.                                     were identified onboard or in a laboratory by photography,
   In this study, we conducted market samplings at the Mai-             basically following Nakabo (2013) but also Kawaguchi &
saka Fishing Port (Maisaka FP) and trawl surveys on bycatch             Shimizu (1978) about Myctophidae and Kai & Tashiro (2019)
species by local trawlers for clarifying the deep-sea fish fauna         about Zeidae. The taxonomic system and scientific names fol-
of the Enshu-nada Sea. Maisaka FP is a major fishing port in             low Motomura (2020). At least a single individual specimen
Hamamatsu City (Fig. 1). The depth of fishing grounds ranges             was kept for each species and inspected in the laboratory dur-
from 50–500 m and the fishing gears, fisheries targets, and               ing the surveys, but large-sized species or species with high
operating depth vary depend on the seasons. From September              commercial value were only photographed. In the laboratory,
to May, bottom trawl fisheries are operated in the continental           specimens were fixed with 10% formalin and subsequently
shelf and upper continental slope at a depth of 100–500 m               replaced with 70% ethanol. All specimens were deposited in
(mainly 300 m), targeting Japanese lobster Metanephrops ja-             The Kagoshima University Museum (KAUM), Kanagawa
ponicus (Tapparone-Canefri, 1873), Deep-sea smelt Glossan-              Prefectural Museum Natural History (KPM-NI), or Waka-
odon semifasciatus (Kishinouye, 1904), and Greeneyes Chlo-              yama Prefectural Museum of Natural History (WMNH).
rophthalmus albatrossis (Jordan & Starks, 1904). From June                 In total, we recorded 208 species in 161 genera in 92 fami-
to August, bottom seine nets are operated on the continental            lies, 142 from trawl surveys and 156 from port sampling,
shelf at a depth shallower than 100 m, targeting large-head             respectively. (Table S2). Forty-six species were first reported
hairtail Trichiurus japonicus (Temminck & Schlegel, 1844)               from the Enshu-nada Sea (Figs. S1–S4; Table S2), of which
and Japanese jack mackerel Trachurus japonicus (Temminck                five species are the northernmost record of each species
& Schlegel, 1844). Thus, we expected that trawl catch landing           [Lophiodes insidiator (Regan, 1921); Lophiodes mutilus (Al-
at Maisaka FP are representatives of the local fish fauna from           cock, 1894); Chlorophthalmus sp. 2 (Japanese name: Bake-
the continental shelf and upper continental slope at 50–500 m           aomeeso sensu Nakabo & Kai 2013); Coelorinchus cingulatus
depth in the western Enshu-nada Sea.                                    Gilbert & Hubbs, 1920; Parascombrops yamanouei Schwar-
   In the market sampling, fish samples were obtained 25                 zhans, Prokofiev & Ho, 2017]. We hypothesized two scenarios
times at Maisaka FP or local fish markets between 2015 and               for these new or northernmost records: one is shifts in distri-
2020 (Table S1), especially, at least once a month between De-          bution range of some demersal fish because of global climate
cember 2018 and April 2020. We also conducted eight trawl               change, as suggested by previous studies (Perry et al. 2005,
surveys on the commercial trawlers “Kyoei-maru” on the                  Engelhard et al. 2014); another is occasional transportation
upper continental slopes at 200–500 m depth in the eastern              of tropical fishes because of the influence of the Kuroshio.

Fig. 1. Topographic map of the Enshu-nada Sea. Solid lines indicate depths at 50 m, and bold lines indicate water depth at 300 m. The
gray circle indicates the Maisaka fishing port (Maisaka FP). Dotted line shows the trawl fishery operating area. This Map was created by first
author using the Generic Mapping Tools, Version 5.4.3 (Wessel et al. 2013).
Deep-sea fish fauna in the Enshu-nada Sea                                                    97

Usually, the Kuroshio strongly affect the fish fauna also in the      this study. The results provided relevant biogeographic data
deep-sea areas in Japanese water bodies through its roles as        to elucidate the species diversity of deep-sea around central
the driver of the water temperature (e.g., Kimura & Sugimoto        Japan, adding northernmost and new records of deep-sea fish-
1990, Nakamura & Uzaki 2007) and the transporter of fish             es in the Enshu-nada Sea. In the future, more detailed ich-
eggs and larvae from southern to northern regions (Kasai et         thyofaunal studies targeting the mesopelagic fauna, seasonal
al. 2002, Sassa et al. 2004). Even in Enshu-nada Sea, high wa-      change of community structure, and exploration of indicator
ter temperature was recorded there than usual when the Kuro-        species for each season will help clarify the species diversity
shio approached the its coastal areas between 2017 and 2020         and its dynamics of the deep-sea area in the Enshu-nada Sea.
(Sugimoto et al. 2021). Although we cannot discuss the rea-
son several northernmost record species were collected at the
                                                                                Electronic supplementary material
present stage in detail, marine environmental events, such as
global climate change and the Kuroshio large meander, might            The online version of this article (doi: xxxxxx/pbr.xx.xxx)
have influenced the distribution of these deep-sea fishes as          contains supplementary material:
well as in this area. Monitoring the local deep-sea fish fauna          Fig. S1. Fresh specimens which represent the first record
for a long period hopefully clarify the factors for the north-      of each species collected from the Enshu-nada Sea. a KPM-
ernmost and new records of deep-sea fishes in the Enshu-nada         NI 40426: Galeus nipponensis; b KPM-NI 40422: Etmopterus
Sea in this study.                                                  molleri; c KPM-NI 42561: Bathycongrus baranesi; d KPM-NI
   In the Enshu-nada Sea, the most abundant family was Mac-         52252: Japonoconger sivicolus; e KPM-NI 51368: Acromycter
rouridae (14 species), followed by Acropomatidae (9 species),       nezumi. Photographs were taken by Hiroshi Senou (a–c) and
                                                                    Akinori Teramura (d, e).
Triglidae (9 species), and Myctophidae (7 species) (Fig. 2).
                                                                       Fig. S2. Fresh specimens which represent the first record
In particular, Macrouridae, Acropomatidae, and Triglidae are
                                                                    of each species collected from the Enshu-nada Sea. a KPM-NI
dominant taxa in the continental shelf and upper continental
                                                                    53030: Diplophos orientalis; b KPM-NI 59908: Astronesthes iji-
slope in this study, which might be explained by the vast
                                                                    mai; c KPM-NI 42564: Chlorophthalmus sp. (Japanese name:
sandy-mud bottoms that they prefer because these families
                                                                    Bake-aomeeso, sensu Nakabo & Kai, 2013); d KPM-NI 53106
were recorded in a similar substrate in other areas (Furuhashi
                                                                    Dasyscopelus asper; e KPM-NI 41862: Diaphus suborbitalis;
et al. 2010, Kubo et al. 2012, Shinohara & Matsuura 1997,           f KPM-NI 59786: Diaphus garmani; g KPM-NI 42823: Poly-
Shinohara et al. 2001, 2005, 2009). Some mesopelagic species        mixia longispina; h KPM-NI 42690: Polymixia berndti; i KPM-
are also recorded from the bottom trawl catches (e.g., Mycto-       NI 52278: Coelorinchus anatirostris; j KPM-NI 40429: Coelo-
phidae and Stomiidae species), while most of the specimens          rinchus jordani; k KPM-NI 51342: Coelorinchus kishinouyei; l
are partially digested. We thus assume that the other fishes         KPM-NI 40420: Coelorinchus longissimus; m KPM-NI 59782:
have vomited them after landing. In addition, the mesopelagic       Coelorinchus cingulatus; n KPM-NI 42795: Malacocephalus
species are less abundant in the present study than in other        nipponensis; o Homostolus acer; p KPM-NI 40418: Lophiodes
regions (off the Pacific Coast of Boso Peninsula: Miya et al.         insidiator; q KPM-NI 59788: Lophiodes mutilus; r KPM-NI
1995, Suruga Bay: Shinohara & Matsuura 1997), probably              59788: Lophiodes mutilus. Photographs were taken by Akinori
due to the fishing gears targeting bottom fishes. Mid-water           Teramura (a, b, d, f, g, i, j, k, m, n, o, r), Hiroshi Senou (c, e, h, l,
trawl surveys targeting mesopelagic fishes may elucidate the         p), and Shuya Kato (q).
fish fauna of the Enshu-nada Sea in future.                             Fig. S3. Fresh specimens which represent the first record
   Overall, we summarized the records of deep-sea fishes             of each species collected from the Enshu-nada Sea. a KPM-NI
caught by bottom trawl fisheries in the Enshu-nada Sea in            59775: Malthopsis kobayashii; b KPM-NI 59776: Malthopsis
                                                                    annulifera; c KPM-NI 47947: Beryx mollis; d KPM-NI 47947:
                                                                    Beryx mollis, a white arrow indicates posterior nasal apertures;
                                                                    e Beryx decadactylus; f KPM-NI 51348: Zenopsis filamentosa; g
                                                                    KPM-NI 39721: Macroramphosus japonicus. Photographs were
                                                                    taken by Akinori Teramura (a–f) and Hiroshi Senou (g).
                                                                       Fig. S4. Fresh specimens of the first record of each species
                                                                    from the Enshu-nada Sea. a KPM-NI 56365: Ebosia bleekeri;
                                                                    b KPM-NI 41788: Pterygotrigla macrorhynchus; c KPM-NI
                                                                    42664: Lepidotrigla hime; d KPM-NI 41642: Peristedion liorhyn-
                                                                    chus; e KPM-NI 41640: Cubiceps whiteleggii; f KPM-NI 47950:
                                                                    Malakichthys barbatus; g KPM-NI 41786: Parascombrops yama-
                                                                    nouei; h KPM-NI 52265: Epinephelus latifasciatus; i KPM-NI
                                                                    52210: Evynnis cardinalis; j KPM-NI 59794: Erythrocles micro-
                                                                    ceps; k KPM-NI 51372: Emmelichthys struhsakeri; l KPM-NI
                                                                    59920: Dexistes rikuzenius; m Symphurus orientalis; n KPM-NI
                                                                    59797: Paratriacanthodes retrospinis. Photographs were taken
Fig. 2. Histogram of the fish composition of the 92 families         by Hiroshi Senou (a, b, d, e, g, h), Akinori Teramura (c, f, i, j, k,
surveyed in the Enshu-nada Sea.
98                                                             A. Teramura et al.

l, m, n).                                                                    English abstract)
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(b) Foetorepus paltivelis without specimens. All Photographs
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were taken by Akinori Teramura.                                            Koslow JA, Boehlert GW, Gordon JDM, Haedrich RL, Lorance P, Parin
   Table S1. Data on surveys on the Maisaka fishing port.                     N (2000) Continental slope and deep-sea fisheries: implications for a
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and slope in the East Enshu-nada Sea between 2015 and 2020.                Miya M, Yamaguchi M, Okiyama M (1995) Midwater Fishes off the
Bold texts and underlined texts indicate new and northernmost                Pacific Coast of Boso Peninsula, Central Japan: Species Composition,
records. Circles represent target species of trawl fishery in the             Abundance, Biomass, and Zoogeographic Affinities. Jpn J Ichthyol 42:
                                                                             237–248.
Enshu-nada Sea.
                                                                           Miyazaki Y, Teramura A, Senou H (2019) Preliminary report on
                                                                             bycatch fish species collected from the Tokyo Submarine Canyon,
                                                                             Japan. ZooKeys 843: 117–128.
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