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New Fossils of Neogene Pricklebacks (Actinopterygii: Stichaeidae) from East Asia Новые Находки Неогеновых Стихеевых (Actinopterygii: Stichaeidae) Из Восточной Азии

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New Fossils of Neogene Pricklebacks (Actinopterygii: Stichaeidae) from East Asia Новые Находки Неогеновых Стихеевых (Actinopterygii: Stichaeidae) Из Восточной Азии ZOOSYSTEMATICA ROSSICA, 24(1): 128–137 25 JUNE 2015 New fossils of Neogene pricklebacks (Actinopterygii: Stichaeidae) from East Asia Новые находки неогеновых стихеевых (Actinopterygii: Stichaeidae) из Восточной Азии M.V. NAZARKIN* & Y. YABUMOTO M.В. НАЗАРКИН, Ё. ЯБУМОТО M.V. Nazarkin, Zoological Institute, Russian Academy of Sciences, 1 Universiteskaya Emb., St Petersburg 199034, Russia. E-mail: [email protected]. *Corresponding author Y. Yabumoto, Kitakyushu Museum of Natural History and Human History, 2–4–1, Higashida, Yahatahigashi-ku, Kitakyushu, 805-0071 Japan. E-mail: [email protected] Two new findings of fossil pricklebacks (Pisces: Stichaeidae) from the Miocene deposits of Honshu Island (Japan) and Kamchatka Peninsula (Russia) are described. Both specimens are incomplete and cannot be identified below either family or subfamily level. Nevertheless, these findings suggest wide distribution of stichaeids in the North-Western Pacific during the Mio- cene. The specimen from the deep-sea Bessho Formation of Honshu represents the first fossil record of pricklebacks of the subfamily Lumpeninae. Описаны две новые находки ископаемых стихеевых рыб (Pisces: Stichaeidae) из отложе- ний миоцена острова Хонсю (Япония) и полуострова Камчатка (Россия). Оба изучен- ных образца очень фрагментарны, и не могут быть определены ниже уровня семейства или подсемейства. Однако эти находки показывают, что семейство стихеевых рыб было широко распространено в середине миоцена в северо-западной части Тихого океана. Эк- земпляр из глубоководной формации Бесшо острова Хонсю является первым обнаруже- нием подсемейства Lumpeninae в ископаемом состоянии. Key words: fossil pricklebacks, Miocene, Russia, Japan, Actinopterygii, Stichaeidae, new finds Ключевые слова: ископаемые стихеевые, миоцен, Россия, Япония, Actinopterygii, Stichaeidae, новые находки INTRODUCTION by long-based dorsal fin, which consists of only or, less often, mostly of spines. Togeth- Recent pricklebacks, family Stichaeidae, er with other morphologically similar fami- are widely distributed in the coastal marine lies, pricklebacks belong to the suborder waters of the Northern Hemisphere. They Zoarcoidei assigned to either Perciformes are common members of littoral and near (Nelson, 2006) or Cottiformes (Imamura & shore fish communities (Makushok, 1958), Yabe, 2002; Wiley & Johnson, 2010). though some species occur at depths more Based on a detailed morphological than 200 m. Most stichaeids inhabit the analysis of the Stichaeidae and closely re- North Pacific, and only few species occur lated families, Makushok (1958) divided in the Northern Atlantic and in the Arctic pricklebacks into the eight subfamilies and (Andriashev, 1954; Makushok, 1958; Lind- proposed a hypothesis on their interrela- berg & Krasjukova, 1975). Pricklebacks are tionships. Mecklenburg & Sheiko (2004) usually small elongated fishes, characterized reviewed later advances in the pricklebacks © 2015 Zoological Institute, Russian Academy of Scienсes NAZARKIN M.V. & YABUMOTO Y. NEOGENE STICHAEIDAE FROM EAST ASIA 129 taxonomy (i.e. Makushok, 1961a, 1961b, 1973; Shiogaki, 1984; Stoddard, 1985; Miki, 1985; Yatsu, 1985, 1986; Miki et al., 1987; Follett & Powell, 1988; Follett & Anderson, 1990; Anderson, 1994; Kimura & Ji- ang 1995; Posner & Lavenberg, 1999) and listed six stichaeid subfamilies diagnosed by mor- phological characters. The re- sults of recent phylogenetic studies based on molecuar data (Kartavtsev et al., 2009; Rad- chenko et al., 2009, 2010, 2012; Turanov et al., 2012; Kwun & Kim, 2013) do not support pre- vious classifications of prickle- backs based on morphology in- cluding osteology. The greater taxonomic di- versity of recent pricklebacks in the Northern Pacific suggests that they had originated from this region. This assumption is consistent with the known pricklebacks fossil findings, which are unique to North- Western Pacific. Until recently, fossil stichaeids were found in the two localities in East Asia: Honshu Island (Japan) and Sakhalin Island (Russia) (Fig. 1). Stichaeus matsubarai Niino Fig. 1. Sketch map of the Sea of Japan and the Sea of Okhotsk (subfamily Stichaeinae) was showing localities of the Miocene pricklebacks: 1 – Kavran- described from the Miocene Utcholok Bay; 2 – Agnevo; 3 – Azuma; 4 – Amikake. deposits near Azuma, Gunma Prefecture, Japan (Niino, 1951; Yabumoto & Uyeno, 1994). Five species of pricklebacks are members of recent sub- fossil pricklebacks were found in the Middle families. Most of fossil species belong to the Miocene (Serravalian) Agnevo Formation recent genera, with only one genus, Nivchia, of western Sakhalin, Russia: Ernogrammus presently extinct. litoralis Gretchina, 1980, Stichaeus brachi- Examination of fossil fish collections de- grammus Nazarkin, 1998, and Stichaeopsis posited in the National Museum of Nature sakhalinensis Nazarkin, 1998 (Stichaeinae); and Science (NSMT, Tokyo, Japan) and Ascoldia agnevica Gretchina, 1980 (Opisto- Borisyak Paleontological Institute, Russian centrinae); and Nivchia makushoki Nazar- Academy of Sciences (PIN, Moscow, Rus- kin, 1998 (Xiphisterinae) (Gretchina, 1980; sia) revealed two previously undescribed Nazarkin, 1998). All mentioned Miocene specimens of fossil pricklebacks from Hon- © 2015 Zoological Institute, Russian Academy of Scienсes, Zoosystematica Rossica 24(1): 128–137 130 NAZARKIN M.V. & YABUMOTO Y. NEOGENE STICHAEIDAE FROM EAST ASIA shu Island (Japan) and Kamchatka Penin- stones and aleurolites, with layers of tuffa- sula (Russia) (Fig. 1). Although both speci- ceous sandstones, lignite and shell-rocks. mens are incomplete, they provide new im- It had been deposited in the comparatively portant information on the distribution and shallow-water conditions (Sinelnikova et diversity of Neogene stichaeids. The length al., 1985). The remains of plants, numer- of dorsal-fin spines in both specimens dis- ous bivalves and gastropods, echinoderms, tinguishes them from the two closely re- worms and sea mammals are abundant in lated families also having the entirely spi- those beds. The rich assemblage of bivalves nous dorsal fin – gunnels (Pholidae) with indicates a relatively warm littoral and sub- a shorter dorsal-fin spines and wolffishes littoral environment (Sinelnikova et al., (Anarhichadidae) with much longer and 1985). The age of this formation is defined slender dorsal-fin spines. In this paper, we as Middle-Late Miocene (Gladenkov, 1978; describe these new findings and provide Gladenkov et al., 1986). some remarks on geographical distribution Osteological features of recent stichaeids of stichaeids in the Miocene. were studied based on X-ray photos of the following specimens from the main ich- MATERIAL, LOCALITIES thyological collection of the Zoological AND METHODS Institute, Russian Academy of Sciences (ZIN; St Petersburg, Russia): Alectrias gal- NSM PV 22683, a single partial skel- linus (Lindberg, 1938): ZIN 52834; Ascold- eton of the caudal region from the black ia variegata Pavlenko, 1910: ZIN 40310, shale (mudstones) of the Bessho Forma- 40361; Bryozoichthys lysimus (Jordan et tion, Honshu, Japan, is represented by the Snyder, 1902): ZIN 44739; Chirolophis sny- molds on the matrix with most skeletal el- deri (Taranetz, 1938): ZIN 54503; Dictyo- ements preserved as whitish mineralized soma burgeri van der Hoeven, 1855: ZIN bones. The Bessho mudstone formation is a 55128; Lumpenella longirostris (Evermann part of the Neogene sequence of Fossa Mag- et Goldsborough, 1907): ZIN 53762; Opis- na. It showed typical flysch facies (Suzuki, thocentrus ocellatus (Tilesius, 1811): ZIN 1982) and yielded fossil bivalves, indicating 54194; Stichaeus punctatus (Fabricius, deep-sea hydrothermal activity (Takeu- 1780): ZIN 54489, 54501, 54502; Stichae- chi, 2004). The Bessho Formation is well opsis nevelskoi (Schmidt, 1904): ZIN 52837. known due to the reach assemblage of fossil Measurements were made with calipers organisms – plants, echinoderms, bivalves to the nearest 0.1 mm. Drawings were based and cephalopods, elasmobranchs and deep- on digital photos. water teleosts, birds and sea mammals – discovered in its beds (Ohe & Koike, 1988; DESCRIPTION Matsuoka et al., 1998; Kawase & Koike, Order PERCIFORMES 2003, 2006; Kohno et al., 2007; Koike et al., 2008a, b; Takakuwa et al., 2009; Ishida et Family STICHAEIDAE Gill, 1864 al., 2009). The age of the Bessho Formation Subfamily LUMPENINAE Jordan is defined as Early-Middle Miocene, 13–16 et Evermann, 1898 mya (Takeuchi, 2004). PIN 3181/1050, a mold of the caudal Gen. et sp. indet. part of a fish skeleton on the thin-grained (Fig. 2) sandstone without any bones remaining, Material. NSM PV 22683, part and counter- from the western coast of Kamchatka, Rus- part of the caudal region; ca. 36°27’N 138°10’E, sia, was found in an outcrop of the Etolona Sakaki Town, Amikake, Oami, Nagano Prefec- Formation. This formation contains alter- ture, Honshu, Japan; Bessho Formation, Early- nation of conglomerates, pebbles, sand- Middle Miocene. © 2015 Zoological Institute, Russian Academy of Scienсes, Zoosystematica Rossica 24(1): 128–137 NAZARKIN M.V. & YABUMOTO Y. NEOGENE STICHAEIDAE FROM EAST ASIA 131 Fig. 2. Lumpeninae gen., sp. indet., specimen NSM PV 22683, photo (a) and outline drawing (b) based on part and counterpart: ar, anal rays; ds, dorsal fin spines; hhy, hypaxial hypural; hy 4+5, fused hypural 4+5; ptf, pterigiophores; uc, utostylar center. Description. The fossil represents an observed above the preural centra 10–12 incomplete caudal portion of the skeleton as a result of
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