Deep-Sea Research II 86–87 (2013) 140–147 Contents lists available at SciVerse ScienceDirect Deep-Sea Research II journal homepage: www.elsevier.com/locate/dsr2 Peanut worms of the phylum Sipuncula from the Sea of Japan with a key to species Anastassya S. Maiorova a,b,n, Andrey V. Adrianov a,b a b A.V. Zhirmunsky Institute of Marine Biology, Far East Branch of Russian Academy of Sciences, Palchevskogo 17, 690059 Vladivostok, Russia Far Eastern Federal University, Sukhanova 8, 690950 Vladivostok, Russia a r t i c l e i n f o a b s t r a c t Available online 6 August 2012 Sipunculan worms from the Russian waters of the Sea of Japan are still poorly investigated while they are much better known from the Japanese coast. The aim of this paper is to describe sipunculans from the Russian coast and from the deepest part of the Sea of Japan near the Primorye Province collected by SoJaBio expedition, and to provide keys for identification of sipunculan species from the Sea of Japan. At the Russian coast of the Sea of Japan only 8 valid species of sipunculans were found and identified: Golfingia margaritacea, G. vulgaris, Nephasoma capilleforme, N. wodjanizkii, Phascolion strombus, Thysanocardia nigra, Themiste hexadactyla ( ¼ T. pyroides), Phascolosoma agassizii. Taking into account 4 other valid species noted for this area, Nephasoma eremite, Thysanocardia catharinae, Themiste blanda and Phascolosoma scolops, which were not found, the sipunculan fauna of the Russian waters of the Sea of Japan now comprises 12 valid species. Nephasoma capilleforme and Nephasoma wodjanizkii are the first records for the North-West Pacific and the Sea of Japan. Species accounts include the most important taxonomic characters and specific biotope data. Accordingly, a key up to species level is provided. Totally, the fauna of the Sea of Japan is now estimated as having 31 valid species of sipunculans. & 2012 Elsevier Ltd. All rights reserved. Keywords: The Sea of Japan Peanut worm Sipunculan Introvert Tentacular crown Hooks Trunk papillae Contractile vessel 1. Introduction Sipunculans, or peanut worms, constitute a well distinguished monophyletic group of exclusively marine non-segmented coelom worms which are currently considered as a separate phylum Sipuncula. They are worldwide in distribution and live in a wide variety of marine habitats from intertidal waters to abyssaldepths and from polar to equatorial seas. After a series of taxonomic revisions, about 150 valid species have been calculated currently (see Cutler, 1994). The body is subdivided into two main regions: barrel-like trunk and long eversible introvert. The anteriormost introvert, termed head, bears a terminal tentacular apparatus, the ciliary tentacles of which function in gas exchange and in feeding. The size of sipunculans varies considerably with trunk of mature specimens (with retracted introvert) from 2–3 mm long in meiobenthic Phascolion psammophilum (see Rice, 1993) and 3 mm in Onchnesoma steenstrupii to 500 mm in Siphonosoma ingens, 550 in Sipunculus indicus (see Stephen and Edmonds, 1972), and even 600 mm in Sipunculus nudus (see Murina, 1977). n Corresponding author at: A.V. Zhirmunsky Institute of Marine Biology, Far East Branch of Russian Academy of Sciences, Palchevskogo 17, 690059 Vladivostok, Russia. E-mail address: anastasia.mayorova@gmail.com (A.S. Maiorova). 0967-0645/$ - see front matter & 2012 Elsevier Ltd. All rights reserved. http://dx.doi.org/10.1016/j.dsr2.2012.08.009 Most sipunculans are deposit feeders, although a few are filter feeders, or sestonophages, with elaborate tentacular crown (species of Thysanocardia and Themiste). Sipunculans consume detritus and fecal material as well as bacteria, algae, protozoans and small invertebrates. In turn, they are eaten by fish, gastropods and cephalopods, carnivorous worms, crabs, starfishes, anemones, and other predators, including men, and also used as a popular ‘fishbait’’. Sipunculans may attain large densities as great as 700 specimens per m2 in reef limestone, and up to about 8000/m2 in some soft sediments (Maiorova and Adrianov, 2010; Murina, 1984; Rice et al., 1983; Williams and Margolis, 1974). In soft sediments, some species live just a few centimeters below the surface but some large species of Sipunculus burrow down a meter in coarse or silty sands, making nearly vertical tunnels. Many species live in sand-filled cracks, crevices, or pockets in the rocks and may also be found in seagrass roots, algal holdfasts and rhizoids, within sponges and under algal mats, in byssal threads of mussels. Some species inhabit empty mollusk shells, polychaete tubes, and foraminiferan tests. Several species can bore into coral, rocks and in decaying whale skulls (Aspidosiphon, Cloeosiphon, Lithacrosiphon). Together with polychaetes, sipunculans are designated to be the most important bioeroders in many recent and fossil reefs. One species, Phascolosoma turnerae, was found in association with submerged wood, A.S. Maiorova, A.V. Adrianov / Deep-Sea Research II 86–87 (2013) 140–147 occupying burrows in the wood (see Rice, 1985). Some small sipunculans (e.g. Phascolion psammophilum) are interstitial in coarse sand and shell hash. One species, Phascolosoma saprophagicum, is found living only on the flesh of a decomposing whale skull at about 900 m (see Gibbs, 1987). Interestingly, that at the north coast of Sakhalin Island in the Okhotsk Sea, sipunculan Golfingia margaritacea (Sars, 1851) is an important component of the diet of grey whales, in some areas comprising more than 50% of benthic biomass (Maiorova and Adrianov, 2010). About 30 species of sipunculans were found in the abyssal depth (below 3000 m). Some eurybathic species are found between 10 and 4000 m over a wide range of temperatures but some species are restricted to cold water deeper than 3000 m and never found above this isobath. At least 13 species are known from the depth below 5000 m: Golfingia anderssoni (Theel, 1911); G. margaritacea (Sars, 1851); Golfingia muricaudata (Southern, 1913); Golfingia vulgaris (de Blainville, 1827); Nephasoma capilleforme (Murina, 1973); Nephasoma diaphanes diaphanes (Gerould, 1913); Nephasoma flagriferum (Selenka, 1885); Nephasoma minutum (Keferstein, 1862); N. schuettei (Augener, 1903); Phascolion lutense (Selenka, 1885); Phascolion pacificum (Murina, 1957); Onchnesoma magnibatha (Cutler, 1969); Apionsoma murinae, (Cutler, 1969), (see Cutler, 1994). Some species are common even at the ultra-abyssal depth (more than 6000 m) (N. minutum, G. muricaudata, G. anderssoni, P. lutense, P. pacificum), but it should be emphasized that the ultra-abyssal zone has no endemic species. Deeper cold-water species Sipunculus norvegicus Danielssen, 1869 is very common in seep communities near the mid-Atlantic ridge at the depth from 1000 to 3500 m and provides a large benthic biomass with high density in the local populations (thousands of specimens per m2). When first collected in such abundance in the deep-water seep communities during the MARECO (Mid-Ocean Ridges) expedition in July 2004, these large worms, all being in the box-corers with retracted introvert, were misinterpreted as priapulids and later identified to a species level (see Maiorova and Adrianov (2010)). Sipunculan fauna of the Far Eastern Seas (North-West Pacific) is still not sufficiently investigated. Data of sipunculan fauna from the Russian waters of the Bering and Okhotsk Seas and the Sea of Japan can be found in a number of reports (see Adrianov and Maiorova, 2010; Makarov, 1950; Maiorova and Adrianov, 2010; Morozov and Adrianov, 2002; Murina, 1975, 1977; Ostroumov, 1909; Popkov, 1993a, 1993b). In the Bering Sea, Makarov (1950) and Murina (1977) have noted 13 species of sipunculans. According to the modern synonymy (see Cutler, 1994), 10 of them are now considered as valid species. These species are G. muricaudata (¼G. appendiculata, in Murina, 1977), G. margaritacea (¼ Phascolosoma margaritaceum, Phascolosoma hudsonianum, Phascolosoma glossipapillosum, in Makarov, 1950), G. vulgaris, Nephasoma minutum, Nephasoma eremita (¼Phascolosoma eremite, in Makarov, 1950 and Golfingia eremite, in Murina, 1977), N. diaphanes (¼ Golfingia schuettei, in Murina, 1977), A. murinae (¼Golfingia murinae, in Murina, 1977), Phascolosoma agassizii (¼P. japonicum, in Makarov, 1950 and in Murina, 1977), Phascolion strombus (¼ Phascolion strombi, in Murina, 1977). In the Okhotsk Sea, about 15 species of sipunculans have been listed (see Makarov, 1950; Murina, 1977), but only 11 of them are now considered to be valid (see Cutler, 1994). These species are G. vulgaris, G. margaritacea ( ¼Phascolosoma margaritaceum, P. hudsonianum, P. wagini, in Makarov, 1950), N. eremita (¼P. eremite, in Makarov, 1950 and G. eremite, in Murina, 1977), N. minutum (¼Golfingia improvisa, in Murina, 1977), N. diaphanes (¼Golfingia schuettei, in Murina, 1977), Thysanocardia catharinae (¼Golfingia catharinae, in Murina, 1977), T. nigra (¼Phascolosoma pavlenkoi, in Makarov, 1950), Themiste blanda, P. agassizii (¼P. japonicum, in 141 Makarov, 1950 and in Murina, 1977;¼P. kurilense, in Murina, 1977), P. scolops, Phascolion strombus (¼P. strombi, in Makarov, 1950 and in Murina, 1977). In the Okhotsk Sea, we found four abundant species of peanut worms—Golfingia vulgaris, G. margaritacea, P. strombus and Phascolosoma agassizii (Maiorova and Adrianov, 2010). P. agassizii is the most common species in the Aniva Bay at the south-east coast of Sakhalin Island, the north frontier of the natural habit (areal) of this species. In the Sea of Japan, Makarov (1950) and Murina (1975, 1977) have listed respectively 10 and 21 species of sipunculans, most of them being noted for Japan and Tsushima strait. Only 17 of them are now considered to be valid (see Cutler, 1994). About eight of these species have been noted for Russian waters from Tatarsky strait to the Peter the Great Bay (Makarov, 1950; Murina, 1977). These species are: Golfingia margaritacea (¼Phascolosoma margaritaceum, P. glossipapillosum, P. hudsonianum, P. okinoseanum, in Makarov, 1950; ¼Golfingia ikedai, Golfingia glossipapillosa, G. nota, in Murina, 1977), G. vulgaris, Thysanocardia catharinae (¼G. catharinae, in Murina, 1977), T. nigra (¼Phascolosoma pavlenkoi, in Makarov, 1950; ¼Golfingia nigra, in Murina, 1977), Themiste blanda, Phascolosoma agassizii (¼ P. japonicum, in Makarov, 1950 and in Murina, 1977; ¼ P. golikovi, in Murina, 1975; ¼P. yesoense, in Murina, 1977), P. scolops, Phascolion strombus (¼P. strombi, in Makarov, 1950 and in Murina, 1977). Two new species, Thysanocardia melanium and Themiste maculosa, described from the Peter the Great Bay by Popkov (1993a, 1993b), later were recognized as junior synonyms of Thysanocardia nigra (see Cutler and Dean, 1997) and of Themiste pyroides (see Morozov and Adrianov, 2002). A number of reports are available on the sipunculan fauna from the Japanese and Korean waters (see Ikeda, 1904, 1924; Sato, 1930, 1934a,1934b, 1937; Cutler and Cutler, 1981; Cutler et al., 1984; Cutler, 1994; Cutler and Dean, 1997; List of Animals in Korea, 1998; Nishikawa and Ueshima, 2006). About 30 species of peanut worms have been noted for the eastern Sea of Japan, near the Japan Island and Tsushima strait (see Cutler et al., 1984). Taking into account taxonomic revisions (see Cutler, 1994), only 23 out of the listed species are considered to be valid. For comparison, at the Pacific (east) coast of Japan about 49 species have been listed (see Cutler et al., 1984). In the List of Animals in Korea (1998), nine species of peanut worms have been noted for Korean waters (Sipunculus nudus, T. nigra, Themiste hexadactyla, Antillesoma antillarum, Phascolosoma agassizii, P. albolineatum, P. japonicum, P. kurilense, P. scolops). According to Cutler (1994), Themiste hexadactyla is a junior synonym of T. pyroides and Phascolosoma japonicum and P. kurilense are considered as junior synonyms of P. agassizii, thus totally giving only seven valid species for this area. The aim of this paper is to describe sipunculans from the Russian coast and from the deepest part of the Sea of Japan near the Primorye Province collected by SoJaBio expedition, and to provide keys for identification of sipunculan species from the Sea of Japan. 2. Materials and methods Sipunculans were collected and identified in the course of field trips in the Russian water of the Sea of Japan during 2008–2010. Deep water sipunculans were collected during the Russian–German deepsea expedition within the bounds of the SoJaBio-project (Sea of Japan Biodiversity Studies) in August–September 2010. During the SoJaBio voyage, benthic specimens were collected alongside the deep-water transects from 500 to 3666 m near the Russian coast of the Sea of Japan. Sipunculans from the Sea of Japan, available in the museum collections, were also checked and identified. 142 A.S. Maiorova, A.V. Adrianov / Deep-Sea Research II 86–87 (2013) 140–147 Species accounts include quantitative characteristics and specific biotope data. The material sampled was maintained in aquaria and photographed alive for illustration of the tentacular apparatus and live coloration of the body, introvert, and tentacles. Worms were fixed with 10% formalin and then postfixed with 70% ethanol. Representatives of all species were examined for internal morphological observations and species identifications. Preparations of introvert hooks were made for all species possessing these structures. Sipunculans were examined using a light (Zeiss Axioplan) and scanning electron microscope (LEO-430). The examined specimens are deposited in the Museum of A.V. Zhirmunsky Institute of Marine Biology, Far Eastern Branch, Russian Academy of Sciences (Vladivostok). In taxonomic accounts we follow the synonymy suggested by Cutler (1994). 3. Results At the Russian coast of the Sea of Japan, we found and identified only eight valid species of sipunculans: G. margaritacea, G. vulgaris, N. capilleforme, N. wodjansky, P. strombus, T. nigra, T. hexadactyla (¼T. pyroides), P. agassizii. Taking into account four other valid species noted by Makarov (1950) and Murina (1977), N. eremite, T. catharinae, T. blanda and P. scolops, which we were not able to find, the sipunculan fauna of the Russian water of the Sea of Japan now comprises 12 valid species. The fauna of the Sea of Japan is now estimated as having 31 valid species of sipunculans. Phascolosoma agassizii, P. scolops, T. nigra and T. hexadactyla ( ¼T. pyroides) usually inhabit the shallow water areas (0–50 m), while G. margaritacea, G. vulgaris, N. capilleforme, N. wodjanizkii, P. strombus are more deep water species, most common at the depth from 50 to 1000 m. The deepest point where we found G. margaritacea is the Bogorov elevation composed of lava stones covered with mud at the depth 1699 m (SoJaBio). Phylum Sipuncula Linnaeus, 1766 Class Sipunculidea E. Cutler and Gibbs, 1985 Order Golfingiiformes E. Cutler and Gibbs, 1985 Family Golfingiidae Stephen and Edmonds, 1972 Genus Golfingia Lankester, 1885 Subgenus Golfingia (Golfingia) Lankester, 1885 Golfingia margaritacea (Sars, 1851) (Fig. 1 A–D) Material. Sea of Japan, near coast of Primorye Province, SoJaBio-2010: sta# A2-8, trawl 44156.0262 N; 137113.2535 E, 582–570 m depth, gravel and muddy sediment, five specimens; sta# B2-5, trawl, 42133.5243b N; 136117.5619 E, 1699 m depth, lava stones with mud, five specimens; sta# B6-9, trawl, 43109.4745 N; 135100.5842 E; 1075 m depth, muddy sediment, eight specimens. Description. Trunk slender, cylindrical, 23–90 mm long and 6–10 mm wide; introvert about equal to the trunk length. Only one young specimen has small scattered hooks, while all other mature specimens lack this character. Trunk is pale and lustrous. Ventral retractor muscles originate about 30% of trunk length to the posterior end. Dorsal retractor muscles originate in the anterior trunk, only 5–6% of trunk length to the posterior end. Gut with up to 35 loops. Nephridia are about 10–15% of trunk length. Discussion. Cutler (1994) has distinguished two subspecies— G. margaritacea margaritacea (Sars, 1851) and G. margaritacea ohlini (Theel, 1911). The last one is known only from the southern hemisphere and differs from the nominate form by having hooks (see Cutler, 1994). The only other species of the genus from the Sea of Japan, G. vulgaris, is well distinguished from G. margaritacea by rugose skin of the trunk and by the presence of well developed hooks in mature specimens. Distribution. This is the most widespread sipunculan species in the North-West Pacific. In the Sea of Japan, this species was previously known from Tatarsky strait and Sakhalin Island in the north to Tsushima strait in the south, alongside Korean, Japanese and Russian coast as well as in the middle of the Sea. Golfingia vulgaris (de Blainville, 1827) Fig. 1. A–D, Golfingia margaritacea. (A) Lateral view, introvert retracted; (B) young specimen, lateral view, introvert everted; (C) head with tentacular apparatus; (D) benthic sample from one Van Veen crab (0.025 m2) with amphipoda Ampelisca macrocephala. Scale bars: A, 20 mm; B, 10 mm; C, 2 mm; D, 50 mm. A.S. Maiorova, A.V. Adrianov / Deep-Sea Research II 86–87 (2013) 140–147 143 Genus Nephasoma Pergament, 1940 Subgenus Nephasoma (Nephasoma) Pergament, 1940 Nephasoma capilleforme (Murina, 1973) Material. Sea of Japan, Peter the Great Bay, near cape Gamov 421200 5500 N; 1311230 4700 E, 200–300 m depth, grab, 6 specimens. Description. Trunk spindle-shaped or thread-like, 3–12 mm long, pale, with dark papillae concentrated at anus and posterior end; introvert about equal to trunk length, with small scattered hooks, tentacular crown with 6 tentacles. Retractor muscles attached about 60% of trunk length to the posterior end. Gut with 12–15 loops. Nephridia about 15% of the trunk length, free. Discussion. This species is well distinguished from two other representatives of the genus Nephasoma from the Sea of Japan, N. eremita and N. wodjanizkii, by the presence of hooks and by elongate thread-like body. Trunk of N. capilleforme is usually swollen and golden colored anteriorly, in the vicinity of anus and nephridiopores, and possesses dark pigmented papillae at the posterior end. Distribution. The species was previously described only from deep water areas of the Pacific, from Southern Hemisphere and North-East Pacific, at the depth from 920 to 3450 m (see Murina, 1973, 1977). This is the first finding of N. capilleforme in the Sea of Japan and the North-West Pacific. Fig. 2. A–C, Themiste hexadactyla. (A) Lateral view, introvert everted (white arrow – collar with patchy distributed pigment; white arrowhead-pigmented anterior introvert); (B) head with tentacular apparatus; (C) hook. D–E, Themiste blanda. (D) ventral view, introvert everted; (E) hook. F–G, Thysanocardia nigra. (F) Lateral view, introvert everted; (G) head with tentacular apparatus. H–I, Phascolosoma agassizii. (H) lateral view, introvert everted; (I) hook (black arrowhead – triangle of hook; black arrow-clear streak). Scale bars: A–B, 5 mm; C, 50 mm; D, 3 mm; E, 50 mm; F, 5 mm; G, 3 mm; H, 5 mm; I, 50 mm. Material. Sea of Japan, Peter the Great Bay, near Nakhodka, 421310 06 N; 1321440 02 E, 200–300 m depth, grab, 10 specimens. Description. Trunk elongated, spindle-shaped, 4–15 mm long and 2–3 mm wide; introvert shorter than trunk, with digitiform tentacles and well developed scattered hooks in mature specimens. Trunk yellow-brown with dark brown papillae more concentrated at the anus and posterior end. Ventral retractor muscles originate about 30% of trunk length to the posterior end. Dorsal retractor muscles originate about 12% of trunk length to the posterior end. Gut with about 16 loops. Nephridia are about 6% of trunk length. Discussion. Cutler (1994) has distinguished two subspecies – G. vulgaris vulgaris (de Blainville, 1827) and G. vulgaris herdmani (Shipley, 1903). Animals belonging to the second subspecies differ from the nominate form by having radiating rows of dark spherical papillae at the posterior end, thus giving appearance of the pseudoshield. This is shallow, warm water subspecies while nominate form is mainly represented by deep and cold water specimens. This species is well distinguished from G. margaritacea by having well developed hooks in mature specimens. As noted by Cutler (1994), G. vulgaris is characterized by dark brown or black heavily papillated areas on the both ends of the trunk while whitish middle trunk looks smooth. In our specimens from the Sea of Japan, these dark brown papillae concentrating at the trunk ends are also scattered over the whole body. Largest of these specimens are also characterized by rugose skin with longitudinal wrinkles in the middle trunk and radial wrinkles at the posterior end, as noted for G. vulgaris herdmani. Distribution. This species is widespread in the North-West Pacific from shallow water to abyssal depth. In the Sea of Japan, this species was previously known only from deep water near the Russian and Korean coasts. This is the first record of this species from the Peter the Great Bay. Nephasoma wodjanizkii (Murina, 1973) Material. Sea of Japan, near coast of Primorye Province, SoJaBio-2010, sta# B7-8, trawl, 43113.6778 N; 135104.4447 E, 532 depth, muddy sediment, community of marine lilies Geliometra glacialis, 4 specimens. Description. Trunk 3.5 –23 mm in length; introvert 2–3 times longer than trunk, with short tentacles, without hooks. Trunk pale, brownish, with black pigmented anal area. Retractor muscles attached about 75–85% of trunk length to the posterior end. Gut with about 10 loops and 2 fixing muscles, oesophagus with 1 fixing muscle; spindle muscle present. Contractile vessel simple. Nephridia 20–25% of the trunk length, dark brown, rough, free; nephridiopores slightly anterior to the anus. Discussion. The species was described by Murina (1973) from the North-East Pacific based on a single specimen collected at 1100 m depth. In contrast to our material, small scattered hooks were found on the introvert of this specimen. Another specimen of N. wodjanizkii, but without hooks, was noted from the Canadian coast by Frank (see Cutler and Cutler, 1986; Frank, 1983). Thompson (1980) has described the new species, N. nicolasi, from Californian, which was revised by Cutler as a junior synonym of N. wodjanizkii (see Cutler and Cutler, 1986). No hooks have been seen in Canadian and Californian specimens. As noted by Cutler (see Cutler, 1994; Cutler and Cutler, 1986), in this genus some species have hooks as young individuals but lose them with age (deciduous hooks). In our specimens, nephridiopores open slightly anterior to the anus while in all other specimens nephridia are situated slightly posterior to the anus. Despite this difference, we identified specimens from the Sea of Japan as representatives of N. wodjanizkii. This species is well distinguished from other representatives of the genus Nephasoma from the Sea of Japan, N. capilleforme and N. eremita. In contrast to our species, N. capilleforme always has hooks and is characterized by thread-like body and loosely wound gut coil. Hookless N. eremita is characterized by nonpapillated trunk with transverse grooves in the body wall and possesses well developed digitiform tentacles. Also, in contrast to N. wodjanizkii, retractor muscles of N. eremita originate in the middle of posterior third of the trunk. 144 A.S. Maiorova, A.V. Adrianov / Deep-Sea Research II 86–87 (2013) 140–147 This is the first finding of N. wodjanizkii in the Sea of Japan and the North-West Pacific. Genus Thysanocardia (Fisher, 1950) Thysanocardia nigra (Ikeda, 1904) (Fig. 2F and G) Material. Sea of Japan, Peter the Great Bay, wide spread, muddy sediments, rhizomes of seagrass Zostera marina, clusters of mussels Crenomytilus grayanus, Modiolus kurilensis and Mytilus trossulus, 1–10 m depth, more than 100 specimens. Description. Trunk spindle-shaped, 20–60 mm long, 2–8 mm wide; introvert one and one-half or two times longer than trunk, with numerous tentacles arranged in dorsal and oral crowns, no hooks. Tentacles in living specimens with wide yellow apical bands followed by brown pigmentation. Trunk smooth, greyyellowish or gray, with minute papillae. Retractor muscles originate about 60% of trunk length to the posterior end. Gut with 18–54 loops, with two fixing muscles. Contractile vessel with numerous pink or red digitiform or branched villi. Nephridia about 30% of trunk length, free. Discussion. This species was initially described from Japan (see Ikeda, 1904) and redescribed in detail by Cutler et al. (1984). Later, this species was noted from the north part of the Sea of Japan (Sakhalin Island) (see Murina, 1977). The species, T. melanium, described by Popkov (1993a) from the Possiet Bay (Peter the Great Bay) in the west part of the Sea of Japan, was revised by Cutler and Dean (1997) as a junior synonym of T. nigra because of only minor differences with the nominate species from Japan. Only other valid species of Thysanocardia, T. catharinae, has been noted in the Sea of Japan. This species is known from Russian (Primorye) and Korean (near Tsushima) waters (see Murina, 1977). In contrast to T. nigra, tentacles of T. catharinae are without brown pigmentation, only with yellow tips and colorless in fixed specimens. This species is also distinguished with characteristic sinuous (zigzag) wrinkles on the trunk which tapers at the posterior end. Distribution. T. nigra is tropical-boreal species, widespread in the Pacific. In Russian waters T. nigra occurs from Tatarsky strait in the north to Possyet Bay (Peter the Great Bay) in the south. In the Sea of Japan, this species is also known from the Japanese and Korean coasts. T. nigra was also noted from the Okhotsk Sea and Pacific coast of Japan. In the West Pacific, the species is distributed to Indonesia. In the East Pacific, it is known from Washington to California. T. nigra is a shallow water species (0–200 m) and was never found in the deep sea. Family Phascolionidae E. Cutler and Gibbs, 1985 Genus Phascolion Theel, 1875 Subgenus Phascolion (Phascolion) Theel, 1875 Phascolion strombus (Montagu, 1804) (Fig. 3A–C) Material. Sea of Japan: near coast of Primorye Province, SoJaBio2010, sta# A2-8, trawl 44156.0262 N; 137113.2535 E, 582–570 m depth, gravel and muddy sediment, 8 specimens; sta# B7-8, trawl, 43113.6778 N; 135104.4447, 532 depth, gravel with muddy sediment, 7 specimens; 47140 N; 1391290 5 E in Tatarsky straight, 80 m depth, 3 specimens; near Moneron Island 461N; 1411110 E, 320 m; 5 specimens. Description. Trunk 15–20 mm long, 3–4 mm wide; introvert is about twice longer than trunk, with claw-like scattered hooks. Trunk with V-shape holdfast papillae on the posterior half. Gut with loose loops, anus anterior to nephridiopores. Retractor muscles originate about 80–90% of trunk length to the posterior end. Nephridium about 25% of trunk length. Discussion. The only other species of this genus in the Sea of Japan is P. lucifugax representing the subgenus Phascolion (Isomya). This species possesses dorsal and ventral retractor muscles of equal size while in the subgenus Phascolion ventral retractors are much thinner than dorsal ones. Cutler (1994) has distinguished two subspecies of P. strombus – P. strombus strombus (Montagu, 1804) and P. strombus cronullae (Edmonds, 1980). The last subspecies is known only from Australian waters and differs from the nominate form by papillae morphology and by proportion of retractor muscles. Distribution. P. strombus is a very widespread species which mainly inhabits various mollusk shells and polychaete tubes. Young specimens can be found in the mouth of mollusk shells occupied by adult worms (see Fig. 3B). In the Pacific, it is known from both northern and southern hemispheres, at the depth from 1 to 4000 m. In the West Pacific, P. strombus is found from the north part of Bering Sea to New Zealand. In the Sea of Japan, this species was previously known from Russian (Primorye coast and Tatarsky strait) and Japanese coasts (Honshu and Tsushima strait. Family Themistidae E. Cutler and Gibbs, 1985 Genus Themiste Gray, 1828 Subgenus Themiste (Themiste) Edmonds, 1980 Themiste blanda (Selenka and de Man, 1883) (Fig. 2D and E) Fig. 3. A–C, Phascolion strombus. (A) Lateral view, retracted introvert; (B) adult specimen with retracted introvert in gastropod shell (white arrow, sipunculan juvenile in the mouth of gastropod shell); (C) holdfast papilla. Scale bars: A–B, 3 mm; C, 50 mm. Material. Sea of Japan, Tsushima strait, Cheju Island 331120 38 N; 1261150 41 E, intertidal, muddy clay, 12 specimens. Description. Trunk pyriform, 3–6 mm long and 1.5–2.5 mm wide; introvert about one-third of half of trunk length, with scattered hooks covering anterior one-third of introvert; tentacular crown with four stems, with brown pigmented tentacules. Gut with 3–4 loops, anus slightly anterior to nephridiopores. Retractor muscles originate about 55% of trunk length to the posterior end. Nephridia smooth, about 30% of trunk length, free. Discussion. Two other valid representatives of the genus Themiste, T. minor and T. pyroides, have been noted from the Sea of Japan. T. hexadactyla is considered by Cutler (1994) as a junior synonym of T. pyroides (see below). The taxonomic status of A.S. Maiorova, A.V. Adrianov / Deep-Sea Research II 86–87 (2013) 140–147 T. maculosa described by Popkov (1993b) is also controversial (see below). In contrast to T. blanda, another representative of the subgenus Themiste (Themiste), T. pyroides, from the Sea of Japan possesses large tentacular crown with six main stems, while representatives from the North-East Pacific have only four main tentacles (see Cutler and Cutler, 1988; Cutler, 1994). T. minor represents another subgenus – Themiste (Lagenopsis) which is characterized by contractile vessel with short digitiform villi while all representatives of the subgenus Themiste possess contractile vessel with long thread-like processes. Distribution. T. blanda is intertidal and shallow water species. In the Sea of Japan, it was initially described from the Honshu and Hokkaido Islands. Murina (1977) also noted this species from the Okhotsk Sea and from the Russian coast of the Sea of Japan. Themiste hexadactyla (Sato, 1930) (Fig. 2A–C) Material. Sea of Japan, Peter the Great Bay, wide spread, clusters of mussels Crenomytilus grayanus, Modiolus kurilensis and Mytilus trossulus, 1–10 m depth, more than 150 specimens. Description. Trunk spindle-shaped, cylindrical or pyriform, 10–60 mm long and 2–15 mm wide; introvert equal or slightly longer than trunk, with black scattered hooks; tentacular crown always with six branched stems in all ages, branches and stems brownish at the aboral surface, separate tentacules with brown bands in the middle. Tentacular crown followed by smooth area (collar) with patchy distributed violet pigment (camouflage drawing); in turn, this collar followed by smooth area with solid violet or blue pigment. Trunk smooth, yellowish-brown or darkbrown, with minute papillae, usually with pointed posterior end. Retractor muscles originate about 60–80% of trunk length to the posterior end. Gut with 16–20 loops, with three fixing muscles; anus and nephridiopores at the same level; wing muscles well developed. Contractile vessel with numerous remarkably long thread-like tubules. Nephridia about 80% of trunk length. Discussion. The careful taxonomic revision appears to be necessary for this genus in the near future to distinguish species from the North-West and North-East Pacific. Representatives of the genus Themiste are very numerous in the Peter the Great Bay and, according to morphological and genetic investigations, correspond to a single population of one species (see Adrianov and Maiorova, 2010; Morozov and Adrianov, 2002; Schulze et al., 2012). Taking into account Cutler’s revision of the genus Themiste (see Cutler, 1994; Cutler and Cutler, 1988) and based on Cutler’s key to a species level (see Cutler, 1994), representatives of this species were identified as T. pyroides (see Morozov and Adrianov, 2002). T. pyroides was initially described from California (see Chamberlin, 1920) and later it was noted from Alaska to Baja California in the North-East Pacific and from Japan (Honshu and Hokkaido Islands) in the North-West Pacific (see Cutler, 1994). Themiste hexadactyla was initially described from Mitsu Bay of Honshu Island in Japan (see Sato, 1930) and later this species was also noted from Hokkaido (see Cutler and Cutler, 1981) and California (see Cutler and Cutler, 1981; Fisher, 1952; Murina, 1977). The principal feature distinguishing T. hexadactyla from T. blanda and T. pyroides is the presence of six main stems in the tentacular crown (see Cutler et al., 1984, p. 282, Fig. 6D; Murina, 1977, p. 245, Fig. 169a; Sato, 1930). Nevertheless, Cutler and Cutler (1988) introduce T. hexadactyla as a junior synonym to T. pyroides and strike off this species from the key for the genus Themiste (Cutler, 1994). In their new keys of the genus (see Cutler, 1994; Cutler and Cutler, 1988), he established for T. pyroides the presence of six main stems of the tentacular crown. 145 To describe Themiste from the Possiet Bay (south part of the Peter the Great Bay) with six tentacular stems, Popkov (1993b) has suggested a new species, Themiste maculosa. He distinguished this species from T. pyroides on the basis of the patchy (not solid) distribution of pigment on the collar and rounded shape of the posterior end of the trunk (see Cutler and Dean, 1997; Popkov, 1993). Despite the hard criticism of Popkov’s description, Cutler and Dean (1997) suggested confirming the validity of this new species thus creating two nominate species with six tentacular stems. Actually, according to our data, most specimens of T. pyroides from American coast possess on the anterior introvert a wide solid purple pigmented area without camouflage drawing, but all these specimens are characterized by four tentacular stems (own observations). In reality, all our specimens of Themiste from the Peter the Great Bay, previously identified as Themiste maculosa with six tentacular stems, possess patchy pigmented anteriormost introvert followed by solid pigmented smooth area that is well seen in living specimens with completely everted tentacular crown. Interestingly, that even in living specimens with completely everted tentacles, this patchy pigmented area can be partly or completely retracted into the anterior introvert thus giving appearance of only solidly colored area of the anteriormost introvert. Moreover, despite the presence of the bluntly pointed posterior end in most living specimens, we have observed some specimens of the same species with rounded posterior end of the trunk. Because of these facts, we do not recommend that only these variable characters be used to introduce a new species of Themiste. In this situation, we suggest to restoring validity of T. hexadactyle for West Pacific Themiste with six tentacular stems and considering T. maculosa as a junior synonym of this species. Nominate T. pyroides and T. blanda are distinguished from T. hexadactyla by the presence of only four main stems in the tentacular crown. According to Cutler (1994, p. 151), T. pyroides differs from T. blanda by much more voluminous tentacular crown and shorter terminal tentacules. Other East-Pacific species of the subgenus Themiste, T. dyscrita and T. hennahi, are well distinguished by the lack of hooks. The last valid representative of the subgenus Themiste, T. alutacea, is known only from Atlantic and characterized by unpigmented collar of the introvert. Distribution. T. hexadactyla is a shallow water East-Pacific species depth range; in the Peter the Great Bay common in between byssus threads and in slit-like spaces in clusters of bivalve mollusks. Class Phascolosomatidea Order Phascolosomatiformes Family Phascolosomatidae Genus Phascolosoma Leuckart, 1828 Subgenus Phascolosoma (Phascolosoma) Leuckart, 1828 Phascolosoma agassizii Keferstein, 1866 (Fig. 2H and I) Material. Sea of Japan, Peter the Great Bay, wide spread, clusters of mussels Crenomytilus grayanus, Modiolus kurilensis and Mytilus trossulus, in clumps of sea grasses Zostera marina and Phillospadix iwatensis on ground between stones, in muddy sand and mud substrates, 0–10 m depth, more than 200 specimens. Description. Trunk cylindrical, 4–40 mm long and 1–10 wide; introvert about equal to trunk length, with numerous hooks arranged in distinct (15–30) rings. Trunk yellow-brown, brown or gray, with large cupola-like dark brown or even black papillae over the whole body, concentrating at anterior and posterior trunk. Tentacles with dark blue or dark brown pigment at the bases. Hooks 45–65 mm tall and 45–70 mm wide, with smooth streak and indistinct triangle, without secondary tooth, with basal warts. Dorsal retractor muscles originate about 50% and ventral retractor muscles about 60–65% of 146 A.S. Maiorova, A.V. Adrianov / Deep-Sea Research II 86–87 (2013) 140–147 trunk length to the posterior end. Gut with 10–18 loops, with 1–3 fixing muscles, spindle muscle attached posteriorly. Anus at the same level or slightly anterior to nephridiopores. Wing muscles well developed. Nephridia about 30–50% of trunk length, about one-third attached to the body wall. Discussion. Four other species of the subgenus Phascolosoma (Phascolosoma), P. scolops, P. pacificum, P. albolineatum and P. granulatum, have been known from the Sea of Japan. In contrast to P. agassizii, P. scolops possesses introvert, usually shorter than trunk, hooks with distinct triangle, and band of red cone-shaped preanal papillae. P. pacificum is characterized by large (more than 100 mm tall) hooks with hump-like secondary tooth and distinct triangle, and possesses very long nephridia about equal to trunk length. P. albolineatum is well distinguished by hooks bent at an obtuse angle. P. granulatum is a species widespread in the Atlantic. P. granulatum, described from the North-West Pacific, is now recognized as P. agassizii. Cutler (1994) has distinguished two subspecies, P. agassizii agassizii (Keferstein, 1866) and P. agassizii kurilense (Sato, 1937). The first subspecies is known from North-East Pacific, from Alaska to Mexican coast, and from Japan. The second subspecies is known only from Kuril Islands and, in contrast to the nominate form, characterized by the presence of secondary lobe in nephridia. In Russian waters, P. agassizii has been noted from the Bering Sea (¼P. japonicum; Komandor Islands), Okhotsk Sea (¼P. kurilense, Kuril Islands; ¼P. japonicum, Sakhalin Island) and the Sea of Japan (¼P. japonicum, Tatarsky strait, Peter the Great Bay, North Korea; ¼P. granulatum, Tsushima Strait). Distribution. Common on both sides of the North Pacific from Mexico to Alaska on the eastern side and from Japan to Russia in the west. Scattered records from Indian Ocean waters, as well as south and west Africa in the Atlantic. 4. Key to sipunculan species In addition to our species, the following identification key also includes all other valid species, lacking in Russian waters, but noted from the Japanese and Korean coasts of the Sea of Japan (see Murina, 1977; Cutler et al., 1984, 1994; List of Animals in Korea, 1998; own collections). 1. Tentacular crown consists of oral crown with peripheral tentacles arranged around mouth and on lateral sides of head; tentacles of dorsal arc present or absent; hooks, if present, usually scattered—class Sipunculidea (2) Tentacular crown consists of dorsal arc encircling nuchal organ; oral crown with peripheral tentacles absent; hooks, if present, in distinct rings—class Phascolosomatidea (24) 2. Longitudinal muscles of body wall gathered into separate or anastomosing bands—order Sipunculiformes, fam. Sipunculidae (3) Longitidinal muscles of body wall in uniform continuous layer—order Golfingiiformes (6) 3. Body wall circular and longitudinal bands anastomosing, spindle muscle attached to the posterior trunk—genus Siphonosoma (4) Body wall circular and longitudinal bands not anastomosing, spindle muscle not attached to the posterior trunk—genus Sipunculus (5) 4. Dorsal retractor muscles originate anterior to ventral pair—S. mourense Retractor muscles originate at the same level—S. cumanense 5. 20–24 longitudinal muscle bands; nephridia free—S. norvegicus 24–34 longitudinal muscle bands; nephridia 10–40% attached—S. nudus 6. Tentacular crown with four or six branched stems—fam. Themistidae, genus Themiste (21) Tentacles unbranched—(7) 7. Two nephridia present—fam. Golfingiidae (8) One nephridium present—fam. Phascolionidae (18) 8. Contractile vessel simple, without villi—(9) Contractile vessel with numerous digitiform villi genus Thysanocardia—(17) 9. Four retractor muscles of introvert genus Golfingia—(10) Two retractor muscles of introvert genus Nephasoma—(13) 10. Posterior trunk with prominent caudal appendage— G. muricaudata Posterior trunk without caudal appendage—(11) 11. Introvert hooks gathered in distinct rings—G. elongata Introvert hooks scattered, if present—(12) 12. Introvert of mature specimens with hooks—G. vulgaris Introvert of mature specimens without hooks—G. margaritacea 13. Tentacles reduced to lobes, trunk with pigmented papillae, trunk length usually exceeds width by more than eight times—N. diaphanes Tentacles well developed, trunk with unpigmented papillae, trunk length usually not exceeds width more than eight times—(14) 14. Hooks present—(15) Hooks absent—(16) 15. Body thread-like; hooks small (20–25 mm), scattered or arranged in rings—N. capilleforme Body not thread-like; hooks (50–150 mm) unique spirally arranged—N. abyssorum 16. Trunk nonpapillated, with transverse grooves; retractor muscles originate in the middle third of the trunk—N. eremite Trunk well papillated, without transverse grooves; retractor muscles originate in the posterior quarter of the trunk— N. wodjanizkii 17. Tentacular crown with dark pigment—T. nigra Tentacular crown without pigment—T. catharinae 18. Retractor muscles fused into a single column; without holdfast papillae—genus Phascolion, subgenus Phascolion (Lesenka), P. rectum Separate dorsal and ventral retractor muscles, with holdfast papillae—(19) 19. Dorsal and ventral retractors of equal size—genus Phascolion, subgenus Phascolion (Isomya) (20) Ventral retractor muscle much thinner than dorsal—genus Phascolion, subgenus Phascolion (Phascolion), P. strombus 20. Hooks blunt, strongly curved; both retractor muscles originate about 15% of distance from the posterior end of trunk, ventral origin slightly posterior to dorsal one— P. lucifugax Hooks blunt, spine-like or slightly bent; two retractor muscles originate at the same level at the posterior end of trunk— P. hedraeum 21. Contractile vessel with long thread-like tubular extensions— subgenus Themiste (Themiste) (22) Contractile vessel with numerous digitiform villi—subgenus Themiste (Lagenopsis), T. minor 22. Tentacular crown with six distinct stems, patchy pigmented collar followed by solid pigmented area—T. hexadactyla Tentacular crown with four distinct stems, collar entirely (solid) pigmented or non-pigmented—(23) 23. Anteriormost introvert with wide solid pigmented area, purple or violet—T. pyroides Anteriormost introvert non-pigmented or only slightly colored apically with narrow band of brown-violet pigment—T. blanda A.S. Maiorova, A.V. Adrianov / Deep-Sea Research II 86–87 (2013) 140–147 24. Anal shield always present, composed of numerous horny plates, longitudinal muscles in continuous layer order Aspidosiphoniformes, fam. Aspidosiphonidae, genus Aspidosiphon, subgenus Aspidosiphon (Aspidosiphon)—(25) Anal shield always absent order Phascolosomatiformes, fam. Phascolosomatidae—(26) 25. Anal shield with extensive grooves and furrows, individual units form into longitudinal ridges—A. muelleri Anal shield without extensive grooves and furrows— A. misakiensis 26. Introvert several times (6 –1 4) longer than trunk; nephridia usually bilobed—genus Apionsoma (27) Introvert usually not more than two times longer than trunk; nephridia unilobed—genus Phascolosoma (28) 27. Tentacles present; hooks with accessory basal spinelets— A. misakianum Tentacles underdeveloped; hooks, if present, without spinelets—A. trichocephalus 28. Hooks remarkably bent at an obtuse angle—P. albolineatum Hooks not bent at an obtuse angle—(29) 29. Hooks with hump-like secondary tooth and distinct triangle; nephridia about equal to trunk length—P. pacificum Hooks without hump-like secondary tooth; nephridia much shorter than trunk length—(30) 30. Introvert usually shorter than trunk; hooks with distinct triangle—P. scolops Introvert usually equal or longer than trunk; hooks with indistinct triangle—P. agassizii Acknowledgments The authors gratefully acknowledge the support of the Russian Foundation of Fundamental Research (grants 11-04-98546-r; 12-04-00263-a), the Far East Branch of the Russian Academy of Sciences (grants 12-I-P30-07, 12-I-0-06-005, 12-I-0-06-004, 12-I-0-06-002, 12-HHC-007), APN Project ARCP2011-10CMY and Ministry of Education and Science of Russian Federation (grant 11.G34.31.0010) during the course of this study. References Adrianov, A.V., Maiorova, A.S., 2010. Reproduction and development of common species of peanut worms (Sipuncula) from the Sea of Japan. Russ. J. Mar. Biol. 36 (1), 1–15. 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