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In this study, we had investigated the benthic deep-sea fauna using the dredge and beam trawl in the region from the continental shelf to the continental slope of the Sea of Kumano, Mie Prefecture, during the No．1722 research voyage of the training ship Seisui-Maru of Mie University. The survey was carried out at 16 stations covering a depth range of 113－1059 m. The results of the survey, 14 phyla had been confirmed. Arthropod, echinoderm, annelid and molluscan macrobenthos were collected from all of the stations. The phylum number of each station was in the range from 4 to 11. The largest number of phyla had been confi rmed at St．10D of boulders bottom（768－800 m depth）. Meiobenthos confi rmed in our sample were kinorhynchs, nematodes, tardigrades, loriciferans and small arthropods such as tanaidaceans, copepods and cumaceans. In addition to free-living species, parasitic copepods, isopods, platyhelminthes, acanthocephalans and nematodes had been found in fish, crustaceans and polychaetes.

Laubieriopsis soyoae sp. nov. Jimi (New Japanese name: Soyo-sube-imomushi-gokai) (Figure 3) Type material. Holotype (NSMT-Pol H-825): 13 mm long, 0.3 mm wide, and 50 paratypes (NSMT-Pol P-826): 10–15 mm long (n=50), 0.3 mm wide (n=50), 24 chaetigers (n=50); free living, sand sediment, off Mikura Island, Tokyo, Japan, by a beam trawl (Fig. 1C, D), at a depth of 1114–1171 m, collected by NJ. One paratypes (ICHUM- 6161): fragmented, used for SEM observation. Description. Holotype (NSMT-Pol H-825) body complete, 13 mm long, 0.3 mm wide, with 24 chaetigers, cylindrical, not swollen in middle region (Fig. 3A, 3B). Living colour transparent (Fig. 3A). Prostomium partially exposed, fused to peristomium (Fig. 3C). Mouth opening ventral, T-shaped. Integument iridescent, smooth, not papillated. Genital papillae unpaired (Fig. 4B), low oval projections, present on right side of posterior margin of chaetiger 8. Anterior segments without annulations. Parapodia biramous, lateral, projected along chaetigers 1–3, inconspicuous in subsequent body region. Interramal papillae, non-pedunculate. All chaetae brownish. Chaetigers 1–3 (Fig. 3D), with chaetae directed anteriorly: notopodia and neuropodia with four curved acicular chaetae per bundle. In chaetiger 4 and following chaetigers (Fig. 3E), chaetae directed laterally or posteriorly: notopodia and neuropodia with short acicular and long capillary chaetae. Acicular chaetae smooth, tips entire, without denticles in tips. Capillary chaetae smooth. Pygidium without anal cirri and papillae (Fig. 3F), withdrawn into posterior segment; chaetae shape same in median chaetigers; anus terminal. Etymology. The species is named after the R/V Soyo -maru. The type specimens from off Mikura Island were collected by the gear present on the ship. The specific name is a noun in the genitive case. Distribution. Only known from the type locality, off Mikura Island, Tokyo, Japan, at a depth of 1114–1171 m. Remarks. Laubieriopsis soyoae sp. nov. can be distinguished from the other members of the genus by the following features: i) 24 chaetigers, ii) chaetigers 1–3 without annulations, and iii) acicular chaetae with tips entire, without denticles. This species most resembles L. arenicola (Riser, 1987) by having acicular chaetae with tapered smooth tips and body with more than 21 chaetigers; however, annulations in chaetigers 1–3 are only present in L. arenicola (Riser 1987; Salazar-Vallejo et al. 2019), and the body has 25 chaetigers. Laubieriopsis hartmanae has been known from Japan (Levenstein 1970; Imajima 2009; Salazar-Vallejo et al. 2019). The new species can be discriminated from L. hartmanae by having 24 chaetigers, instead of only 16, and genital papillae on chaetiger 8, instead of being present between chaetigers 6–7.

Genus Fauveliopsis McIntosh, 1922 (New Japanese name: yadokari-imomushi-gokai-zoku), Published as part of Jimi, Naoto, Fujimoto, Shinta, Ogawa, Akito, Fujita, Yoshihisa, Shigenobu, Yuya & Imura, Satoshi, 2020, Two new species of Fauveliopsidae (Annelida, Sedentaria) from submarine caves and deep sea, Japan, pp. 385-391 in Zootaxa 4878 (2) on page 387, DOI: 10.11646/zootaxa.4878.2.11, http://zenodo.org/record/4424921, {"references":["McIntosh, W. C. (1922) Notes from the Gatty Marine Laboratory, St. Andrews, 44: 1. On new and rare Polychaeta, Gephyrea, etc., from various regions; 2. Recent additions to the British marine Polychaeta (continued). Annals and Magazine of Natural History, Series 9, 9, 1 - 30. https: // doi. org / 10.1080 / 00222932208632638"]}

Fauveliopsis antri sp. nov. Jimi (New Japanese name: doukutsu-yadokari-imomushi-gokai) (Figure 2) Type material. Holotype (NSMT-Pol H-822): 1.2 mm long, 0.2 mm wide; 5 paratypes (NSMT-Pol P-823): 1.0��� 1.2 mm long (n=5), 0.2 mm wide (n=5), 32 chaetigers (n=5); all specimens were extracted from small molluscan shells by using forceps, silt sediment, Daidokutsu cave, off Ie Island, Okinawa, Japan, at a depth of 26 m; collected by NJ and SF. 15 additional paratypes (NSMT-Pol P-824): 1.0��� 1.5 mm long (n=15), 0.2 mm wide (n=15), 32 chaetigers (n=15); obtained from small molluscan shells by washing with fresh water, silt sediment, Shodokutsu cave (���unnamed cave���), at a depth of 18 m (near to Daidokutsu cave). Description. Holotype (NSMT-Pol H-822) body complete, 1.2 mm long, 0.2 mm wide, 32 chaetigers, anterior and median parts cylindrical, posteriorly swollen, divided into three different regions by body colour and chaetal features, borderline of median and posterior regions indistinct. (Fig. 2A). Body colour whitish in anterior and medial parts, posteriorly transparent, without pigmentation, posterior gut yellowish (Fig. 2A). Prostomium partially exposed, fused to peristomium, with a wide, densely ciliated transverse band (Fig. 2B, C). Nuchal organ present on dorsal side of peristomium. Mouth opening ventral, T-shaped opening. Integument opaque, rugose, papillated on dorsal and ventral surface. Body papillae 2���3 ��m in length (Figs. 2F, 4A), digitate, sparsely present on dorsal and ventral sides. Posterior region transparent, rugose; last chaetiger with a transverse band of large, rounded papillae (Fig. 2F). Genital papillae not seen. Parapodia lateral, inconspicuous in all body regions, orange colour in life. Internal papilla pedunculate, on middle of parapodia, along anterior region; smaller and closer to notopodia in median and posterior regions. Chaetigers 1���3 with chaetae directed anteriorly (Fig. 2C): notopodia with straight, thick capillary chaetae (upper) and curved acicular (lower) chaetae; neuropodia with two types: acicular, stout sigmoid acicular (upper) and short sigmoid acicular chaetae. Chaetigers 4 and following with chaetae directed laterally or posteriorly: notopodia and neuropodia with long curved acicular and long capillary chaetae (Fig. 2D). All chaetigers with two notochaetae and two neurochaetae (Fig. 2E). Pygidium without anal cirri; anus terminal with some blunt, globular or digitate papillae surrounding anus (Fig. 2F). Etymology. Latin ��� antrum ��� means ���cave��� in English. The name is derived from the habitat of this new species. The specific name is a noun in the genitive case. Distribution. Only known from submarine caves, off Ie Island, Okinawa, Japan, at a depth of 18��� 26 m. Remarks. Fauveliopsis antri sp. nov. can be distinguished from the other members of the genus by the following features: i) 32 chaetigers, ii) chaetigers 1���3 with stout hooks, iii) minute body (about 1 mm in length), iv) all parapodia with same number of chaetae (2 notochaetae; 2 neurochaetae), and v) presence of dorsal and ventral papillae. The most similar species is F. jameoaquensis N����ez in N����ez, Oca��a & Brito, 1997, also described from caves, by having two chaetae per rami along its minute body. However, the main difference is that F. antri sp. nov., has curved acicular chaetae in chaetigers 1���3, while F. jameoaquensis has straight acicular chaetae (N����ez et al. 1997). The species also resembles F. brevipodus Hartman, 1971 and Fauveliopsis sp. A Wolf, 1984 by having two chaetae per rami along its body, body with more than 15 chaetigers, and papillae on pygidium. The new species differs from F. brevipodus in the chaetiger number and papillae of pygidium: the new species has 32 chaetigers and multi-papillated pygidium, whereas F. brevipodus has 27���29 chaetigers and smooth pygidium. Fauveliopsis sp. A also has multi-papillated pygidium but the new species has minute body (about 3.0 mm long in Fauveliopsis sp. A) with 32 chaetigers (18���30 chaetigers in Fauveliopsis sp. A). Some specimens of Fauveliopsis levensteinae SalazarVallejo, 2019 also have 32 chaetigers but have many chaetae on posterior chaetigers. The new species can be distinguished from two other species recorded from Japan, F. glabra and F. challengeriae (Levenstein 1970; Imajima 2009), by having a minute body and same number of chaetae in all parapodia. Fauveliopsis glabra and F. challengeriae have large bodies (about 5 mm in body length) and the number of chaetae varies depending on the body area., Published as part of Jimi, Naoto, Fujimoto, Shinta, Ogawa, Akito, Fujita, Yoshihisa, Shigenobu, Yuya & Imura, Satoshi, 2020, Two new species of Fauveliopsidae (Annelida, Sedentaria) from submarine caves and deep sea, Japan, pp. 385-391 in Zootaxa 4878 (2) on pages 387-388, DOI: 10.11646/zootaxa.4878.2.11, http://zenodo.org/record/4424921, {"references":["Nunez, J., Ocana, O. & Brito, M. C. (1997) Two new species (Polychaeta: Fauveliopsidae and Nerillidae) and other polychaetes from the marine lagoon cave of Jameos del Agua, Lanzarote (Canary Islands). Bulletin of Marine Science, 60, 252 - 260. [https: // www. ingentaconnect. com / contentone / umrsmas / bullmar / 1997 / 00000060 / 00000002 / art 00007]","Hartman, O. (1971) Abyssal polychaetous annelids from the Mozambique Basin off Southeast Africa, with a compendium of abyssal polychaetous annelids from world-wide areas. Journal of the Fisheries Research Board of Canada, 28, 1407 - 1428. https: // doi. org / 10.1139 / f 71 - 219","Wolf, P. S. (1984) Family Fauveliopsidae Hartman, 1971. In: Uebelacker, J. M. & Johnson, P. G. (Eds.), Taxonomic Guide to the Polychaete of the Northern Gulf of Mexico. 7 Vols. Barry A. Vittor & Associates, Mobile, Alabama, pp. 1 - 37.","Levenstein, R. Y. (1970) Novi i redkie vidi mnogoshchetinkovikh chervei glubokovodnogo roda Fauveliopsis i osobennosti ikh rasprostraneniya. Trudy Instituta Okeanologii, Akademiya Nauk SSSR, 88, 227 - 235.","Imajima, M. (2009) Deep-sea benthic polychaetes off Pacific coast of the northern Honshu, Japan. Monographs, National Museum of Nature and Science, Tokyo, 39, 39 - 192. [https: // www. kahaku. go. jp / research / publication / monograph / download / 39 / p 039 - 192. pdf]"]}

Family Fauveliopsidae Hartman, 1971 (New Japanese name: imomushi-gokai-ka), Published as part of Jimi, Naoto, Fujimoto, Shinta, Ogawa, Akito, Fujita, Yoshihisa, Shigenobu, Yuya & Imura, Satoshi, 2020, Two new species of Fauveliopsidae (Annelida, Sedentaria) from submarine caves and deep sea, Japan, pp. 385-391 in Zootaxa 4878 (2) on page 387, DOI: 10.11646/zootaxa.4878.2.11, http://zenodo.org/record/4424921, {"references":["Hartman, O. (1971) Abyssal polychaetous annelids from the Mozambique Basin off Southeast Africa, with a compendium of abyssal polychaetous annelids from world-wide areas. Journal of the Fisheries Research Board of Canada, 28, 1407 - 1428. https: // doi. org / 10.1139 / f 71 - 219"]}

Genus Laubieriopsis Petersen, 2000 (New Japanese name: sube-imomushi-gokai-zoku), Published as part of Jimi, Naoto, Fujimoto, Shinta, Ogawa, Akito, Fujita, Yoshihisa, Shigenobu, Yuya & Imura, Satoshi, 2020, Two new species of Fauveliopsidae (Annelida, Sedentaria) from submarine caves and deep sea, Japan, pp. 385-391 in Zootaxa 4878 (2) on page 389, DOI: 10.11646/zootaxa.4878.2.11, http://zenodo.org/record/4424921, {"references":["Petersen, M. E. (2000) A new genus of Fauveliopsidae (Annelida Polychaeta), with a review of its species and redescription of some described taxa. Bulletin of Marine Science, 67, 491 - 515. [https: // www. ingentaconnect. com / content / umrsmas / bullmar / 2000 / 00000067 / 00000001 / art 00041? crawler = true]"]}

We herein present the results of a survey which assessed the benthic fauna from subtidal to continental shelf depth in the Shirahama area from 2012 to 2016. Our research resulted in the identification of 132 species from 75 families in seven phyla, Cnidaria, Annelida, Tardigrada, Arthropoda, Mollusca, Echinodermata and Chordata. This includes 24 newly recorded species to Shirahama. Two species were also new records for Japanese waters. Furthermore, six undescribed species and five potentially undescribed species were recorded. We provide a selection of relevant photos for future taxonomic studies and monitoring of environmental changes., Issued online on Dec. 7, 2016; paper version on Aug. 30, 2018

Angursa antarctica Villora-Moreno, 1998 Emended diagnosis Angursa with bulbous primary clavae shorter than lateral cirri; pedestals of primary clavae and lateral cirri absent; morphology of secondary clavae unknown; flat tertiary clavae around external cirri; tapering cirri E with annulated scapi situated on short cirrophores; leg I sensory organs present; legs II and III sensory organs absent; leg IV sensory organs each as papilla enveloped in hemispherical cuticular sheath with short, apical spine; weak anal papillae present. Remarks We amend the diagnosis of A. antarctica to include the information on the anal papillae noted in the original description (Villora-Moreno 1998) as well as the common pedestals of the primary clavae and the lateral cirri that Villora-Moreno (1998) considered as a distinguishing character but did not include in the diagnosis of A. antarctica. Furthermore, Villora-Moreno (1998) reported the presence of the secondary clavae around the external cirri. As the external cirri are positioned ventrally, we believe that the clavae Villora-Moreno (1998) observed should be referred to as tertiary clavae which are always positioned ventrally in the genus., Published as part of Fujimoto, Shinta & Hansen, Jesper Guldberg, 2019, Revision of Angursa (Arthrotardigrada: Styraconyxidae) with the description of a new species from Japan, pp. 1-19 in European Journal of Taxonomy 510 on page 9, DOI: 10.5852/ejt.2019.510, http://zenodo.org/record/2616378, {"references":["Villora-Moreno S. 1998. Deep-sea Tardigrada from South Shetland Islands (Antarctica) with description of Angursa antarctica sp. nov. (Arthrotardigrada, Halechiniscidae). Polar Biology 19: 336 - 341. https: // doi. org / 10.1007 / s 003000050255"]}

Angursa capsula Bussau, 1992 Fig. 5 Material examined All three specimens were collected from a manganese nodule area off Peru, but it is not clear whether the paratypes observed in this study were collected with the holotype or at a nearby site on a different day since the museum catalogue numbers were not mentioned in the original description (Bussau 1992). Holotype PACIFIC OCEAN ��� adult ♀; tropical SE Pacific, Peru Basin, collected from a manganese nodule area off Peru, 4140���4170 m deep; 07��03���84��� S, 88��28���15��� W; 17 Feb. 1989; SMF 28. Paratypes PACIFIC OCEAN ��� 2 specimens collected from a manganese nodule area off Peru; SMF 29, SMF 30. Remarks The re-examination of the type material revealed the presence of the paired globular body fringing the mouth in the holotype (Fig. 5A), the shallow bases of the primary clavae and the lateral cirri which we do not regard as pedestals, and the morphology of the leg IV sensory organs (Fig. 5B) to be not significantly different from those of A. abyssalis and A. clavifera (Renaud-Mornant 1981; Noda 1985). The third point is especially important as its ���capsule-shaped��� leg IV sensory organ has been the primary diagnostic character for this species (Bussau 1992). The seconday and tertiary clavae were now very difficult to observe in the holotype. They were observed in SMF 29, but their complete contours were difficult to recognize., Published as part of Fujimoto, Shinta & Hansen, Jesper Guldberg, 2019, Revision of Angursa (Arthrotardigrada: Styraconyxidae) with the description of a new species from Japan, pp. 1-19 in European Journal of Taxonomy 510 on page 8, DOI: 10.5852/ejt.2019.510, http://zenodo.org/record/2616378, {"references":["Bussau C. 1992. New deep-sea Tardigrada (Arthrotardigrada, Halechiniscidae) from manganese nodule area of the eastern South Pacific. Zoologica Scripta 21: 79 - 91. https: // doi. org / 10.1111 / j. 1463 - 6409.1992. tb 00311. x","Renaud-Mornant J. 1981. Deux nouveaux Angursa Pollock, 1979, du domaine abyssal (Tardigrada, Arthrotardigrada). Tethys 10: 161 - 164.","Noda H. 1985. Description of a new subspecies of Angursa bicuspis Pollock (Heterotardigrada, Halechiniscidae) from Tanabe Bay, Japan. Publications of the Seto Marine Biological Laboratory 30: 269 - 276. Available from https: // hdl. handle. net / 2433 / 176110 [accessed 4 Feb. 2019]."]}

Angursa lanceolata Renaud-Mornant, 1981 Fig. 4 Emended diagnosis Angursa with bulbous primary clavae terminating in apices shorter than lateral cirri; pedestals of primary clavae and lateral cirri present; flat secondary clavae present; morphology of tertiary clavae unknown; lance-like cirri E; Leg I sensory organs present; legs II and III sensory organs absent; leg IV sensory organs each as elongate papilla with short apical spine; anal papillae absent. Material examined Paratype ATLANTIC OCEAN ��� 1 adult; mounted with the dried out holotype; Dinet leg.; AE128. Remarks The primary clavae were described as leaf-shaped by Renaud-Mornant (1981), but the primary clavae of the paratype we observed were bulbous papillae (8 ��m long, 6 ��m in diameter) with blunt apices (the apex were only recognised in one of the clavae) (Fig. 4A). Their neighbouring sensory organs, the lateral cirri (17 ��m) (Fig. 4A), were apparently longer than reported in the original description (11 ��m) (Renaud-Mornant 1981). As it was drawn in the original description, the pedestals that support the primary clavae and the lateral cirri were apparent (Fig. 4A). We recognised the flat secondary clavae (Fig. 4A) which had already been mentioned as slightly hollow formations by Renaud-Mornant (1981), but their contours were difficult to observe. The presence and morphology of the tertiary clavae could not be revealed in this study. The characteristic cirri E (Fig. 4B) were confirmed, but their proportion seem slightly different that the drawing provided by Renaud-Mornant (1981). We recognised the short, apical spines of the leg IV sensory organs (Fig. 4B) which were overlooked in the original description (Renaud-Mornant 1981). Other measurements of the paratype are as follows: body length. 136 ��m; median cirrus, 3 ��m; internal cirrus, 6 ��m; external cirrus, 6 ��m; cirrus E, 11 ��m; leg I sensory organ, 6 ��m; leg IV sensory organ, 5 ��m. Angursa lanceolata closely resembles A. lingua by the bulbous primary clavae shorter than the lateral cirri, the apparent pedestals that support these two sensory organs, and the elongate leg IV sensory organs with apical spines (Renaud-Mornant 1981; Bussau 1992). However, these two species are easily distinguished from each other by the longer cephalic cirri and cirri E in A. lingua (median cirrus, 10 ��m; internal cirrus, 9 ��m; external cirrus 13 ��m; lateral cirrus 28 ��m; cirrus E, 20 ��m (body length, 145 ��m)) and by the lance-shaped cirri E in A. lanceolata versus tongue-shaped cirri E in A. lingua (Renaud- Mornant 1981; Bussau 1992)., Published as part of Fujimoto, Shinta & Hansen, Jesper Guldberg, 2019, Revision of Angursa (Arthrotardigrada: Styraconyxidae) with the description of a new species from Japan, pp. 1-19 in European Journal of Taxonomy 510 on pages 7-8, DOI: 10.5852/ejt.2019.510, http://zenodo.org/record/2616378, {"references":["Renaud-Mornant J. 1981. Deux nouveaux Angursa Pollock, 1979, du domaine abyssal (Tardigrada, Arthrotardigrada). Tethys 10: 161 - 164.","Bussau C. 1992. New deep-sea Tardigrada (Arthrotardigrada, Halechiniscidae) from manganese nodule area of the eastern South Pacific. Zoologica Scripta 21: 79 - 91. https: // doi. org / 10.1111 / j. 1463 - 6409.1992. tb 00311. x"]}

Identification key to Angursa species 1. Club-shaped primary clavae longer than lateral cirri........................................ 2 (A. bicuspis group) ��� Bulbous primary clavae shorter than lateral cirri...........................................6 (A. lanceolata group) 2. Well developed anal papillae..................................................................... A. bicuspis Pollock, 1979 ��� Anal papillae absent........................................................................................................................... 3 3. Legs II and III sensory organs present; spiral seminal receptacle ducts........ A. clavifera Noda, 1985 ��� Leg II and III sensory organs absent.................................................................................................. 4 4. Three-lobed tertiary clavae; D-shaped secondary clavae; seminal receptacles present........................................................................................................................................... A. seisuimaruae sp. nov. ��� Tertiary clavae with no indentation or only slightly indented around external cirri.......................... 5 5. Secondary clavae situated close together (ca 3 ��m); short (10 ��m) cirrus E.......................................................................................................................................... A. abyssalis Renaud-Mornant, 1981 ��� Secondary clavae situated apart (ca 9 ��m); long (19 ��m) cirrus E............... A. capsula Bussau, 1992 6. Leg IV sensory organs as papillae enveloped in hemispherical sheaths; cirri E with annulated scapi............................................................................................ A. antarctica Villora-Moreno, 1998 ��� Leg IV sensory organs as elongate papillae....................................................................................... 7 7. Cirri E lance-shaped................................................................ A. lanceolata Renaud-Mornant, 1981 ��� Cirri E tongue-shaped..................................................................................... A. lingua Bussau, 1992, Published as part of Fujimoto, Shinta & Hansen, Jesper Guldberg, 2019, Revision of Angursa (Arthrotardigrada: Styraconyxidae) with the description of a new species from Japan, pp. 1-19 in European Journal of Taxonomy 510 on page 13, DOI: 10.5852/ejt.2019.510, http://zenodo.org/record/2616378, {"references":["Pollock L. W. 1979. Angursa bicuspis n. g., n. sp., a new marine arthrotardigrade from the western North Atlantic. Transactions of the American Microscopical Society 98: 558 - 562. https: // doi. org / 10.2307 / 3225907","Noda H. 1985. Description of a new subspecies of Angursa bicuspis Pollock (Heterotardigrada, Halechiniscidae) from Tanabe Bay, Japan. Publications of the Seto Marine Biological Laboratory 30: 269 - 276. Available from https: // hdl. handle. net / 2433 / 176110 [accessed 4 Feb. 2019].","Renaud-Mornant J. 1981. Deux nouveaux Angursa Pollock, 1979, du domaine abyssal (Tardigrada, Arthrotardigrada). Tethys 10: 161 - 164.","Bussau C. 1992. New deep-sea Tardigrada (Arthrotardigrada, Halechiniscidae) from manganese nodule area of the eastern South Pacific. Zoologica Scripta 21: 79 - 91. https: // doi. org / 10.1111 / j. 1463 - 6409.1992. tb 00311. x","Villora-Moreno S. 1998. Deep-sea Tardigrada from South Shetland Islands (Antarctica) with description of Angursa antarctica sp. nov. (Arthrotardigrada, Halechiniscidae). Polar Biology 19: 336 - 341. https: // doi. org / 10.1007 / s 003000050255"]}

Genus Angursa Pollock, 1979 Emended diagnosis Styraconyxidae with slender body; median cirrus present; papillate primary clavae; flat secondary and tertiary clavae; leg I sensory organs present; legs II and III sensory organs occasionally present; leg IV sensory organs papillate; cirri E near caudal margin of body; each digit terminates in claw with two divergent points; peduncles present at base of external digits; seminal receptacles present; terminal positioned anus occasionally with anal papillae., Published as part of Fujimoto, Shinta & Hansen, Jesper Guldberg, 2019, Revision of Angursa (Arthrotardigrada: Styraconyxidae) with the description of a new species from Japan, pp. 1-19 in European Journal of Taxonomy 510 on page 3, DOI: 10.5852/ejt.2019.510, http://zenodo.org/record/2616378, {"references":["Pollock L. W. 1979. Angursa bicuspis n. g., n. sp., a new marine arthrotardigrade from the western North Atlantic. Transactions of the American Microscopical Society 98: 558 - 562. https: // doi. org / 10.2307 / 3225907"]}

Angursa lingua Bussau, 1992 Material examined Out of the 19 specimens designated as type material (Bussau 1992), paratypes SMF 34, SMF 38, SMF 39, SMF 42, and SMF 43 were observed. However, the mounting media were dried out in SMF 34, SMF 38, SMF 39 and SMF 42. Due to the same problem discussed for A. capsula, the collection locality of each specimen is not clear. Remarks No new information was obtained from the detailed observation of SMF 43., Published as part of Fujimoto, Shinta & Hansen, Jesper Guldberg, 2019, Revision of Angursa (Arthrotardigrada: Styraconyxidae) with the description of a new species from Japan, pp. 1-19 in European Journal of Taxonomy 510 on page 9, DOI: 10.5852/ejt.2019.510, http://zenodo.org/record/2616378, {"references":["Bussau C. 1992. New deep-sea Tardigrada (Arthrotardigrada, Halechiniscidae) from manganese nodule area of the eastern South Pacific. Zoologica Scripta 21: 79 - 91. https: // doi. org / 10.1111 / j. 1463 - 6409.1992. tb 00311. x"]}

Angursa abyssalis Renaud-Mornant, 1981 Figs 2���3 Emended diagnosis Angursa with club-shaped primary clavae longer than lateral cirri; pedestals of primary clavae and lateral cirri absent; flat, distorted-oval secondary clavae; flat, oval tertiary clavae slightly indented near external cirri; tapering cirri E with basal scapi; leg I sensory organs present; legs II and III sensory organs absent; leg IV sensory organs each as papilla enveloped in hemispherical cuticular sheath with short, apical spine; anal papillae absent. Material examined Holotype ATLANTIC OCEAN ��� adult ♀; Southeast Atlantic, collected from off Angola; 2063 m deep site; 12��03��� 3 S, 12��20��� 5 E; Dinet leg.; MNHN AF 05/77 Ma. Additional specimens ATLANTIC OCEAN ��� 1 adult ♀; North Atlantic; 2205 m deep; Dinet & Vivier leg.; MNHN AP 345/560Ma. MEDITERRANEAN SEA ��� 1 adult; Cassidaigne Canyon; 460 m deep; Vitiello & Vivier leg.; MNHN AE 120 /78Ma. Remarks There were two primary purposes for the re-examination of this taxon. One purpose was to identify the secondary and tertiary clavae, which were not reported in the original description (Renaud-Mornant 1981). The second purpose was to accurately understand the morphology of the leg IV sensory organs, which was described as hemispherical papillae in the text, but drawn as elongate papillae in the figure of the original description (Renaud-Mornant 1981). Unfortunately, the secondary and tertiary clavae were not recognised in the holotype, MNHN AF 05/77Ma. However, they were evident in the additional specimen, MNHN AP 345/560Ma. In this specimen, the paired secondary clavae were positioned in the dorsal area between the median cirrus and the primary clavae and each was in the form of a flat, distorted-oval lobe (the complete contour was not recognisable) (Figs 2, 3A). The paired tertiary clavae were position ventrally, immediately posterior to the external cirri and each was in the form of a flat, transversally-elongated lobe with a slight indentation where the external cirrus arises (Figs 2, 3B). In the other additional specimen, MNHN AE 120/78Ma, the secondary clavae were not recognised, but a paired structure indicative of clavae (tertiary clavae?) was observed in the latero-ventral position of the cephalic region (Fig. 3C). Further, a paired globular body fringing the mouth was recognised (Fig. 3D). The leg IV sensory organs were recognised in all three specimens.The holotype and MNHN AP345/560Ma had leg IV sensory organs consisting of papillae enveloped in hemispherical cuticular sheaths with short, apical spines (Fig. 3E) similar to that described in the text of the original description (Renaud-Mornant 1981). On the other hand, the leg IV sensory organs of MNHN AE120/78Ma lacked the enveloping cuticular sheaths and the papillae appeared slightly elongate and also constricted at the base (Fig. 3F) as in fig. 1A of Renaud-Mornant (1981). Further, it is clear that fig. 1C of Renaud-Mornant (1981) is based on MNHN AE120/78Ma (Fig. 3F). Other morphological characters of the three specimens confirm the original description (Renaud-Mornant 1981) except that a scapus was observed in one of the external cirri of the MNHN AP345/560Ma (Fig. 2). Here we provide measurements of the holotype missing in the original description: internal cirrus, 3 ��m; external cirrus, 6 ��m; primary clava, 10 ��m; leg IV sensory organ, 5 ��m long and 3 ��m in diameter. Our re-examination reveals that the term ���balloon-shaped��� used by Bussau (1992) and Villora-Moreno (1998) (probably based on the figure in the original description of Renaud-Mornant 1981) to describe the sensory organs of leg IV of A. abyssalis is inadequate since those of the holotype and MNHN AP345/560Ma had a hemispherical appearance. Further, we consider MNHN AE120/78Ma with ���balloon-shaped��� leg IV sensory organs (in the sense of Bussau 1992 and Villora-Moreno 1998) as an undescribed species. The difference in the morphology of the tertiary clavae also supports this view. However, MNHN AE120/78Ma is not in an acceptable condition for species description. Based on the observation of the holotype and MNHN AP345/560Ma, Angursa abyssalis resembles A. clavifera and A. capsula by the club-shaped primary clavae longer than the lateral cirri, the leg IV sensory organs in hemispherical shape, and the lack of the anal papillae (Noda 1985; Bussau 1992). However, it differs from A. clavifera by the larger secondary clavae, the tertiary clavae slightly indented around the external cirri, and the absence of sensory organs on legs II and III (Noda 1985). It differs from A. capsula by the larger and closer situated pair of secondary clavae (3 ��m apart in MNHN AP345/560Ma and 9 ��m apart in A. capsula measured from the drawing of a paratypic female in the original description see Bussau 1992), the tertiary clavae slightly indented around the external cirri (Bussau 1992) and a much shorter cirrus E (10 ��m (body length, 141 ��m) in A. abyssalis versus 19 ��m (170 ��m) in A. capsula)., Published as part of Fujimoto, Shinta & Hansen, Jesper Guldberg, 2019, Revision of Angursa (Arthrotardigrada: Styraconyxidae) with the description of a new species from Japan, pp. 1-19 in European Journal of Taxonomy 510 on pages 5-7, DOI: 10.5852/ejt.2019.510, http://zenodo.org/record/2616378, {"references":["Renaud-Mornant J. 1981. Deux nouveaux Angursa Pollock, 1979, du domaine abyssal (Tardigrada, Arthrotardigrada). Tethys 10: 161 - 164.","Bussau C. 1992. New deep-sea Tardigrada (Arthrotardigrada, Halechiniscidae) from manganese nodule area of the eastern South Pacific. Zoologica Scripta 21: 79 - 91. https: // doi. org / 10.1111 / j. 1463 - 6409.1992. tb 00311. x","Villora-Moreno S. 1998. Deep-sea Tardigrada from South Shetland Islands (Antarctica) with description of Angursa antarctica sp. nov. (Arthrotardigrada, Halechiniscidae). Polar Biology 19: 336 - 341. https: // doi. org / 10.1007 / s 003000050255","Noda H. 1985. Description of a new subspecies of Angursa bicuspis Pollock (Heterotardigrada, Halechiniscidae) from Tanabe Bay, Japan. Publications of the Seto Marine Biological Laboratory 30: 269 - 276. Available from https: // hdl. handle. net / 2433 / 176110 [accessed 4 Feb. 2019]."]}

Angursa bicuspis Pollock, 1979 Fig. 1 Emended diagnosis Angursa with presence of median cirrus uncertain; primary clavae longer than lateral cirri; pedestals of primary clavae and lateral cirri absent; morphology of secondary and tertiary clavae unknown; simple, tapering cirri E present; leg I sensory organs present; presence of legs II and III sensory organs uncertain; leg IV sensory organs each as papilla with short, apical spine; robust anal papillae present; presence of seminal receptacles uncertain; presence of proximal pads of internal digits uncertain. Material examined Holotype UNITED STATES OF AMERICA ��� 1 adult; Massachusetts, Buzzard Bay side of Penzance Point, Woods Hole; 41��31���41��� N, 70��40���41��� W; 9 Jul. 1968; Hummon & Pollock leg.; USNM W 50900. Remarks Angursa bicuspis was described based on the holotype and an additional specimen (Pollock 1979). However, their conspecificity has been doubted by Kaczmarek et al. (2015) owing to the fact that the two specimens were collected from two different environments of distant localities and that they showed morphological differences (Pollock 1979). As there is no positive evidence to regard the two specimens as conspecific, we support the opinion of Kaczmarek et al. (2015) and consider it as sufficient to base our attempt to re-diagnose A. bicuspis on only the holotype. The holotype is in bad condition. However, our observation confirmed that of Pollock (1979) in the overall body shape (Fig. 1A) and the presence of the internal cirri (Fig. 1B), the external cirri (which is 8 ��m, not 5.5 ��m as reported by Pollock 1979) (Fig. 1B), the lateral cirri, the primary clavae (Fig. 1B), the leg I sensory organs, the cirri E, the characteristic claws, and the two papillae posterior to the anus (Fig. 1C). Pollock (1979) considered the latter structure as not useful for distinguishing species referring to the variability of the caudal projection of Batillipes Richters, 1909 and the fact that it was not observed in the additional specimen. Villora-Moreno (1998) supported this view, but Noda (1985) and Bussau (1992) used this structure to distinguish this species from its congeners (reason not explained). We agree with Noda (1985) and Bussau (1992) for the following two reasons. Firstly, we consider that this character is not homologous to the caudal projection of Batillipes as Pollock (1979) suggested, but a developed form of the cuticular fold that closes the anus, homologous to the ���two weak protuberances��� of A. antarctica described by Villora-Moreno (1998). Secondly, as discussed in the previous paragraph, this character���s presence/absence in the two specimens of Pollock (1979) is likely an interspecific variation. Thus, A. bicuspis can be easily distinguished from its congeners by the presence of the well-developed anal papillae (Renaud-Mornant 1981; Noda 1985; Bussau 1992; Villora-Moreno 1998), consequently supporting the species status of A. abyssalis proposed by Bussau (1992). In addition to confirming and re-interpreting the original observation (Pollock 1979), we recognize three overlooked characters: a paired globular body fringing the mouth (Fig. 1 B), the leg IV sensory organs and the peduncles (3 ��m) that do not reach the claws (Fig. 1D). Each leg IV sensory organ consists of a spherical papilla and a short, apical spine (Fig. 1 C). An enveloping cuticular sheath of the papilla was not evident, but may have been obscured due to the squeezed state of the specimen. The bad condition of the specimen also hindered our decisions on the presence/absence of the median cirrus, the secondary and tertiary clavae, legs II and III sensory organs, proximal pads of the internal digits and the seminal receptacles. Additional collection of specimens from the type locality is necessary for more detailed discussion on this species morphology., Published as part of Fujimoto, Shinta & Hansen, Jesper Guldberg, 2019, Revision of Angursa (Arthrotardigrada: Styraconyxidae) with the description of a new species from Japan, pp. 1-19 in European Journal of Taxonomy 510 on pages 3-4, DOI: 10.5852/ejt.2019.510, http://zenodo.org/record/2616378, {"references":["Pollock L. W. 1979. Angursa bicuspis n. g., n. sp., a new marine arthrotardigrade from the western North Atlantic. Transactions of the American Microscopical Society 98: 558 - 562. https: // doi. org / 10.2307 / 3225907","Kaczmarek L., Bartels P. J., Roszkowska M. & Nelson D. R. 2015. The zoogeography of marine tardigrades. Zootaxa 4037: 1 - 189. https: // doi. org / 10.11646 / zootaxa. 4037.1.1","Villora-Moreno S. 1998. Deep-sea Tardigrada from South Shetland Islands (Antarctica) with description of Angursa antarctica sp. nov. (Arthrotardigrada, Halechiniscidae). Polar Biology 19: 336 - 341. https: // doi. org / 10.1007 / s 003000050255","Noda H. 1985. Description of a new subspecies of Angursa bicuspis Pollock (Heterotardigrada, Halechiniscidae) from Tanabe Bay, Japan. Publications of the Seto Marine Biological Laboratory 30: 269 - 276. Available from https: // hdl. handle. net / 2433 / 176110 [accessed 4 Feb. 2019].","Bussau C. 1992. New deep-sea Tardigrada (Arthrotardigrada, Halechiniscidae) from manganese nodule area of the eastern South Pacific. Zoologica Scripta 21: 79 - 91. https: // doi. org / 10.1111 / j. 1463 - 6409.1992. tb 00311. x","Renaud-Mornant J. 1981. Deux nouveaux Angursa Pollock, 1979, du domaine abyssal (Tardigrada, Arthrotardigrada). Tethys 10: 161 - 164."]}

Angursa seisuimaruae sp. nov. urn:lsid:zoobank.org:act: 208985F1-BBB3-40B4-A2E4-26E8662277DD Figs 6���7 Angursa sp. ��� Kimura et al. 2018: table 23. Diagnosis Angursa with club-shaped primary clavae longer than lateral cirri; pedestals of primary clavae and lateral cirri absent; D-shaped secondary clavae; three-lobed tertiary clavae; leg I sensory organs as subdivided spines; legs II and III sensory organs absent; leg IV sensory organs each as papilla enveloped in hemispherical cuticular sheath with short, apical spine; anal papillae absent; seminal receptacles open latero-ventrally with short, sinuous ducts. Etymology The specific epithet refers to the T/V Seisuimaru of Mie University. Type material Holotype PACIFIC OCEAN ��� adult ♀; Northwestern Pacific, Japan, Sea of Kumano; between 33��59.5��� N, 136��56.7��� E and 33��59.7��� N, 136��56.7��� E; 768���796 m below sea level; 10 Nov. 2017; mounted in glycerol; Fujimoto leg.; KUZ Z2066. Paratype PACIFIC OCEAN ��� 1 adult ♀; same data as for holotype; KUZ Z2067. Type locality Sea of Kumano, Northwestern Pacific (between 33��59.5��� N, 136��56.7��� E and 33��59.7��� N, 136��56.7��� E; depth range, 768���796 m) (Kimura et al. 2018). Description of holotype Adult female with unarmoured, slender body (143 ��m long; 35 ��m wide between legs II and III) (Figs 6, 7A). Epicuticular pillars visible. Cephalic region with unpaired median cirrus, paired internal cirri, paired external cirri, paired lateral cirri, paired primary clavae, paired secondary clavae, paired tertiary clavae, and mouth opening. Median cirrus (1 ��m) positioned dorsally. Internal cirri (5 ��m) inserted on anterior margin of body each consisting of scapus (2 ��m) and distal flagellar part with blunt tip (2 ��m). External cirri (9 ��m) inserted ventrally each consisting of scapus (5 ��m) and distal flagellar part with blunt tip (5 ��m). Lateral cirri (4 ��m) positioned dorso-laterally. Primary clavae (10 ��m) positioned immediately ventral to lateral cirri and both organs inserted directly on body with no apparent pedestal (only very slight rise). Primary clavae club-shaped and distal half slightly bent posteriorly. Secondary clavae (Figs 6, 7B) present posterior to median cirrus as flat, D-shaped structures without epicuticular pillars. Tertiary clavae (Figs 6, 7C) indented around external cirri, giving impression of three-lobed structures (one lobe directed anteriorly, interior to external cirri; second lobe developed interior to first lobe; third lobe developed posterior to external cirri) without epicuticular pillars. Anteriorly directed mouth at antero-most position of body. Buccal apparatus dissolved and only pharyngeal bulb recognised (length 14 ��m, width 11 ��m). Paired cirri E (18 ��m) (Figs 6, 7D) as simple, tapering spines without recognisable annulation, inserted dorsally slightly anterior to leg IV. Each leg consists of indistinct coxa, femur, indistinct tibia, and tarsus bearing four digits terminating in claws with two divergent points. Leg I sensory organ (10 ��m) (Figs 6, 7E) present on femur as spine consisting of proximal portion and distal portion with blunt tip. Legs II and III lack sensory organs. Leg IV sensory organs (4 ��m) (Figs 6, 7D) present at base of leg IV each as papilla enveloped in hemispherical cuticular sheath with short, apical spine. Internal digits have proximal pads and lack peduncles (Figs 6, 7E). External digits have arched peduncles at their base which never reach claws (Figs 6, 7E). Internal digits always longer than external digits and leg IV has longer digits than those of legs I���III (leg I, internal digits 9 ��m/external digits 6 ��m; leg II, 9/6; leg III, 10/6; leg IV, 13/7). Rosette-like gonopore (4 ��m in diameter) (Figs 6, 7F) situated 2 ��m anterior to anus. Longitudinal cuticluar fold of anus reach dorsal side (Figs 6, 7D). Paired seminal receptacles (Figs 6, 7F) positioned near latero-ventral margin of body, at level slightly anterior to gonopore. Each seminal receptacle duct runs interiorly, antero-dorsally, and then postero-ventrally terminating in round vesicle. Description of paratype Cephalic region accidentally destroyed during cleaning of specimen before observation and mouth opening, buccal apparatus, median cirrus, secondary clavae, and tertiary clavae not recognized. Primary clava (11 ��m) and scapi of internal, external, and lateral cirri recognized on torn cuticle. Presence of scapi suggests subdivision of lateral cirri as in the internal and external cirri. Gonopore and anus recognised before permanent preparation in lateral positioned body. Other characters confirm that of holotype (cirrus E, 17 ��m; leg I sensory organ, 10 ��m; leg IV sensory organ, 4 ��m). Remarks Angursa seisuimaruae sp. nov. resembles A. abyssalis, A. clavifera, and A. capsula by the club-shaped primary clavae longer than the lateral cirri, the leg IV sensory organs in hemispherical shape, and the lack of the anal papillae (Renaud-Mornant 1981; Noda 1985; Bussau 1992). However, the new species is distinguished from all three species by its D-shaped secondary clavae and its three-lobed tertiary clavae (Noda 1985; Bussau 1992). Further, the new species���s seminal receptacle ducts are shorter than A. clavifera ���s spiral ones (Noda 1985). Identification key to Angursa species 1. Club-shaped primary clavae longer than lateral cirri........................................ 2 (A. bicuspis group) ��� Bulbous primary clavae shorter than lateral cirri...........................................6 (A. lanceolata group) 2. Well developed anal papillae..................................................................... A. bicuspis Pollock, 1979 ��� Anal papillae absent........................................................................................................................... 3 3. Legs II and III sensory organs present; spiral seminal receptacle ducts........ A. clavifera Noda, 1985 ��� Leg II and III sensory organs absent.................................................................................................. 4 4. Three-lobed tertiary clavae; D-shaped secondary clavae; seminal receptacles present........................................................................................................................................... A. seisuimaruae sp. nov. ��� Tertiary clavae with no indentation or only slightly indented around external cirri.......................... 5 5. Secondary clavae situated close together (ca 3 ��m); short (10 ��m) cirrus E.......................................................................................................................................... A. abyssalis Renaud-Mornant, 1981 ��� Secondary clavae situated apart (ca 9 ��m); long (19 ��m) cirrus E............... A. capsula Bussau, 1992 6. Leg IV sensory organs as papillae enveloped in hemispherical sheaths; cirri E with annulated scapi............................................................................................ A. antarctica Villora-Moreno, 1998 ��� Leg IV sensory organs as elongate papillae....................................................................................... 7 7. Cirri E lance-shaped................................................................ A. lanceolata Renaud-Mornant, 1981 ��� Cirri E tongue-shaped..................................................................................... A. lingua Bussau, 1992 Identification key to Angursa species 1. Club-shaped primary clavae longer than lateral cirri........................................ 2 (A. bicuspis group) ��� Bulbous primary clavae shorter than lateral cirri...........................................6 (A. lanceolata group) 2. Well developed anal papillae..................................................................... A. bicuspis Pollock, 1979 ��� Anal papillae absent........................................................................................................................... 3 3. Legs II and III sensory organs present; spiral seminal receptacle ducts........ A. clavifera Noda, 1985 ��� Leg II and III sensory organs absent.................................................................................................. 4 4. Three-lobed tertiary clavae; D-shaped secondary clavae; seminal receptacles present........................................................................................................................................... A. seisuimaruae sp. nov. ��� Tertiary clavae with no indentation or only slightly indented around external cirri.......................... 5 5. Secondary clavae situated close together (ca 3 ��m); short (10 ��m) cirrus E.......................................................................................................................................... A. abyssalis Renaud-Mornant, 1981 ��� Secondary clavae situated apart (ca 9 ��m); long (19 ��m) cirrus E............... A. capsula Bussau, 1992 6. Leg IV sensory organs as papillae enveloped in hemispherical sheaths; cirri E with annulated scapi............................................................................................ A. antarctica Villora-Moreno, 1998 ��� Leg IV sensory organs as elongate papillae....................................................................................... 7 7. Cirri E lance-shaped................................................................ A. lanceolata Renaud-Mornant, 1981 ��� Cirri E tongue-shaped..................................................................................... A. lingua Bussau, 1992, Published as part of Fujimoto, Shinta & Hansen, Jesper Guldberg, 2019, Revision of Angursa (Arthrotardigrada: Styraconyxidae) with the description of a new species from Japan, pp. 1-19 in European Journal of Taxonomy 510 on pages 10-13, DOI: 10.5852/ejt.2019.510, http://zenodo.org/record/2616378, {"references":["Kimura T., Kimura S., Jimi N., Kakui K., Tomioka S., Oya Y., Matsumoto Y., Tanabe Y., Hasegawa N., Hookabe N., Homma R., Hosoda Y., Fujimoto S., Kuramochi T., Fujita T., Ogawa A., Kobayashi I., Ishida Y., Tanaka H., Onishi H., Shimetsugu M., Yoshikawa A., Tanaka M., Kushida Y., Maekawa Y., Nakamura T., Okumura J. & Tanaka K. 2018. Benthic deep-sea fauna in the Sea of Kumano, Mie Prefecture, Japan. Annals of Field Research and Technology Mie University 16: 1 - 32. [In Japanese.]","Renaud-Mornant J. 1981. Deux nouveaux Angursa Pollock, 1979, du domaine abyssal (Tardigrada, Arthrotardigrada). Tethys 10: 161 - 164.","Noda H. 1985. Description of a new subspecies of Angursa bicuspis Pollock (Heterotardigrada, Halechiniscidae) from Tanabe Bay, Japan. Publications of the Seto Marine Biological Laboratory 30: 269 - 276. Available from https: // hdl. handle. net / 2433 / 176110 [accessed 4 Feb. 2019].","Bussau C. 1992. New deep-sea Tardigrada (Arthrotardigrada, Halechiniscidae) from manganese nodule area of the eastern South Pacific. Zoologica Scripta 21: 79 - 91. https: // doi. org / 10.1111 / j. 1463 - 6409.1992. tb 00311. x","Pollock L. W. 1979. Angursa bicuspis n. g., n. sp., a new marine arthrotardigrade from the western North Atlantic. Transactions of the American Microscopical Society 98: 558 - 562. https: // doi. org / 10.2307 / 3225907","Villora-Moreno S. 1998. Deep-sea Tardigrada from South Shetland Islands (Antarctica) with description of Angursa antarctica sp. nov. (Arthrotardigrada, Halechiniscidae). Polar Biology 19: 336 - 341. https: // doi. org / 10.1007 / s 003000050255"]}

Fujimoto, Shinta (2018): A New Species of Tanarctus (Heterotardigrada: Arthrotardigrada: Tanarctidae) from Oku-Matsushima, Japan. Species Diversity 23: 209-213, DOI: 10.12782/specdiv.23.209

Fujimoto, S. (2015) Quisarctus yasumurai gen. et sp. nov. (Arthrotardigrada: Halechiniscidae) from a submarine cave, off Iejima, Ryukyu Islands, Japan. Zootaxa. 3948 (1): 145-150. Fujimoto, S. (2014) A new Stygarctus (Arthrotardigrada: Stygarctidae) from Japan, with entangled seminal receptacle ducts. Zootaxa. Magnolia Press. 3784 (2): 187-195.

Quisarctus gen. nov. Diagnosis. Halechiniscidae with cylindrical body; primary clava and lateral cirrus arise from common cirrophore; primary clava longer than lateral cirrus; secondary clava undeveloped; stylet supports present; leg IV sense organ as elongate papilla with distal spine; legs terminate in digits without folds, peduncles, proximal pads, pretarsi or wrinkles; internal digits longer than external digits; each digit terminates in sheathed, small, crescent-shaped claw with minute calcar; pair of ventrally opening seminal receptacles each with slender, sinuous duct terminating in spherical vesicle. Etymology. The generic epithet is a combination of Quis, a latin word for ���Who?��� or ���What?��� referring to the author���s first impression of this animal, and arctus, a Latinised Greek word for ���bear��� referring to the common name of tardigrades. Type species. Quisarctus yasumurai sp. nov., Published as part of Fujimoto, Shinta, 2015, Quisarctus yasumurai gen. et sp. nov. (Arthrotardigrada: Halechiniscidae) from a submarine cave, off Iejima, Ryukyu Islands, Japan, pp. 145-150 in Zootaxa 3948 (1) on page 146, DOI: 10.11646/zootaxa.3948.1.10, http://zenodo.org/record/239751

Quisarctus yasumurai sp. nov. (Figs. 1, 2; Table 1) Diagnosis. Halechiniscidae with cylindrical body; primary clava and lateral cirrus arise from common cirrophore; primary clava longer than lateral cirrus; secondary clava undeveloped; stylet supports present; leg I sense organ consists of three parts; leg II sense organ as unarticulated spine; leg III sense organ as unarticulated spine; leg IV sense organ as elongate papilla with distal spine; all legs terminate in four digits without folds, peduncles, proximal pads, pretarsi or wrinkles; internal digits longer than external digits; each digit terminates in sheathed, small, crescent-shaped claw with minute calcar; pair of ventrally opening seminal receptacles each with slender, sinuous duct terminating in spherical vesicle. Material examined. Holotype: KUZ Z 1649, adult female. Paratypes: KUZ Z 1650, adult female; KUZ Z 1651, four-digit juvenile. Type locality: Water depth of 29 m, ���Daidokutsu��� cave, off Iejima, Okinawa Islands, Ryukyu Islands, Japan (26 �� 43 ' 18 "N, 127 �� 50 '00"E). Coll. by Koshin Yasumura on 6 th November 2013. Type depository: The type series is deposited in the Zoological Collection of Kyoto University. Description of holotype (Figs. 1, 2 A, 2 B). Adult female, body length 106 ��m. Cylindrical body with smooth cuticle. Round cephalic region without lobes. Complete set of three parted cephalic cirri present. Dorsal median cirrus with scapus (2 ��m), tubular portion (7 ��m) and flagellum (3 ��m) inserted 13 ��m posterior to anterior margin of body. Dorsal internal cirrus with scapus (2 ��m), tubular portion (5 ��m) and flagellum (4 ��m) inserted on short cirrophore. Ventral external cirrus with scapus (4 ��m), tubular portion (4 ��m) and flagellum (2 ��m) slightly exterior to internal cirrus. Internal cirri and external cirri situated near anterior margin of body. Lateral cirrus with scapus (3 ��m), tubular portion (5 ��m) and flagellum (4 ��m) inserted on short lateral cirrophorus with primary clava. Elongated, proximally-inflated primary clava (27 ��m) with distal pore. Lateral cirrus situated antero-dorsal to primary clava. Secondary clava undeveloped. Mouth directed anteriorly. Observation of bucco-pharyngeal apparatus incomplete due to limited visibility in mounting media. Bucco-pharyngeal apparatus with buccal tube (21 ��m), three placoids, pharyngeal bulb (11 ��m �� 11 ��m) and two stylets (furca not observed). Stylet supports present but not observed in holotype. Cirrus E (20 ��m) with proximal portion, distal portion and tapering spine inserted on short cirrophore. Leg I sense organ (5 ��m) articulated into three parts. Leg II and III sense organs (8 ��m, 7 ��m) as unarticulated spines. Leg IV sense organ (5 ��m) with proximal papilla and short, distal tapering. Each leg terminates in four digits without folds, peduncles, proximal pads, pretarsi or wrinkles. Internal digits longer than external digits. Digits on leg IV longer than that of legs I���III. Each digit terminates in sheathed, small, crescent-shaped claw without dorsal spur. Minute calcar recognised in well-positioned claws of internal and external digits. Rossette-like female gonopore opens 5 ��m anterior to caudal anus. Pair of seminal receptacles present. Slender, sinuous seminal receptacle duct opens ventrally near lateral margin of body slightly posterior to level of gonopore and runs exteriorly and then dorsally terminating in spherical vesicle. In holotype, seminal vesicle directed dorsally gives spherical appearance. Remarks on paratypes. In the paratypic female, KUZ Z 1650, the digits were in better position for observation (Figs. 2 C, D). In addition, the stylet supports were more obvious (Fig. 2 E) and the seminal vesicles were observed from lateral view (Fig. 2 F). The other paratype, KUZ Z 1651, was identified as a four-digit juvenile based on difficultly in observing the gonopore region and absence of both seminal receptacles and spermatozoa. Etymology. The specific epithet, yasumurai, is named in honor of Koshin Yasumura who collected the sediment samples from which the new species was obtained., Published as part of Fujimoto, Shinta, 2015, Quisarctus yasumurai gen. et sp. nov. (Arthrotardigrada: Halechiniscidae) from a submarine cave, off Iejima, Ryukyu Islands, Japan, pp. 145-150 in Zootaxa 3948 (1) on pages 146-148, DOI: 10.11646/zootaxa.3948.1.10, http://zenodo.org/record/239751

Fujimoto, Shinta (2015): Quisarctus yasumurai gen. et sp. nov. (Arthrotardigrada: Halechiniscidae) from a submarine cave, off Iejima, Ryukyu Islands, Japan. Zootaxa 3948 (1): 145-150, DOI: http://dx.doi.org/10.11646/zootaxa.3948.1.10

Stygarctus ayatori sp. nov. (Figs. 2���6, Table 1) Diagnosis. Stygarctus with club shaped primary clavae and bent sausage-like secondary clavae; funnel shaped lateral processes without micro-spines; pair of dorsal spines with terminal bifurcation; caudal plate with pair of antero-lateral lobes, pair of round lateral processes with minute spike and pair of caudal spikes; ventral surface without plates; female genital structure with seminal receptacle ducts entangled near exterior opening and internal thickening situated peripheral to female gonopore and between female gonopore and anus. Materials examined. Holotype: KUZ Z 677: adult female mounted in Fluoromount-G TM. Paratypes: KUZ Z 678: adult female mounted in glycerol; KUZ Z 679, Z 680, Z 681: three adult females mounted in Hoyer���s medium; KUZ Z 682: adult male mounted in Fluoromount-G TM; KUZ Z 683: adult male mounted in Hoyer���s medium. Type Locality: Okinoshima, Tateyama Bay, Boso Peninsula, Honshu, Japan (34 �� 59 ��� 27 ���N, 139 �� 49 ��� 24 ���E). Coll. Shinta Fujimoto on 18 th April 2013. Type depositories: The type series is deposited in the Zoological Collection of Kyoto University (KUZ). The two specimens mounted in distilled water and the four specimens on a SEM stub are deposited in the author���s personal collection. Description of holotype. Adult female with body length of 112 ��m. Dorsal surface with sculptured dorsal plates (Figs. 2, 5 A); ventral surface smooth without plates or cuticle folds (Fig. 3). Cephalic plate with medial lobe, pair of medio-lateral lobes and pair of lateral lobes. Ventral lobes on each side of anteriorly protruded mouth cone. FIGURE 2. Drawing of Stygarctus ayatori sp. nov., holotype female [KUZ Z 677] (dorsal view). cE, cirrus E; cs, caudal spike; ds, dorsal spine; ec, external cirrus; ic, internal cirrus; lc, lateral cirrus; mc, median cirrus; ms, minute spike; pcl, primary clava; scl, secondary clava; s 4, leg IV sensory organ. FIGURE 3. Drawing of Stygarctus ayatori sp. nov., holotype female [KUZ Z 677] (ventral view). a, anus; cE, cirrus E; cs, caudal spike; ds, dorsal spine; ec, external cirrus; fg, female gonopore; ic, internal cirrus; it, internal thickening; lc, lateral cirrus; ms, minute spike; pcl, primary clava; scl, secondary clava; sr, seminal receptacle; s 4, leg IV sensory organ. Bucco-pharyngeal apparatus dissolved before observation. Unpaired median cirrus (12 ��m) with short scapus (2 ��m) and flagellum (10 ��m) inserted medially at level of incision between medio-lateral and lateral lobes. Medial lobe bears pair of internal cirri (12 ��m) each with short scapus (3 ��m) and flagellum (9 ��m) arising from cirrophore. Lateral lobe bears lateral cirrus (14 ��m) with short scapus (3 ��m) and flagellum (11 ��m) and club shaped primary clava (6 ��m) inserted dorso-posterior to lateral cirrus. Thin sheet present exterior to lateral lobe. Ventral lobe bears external cirrus (11 ��m) with short scapus (2 ��m) and flagellum (9 ��m) and sausage-like secondary clava (16 ��m) inserted interior to external cirrus. Three dorsal body plates with funnel shaped lateral processes, reduced to thin sheets at lateral margin and micro-spines absent. Two intersegmental plates present. Pair of dorsal spines (23 ��m) with terminal bifurcation fringed with thin cuticle inserted on posterior ridge of second dorsal body plate. Caudal plate with pair of anterolateral lobes, pair of round lateral processes each with minute spike and pair of caudal spikes. Round lateral process reduced to thin sheet at lateral margin. Pair of Cirri E (24 ��m) each with proximal portion (6 ��m) and flagellum (18 ��m) inserted between antero-lateral lobe and round lateral process. Papilla shaped leg IV sense organ (2 ��m) inserted at base of leg IV. Rosette-like female gonopore opens ventrally 8 ��m anterior to anus. Pair of seminal receptacles each consists of sinuous duct opening 2 ��m posterior from level of gonopore and small vesicle positioned 11 ��m laterally from the gonopore. Seminal receptacle duct forming a three-dimensional nodule of entanglement near duct opening. Terminal anus consists of two cuticle lobes. Region peripheral to gonopore and between gonopore and anus with internal thickening. Four claws inserted directly on each leg with dorsal, long, thin filaments on median claws. Claws of leg IV (7 ��m) longer than those of leg I���III (5 ��m). Etymology. The specific epithet, ayatori, is a Japanese word for ���string figure��� referring to the seminal receptacle ducts of the new species. Remarks on paratypes and additional material. The bucco-pharyngeal apparatus was observed in specimens mounted in distilled water (Figs. 4 A, 5 B). It consists of a buccal tube (22 ��m), two stylets with furca, three apophyses (3 ��m) and three placoids (3 ��m) for a specimen with body length of 111 ��m. The pharyngeal bulb was not observed. The female genital structure was identical in all material observed with no variation in the number of turns or loops in the ducts or in the shape of the internal thickening (Figs. 4 B, 5 C). The male gonopore, situated immediately anterior to the anus, is a simple round opening of a posteriorly directed 4 ��m long tube (Fig. 4 C). The difference in the genital structures of the female and male was the only sexual dimorphism recognized in the new species. Results of electron scanning microscopy are described as follows. The dorsal plates are strongly sculptured and each plate is partitioned only by the elevation of the cuticle (Figs. 6 A, B). The three-dimensional appearance of the cephalic region is better observed than under DIC microscopy (Fig. 6 C). The female gonopore opens on a triangular button shaped lobe, which is not visible with DIC (Fig. 6 D). The smooth surface posterior to the gonopore indicates the internal thickening observed with DIC is an internal structure (Fig. 6 D). Differential diagnosis. Stygarctus ayatori sp. nov. is distinguished from S. arbornatus (for which there is no information on the genital structure) by the lack of dorsal spines, which are present in the new species. Stygarctus bradypus is most similar to S. ayatori sp. nov. in the morphology of sensory organs, dorsal plates, lateral processes, dorsal spines and caudal spikes but the new species is distinguished from S. bradypus, and the other congeners, by the entangled seminal receptacle ducts and internal thickening in the female genital region. In contrast to the new species, these congeners have simple ducts with loops or spirals and no internal thickening reported with the exception of a v-shaped thickening situated between the pair of seminal receptacles in S. gourbaultae Renaud- Mornant, 1981 (Pollock 1970 a, b; Renaud-Mornant 1970, 1981; Grimaldi de Zio et al. 1987; Hansen et al. 2012)., Published as part of Fujimoto, Shinta, 2014, A new Stygarctus (Arthrotardigrada: Stygarctidae) from Japan, with entangled seminal receptacle ducts, pp. 187-195 in Zootaxa 3784 (2) on pages 188-194, DOI: 10.11646/zootaxa.3784.2.8, http://zenodo.org/record/230089, {"references":["Renaud-Mornant, J. (1981) Stygarctus gourbaultae n. sp. un nouveau tardigrade marin (Arthrotardigrada) de la Guadeloupe. Bulletin du Museum National d'Histoire Naturelle, Paris 4 e serie 3 section A (Zoologie, Biologie et Ecologie Animales), no. 1, 175 - 180.","Pollock, L. W. (1970 a) Batillipes dicrocerus n. sp., Stygarctus granulatus n. sp. and other tardigrada from Woods Hole, Massachusetts, USA. Transactions of the American Microscopical Society, 89, 38 - 52.","Pollock, L. W. (1970 b) Reproductive anatomy of some marine Heterotardigrada. Transactions of the American Microscopical Society, 89, 308 - 316.","Renaud-Mornant, J. (1970) Parastygarctus sterreri n. sp., tardigrade marin nouveau de l'Adriatique. Cahiers de Biologie Marine, 11, 355 - 360.","Hansen, J. G., Kristensen, R. M. & Jorgensen, A. (2012) The armoured marine tardigrades (Arthrotardigrada, Tardigrada). Scientia Danica. Series B, Biologica, 2, 1 - 91."]}