Deep-sea Fauna and Pollutants off Pacific Coast of Northern Japan, edited by T. Fujita, National Museum of Nature andScience Monographs, No. 39, pp. 39 192, 2009

Deep-sea Benthic Polychaetes off Pacific Coast of
the Northern Honshu, Japan
Minoru Imajima
Department of Zoology, National Museum of Nature and Science,
3 23 1 Hyakunin-cho, Shinjuku-ku, Tokyo, 169 0073 Japan
E-mail: imajima@kahaku.go.jp

Abstract: Polychaetous annelids of the deep-sea off Pacific coast of the northern Honshu through the research
project conducted during 2005 2007 by the National Museum of Nature and Science, Tokyo, were taxonomically examined. A total of 211 identified species and 32 indeterminable species in 43 families of polychaetes
are recognized. Fourteen species are new to science: Weberia abyssicola (Polynoidae), Sigalion orientalis(Sigalionidae), Glycera neorobusta, Glycera okai and Glycera semibranchiopoda (Glyceridae), Nereis abyssa
and Rullierinereis profunda (Nereididae), Euphrosine digitalis and Euphrosine pseudonotialis (Euphrosinidae), Paradiopatra gracilis (Onuphidae), Naineris japonica (Orbiniidae), Laubieriopsis brevis japonica (Fauveliopsidae), Pseudoscalibregma orientalis (Scalibregmatidae) and Anobothrus wakatakamaruae (Ampharetidae). Thirty-seven species are newly added to the Japanese polychaetous fauna. Eighty-six species are
known also from the northwestern Pacific Ocean, including Sea of Okhotsk and Bering Sea. Forty species are
known only from Japan, or endemic. The Fauveliopsidae is newly added to the Japanese fauna.
Key words: Off northern Honshu, deep-sea, polychaetes, new species, taxonomy.

Introduction
The deep-sea research project “Study on Deep-Sea Fauna and Conservation of Deep-Sea
Ecosystem” was conducted during the period 2005 2007 off Pacific coast of the northern Honshu
by the National Museum of Nature and Science, Tokyo. The Pacific coast of the northern Honshu
is influenced by the cold Oyashio Current which originates in Arctic seas. The Oyashio Current
goes toward the Boso Peninsula and causes a rapid decrease in temperature of the water which has
been flowing northward as the warm Kuroshio Current. Its effects are greatest in winter when thecold current approaches the Boso Peninsula.
In this survey, polychaetes were collected from 232 stations, in depths between 150 and 5268
m, distributed all over off Pacific coast of the northern Honshu. Details regarding sampling sites(station, latitude and longitude, depth, date) are given in Table 1. The detailed map of all localities
is mentioned in the text (Fig. 1). The samples were collected by the R/Vs Wakataka-maru andSoyo-maru of the Fisheries Research Agency (FRA) and R/V Tansei-maru of the Japan Agency for
Marine-Earth Science and Technology (JAMSTEC). Sampling was carried out by various types of
biological dredges and trawls.
The polychaete fauna from the deep-sea off Pacific coast of the northern Honshu was still
poorly known. In this survey, a total of 211 identified species and 32 indeterminable species in 43
families of polychaetes were recognized. Fourteen species, Weberia abyssicola (Polynoidae), Sigalion orientalis (Sigalionidae), Glycera neorobusta, Glycera okai and Glycera semibranchiopoda(Glyceridae), Nereis abyssa and Rullierinereis profunda (Nereididae), Euphrosine digitalis andEuphrosine pseudonotialis (Euphrosinidae), Paradiopatra gracilis (Onuphidae), Naineris japonica (Orbiniidae), Laubieriopsis brevis japonica (Fauveliopsidae), Pseudoscalibregma orientalis

40

Minoru Imajima

Fig. 1. Map showing collected stations of polychaetes in the benthic surveys operated by Wakataka-maru (●), Soyomaru (▲) and Tansei-maru (×) during 2005 2007.

41

Deep-sea Benthic Polychaetes off Northern Honshu

Table 1. Collection data for stations at where polychaetes were collected in surveys during 2005 2007.
StationWA05-E410
WA05-E425
WA05-E450
WA05-E480
WA05-E510
WA05-E1000D
WA05-E550
WA05-E750
WA05-E900
WA05-F425
WA05-F450
WA05-F480
WA05-F510
WA05-F1200
WA05-F900
WA05-G150
WA05-G210
WA05-G280
WA05-G310
WA05-H150
WA05-H310
WA05-H350
WA05-H450
WA05-H510
WA05-H380
WA05-H650
WA05-H750
WA05-H900D
WA05-G410
WA05-G550
WA05-F350
WA05-F380
WA05-F410
WA05-G425
WA05-G450
WA05-G510
WA05-G1500D
WA05-G750
WA05-G900
WA05-GH250
WA05-GH280
WA05-GH310
WA05-GH350
WA05-GH510
WA05-GH510D
WA05-GH380
WA05-GH380D
WA05-GH425
WA05-GH450
WA05-GH480
WA05-FG250D
WA05-FG350
WA05-FG410
WA05-FG450
WA05-FG480

Position
38˚23.69´N,142˚02.59´E 38˚22.12´N,142˚03.29´E
38˚24.09´N,142˚03.01´E 38˚25.74´N,142˚02.65´E
38˚23.56´N,142˚03.98´E 38˚25.15´N,142˚03.66´E
38˚22.64´N,142˚05.29´E 38˚20.92´N,142˚05.97´E
38˚22.50´N,142˚06.32´E 38˚23.58´N,142˚05.76´E
38˚26.74´N,142˚23.82´E
38˚22.62´N,142˚07.26´E 38˚23.64´N,142˚07.55´E
38˚22.11´N,142˚13.89´E 38˚23.08´N,142˚14.47´E
38˚28.88´N,142˚21.39´E 38˚29.69´N,142˚21.61´E
37˚44.32´N,141˚54.82´E 37˚42.55´N,141˚55.03´E
37˚43.65´N,141˚56.61´E 37˚45.31´N,141˚56.38´E
37˚41.93´N,141˚59.00´E 37˚40.20´N,141˚59.02´E
37˚39.44´N,142˚01.15´E 37˚38.22´N,142˚01.09´E
37˚47.60´N,142˚37.12´E 37˚47.40´N,142˚37.17´E
37˚46.70´N,142˚18.75´E 37˚45.73´N,142˚19.06´E
36˚59.79´N,141˚17.45´E 37˚01.31´N,141˚17.80´E
36˚57.13´N,141˚22.78´E 36˚58.48´N,141˚23.86´E
36˚55.43´N,141˚24.87´E 36˚54.05´N,141˚24.17´E
36˚56.28´N,141˚26.93´E 36˚54.64´N,141˚26.50´E
36˚29.90´N,140˚57.05´E 36˚31.28´N,141˚58.14´E
36˚29.03´N,140˚59.48´E 36˚30.51´N,141˚00.43´E
36˚28.11´N,140˚59.64´E 36˚29.16´N,141˚00.34´E
36˚29.47´N,141˚02.66´E 36˚30.30´N,141˚03.57´E
36˚30.63´N,141˚05.21´E 36˚31.41´N,141˚06.12´E
36˚29.09´N,141˚00.84´E 36˚30.04´N,141˚01.66´E
36˚30.84´N,141˚11.47´E 36˚31.63´N,141˚12.55´E
36˚33.63´N,141˚20.09´E 36˚34.12´N,141˚21.17´E
36˚30.87´N,141˚20.98´E
36˚56.76´N,141˚33.30´E 36˚58.10´N,141˚34.44´E
36˚58.17´N,141˚37.95´E 36˚59.05´N,141˚38.63´E
37˚37.54´N,141˚47.32´E 37˚39.17´N,141˚47.35´E
37˚38.58´N,141˚50.55´E 37˚40.28´N,141˚50.65´E
37˚43.09´N,141˚53.92´E 37˚44.84´N,141˚53.54´E
36˚53.21´N,141˚29.20´E 36˚52.11´N,141˚27.67´E
36˚51.57´N,141˚28.68´E 36˚52.75´N,141˚30.01´E
36˚51.57´N,141˚30.34´E 36˚52.36´N,141˚31.43´E
36˚48.39´N,141˚47.74´E
36˚46.16´N,141˚35.36´E 36˚45.58´N,141˚34.75´E
36˚49.87´N,141˚40.98´E 36˚49.31´N,141˚40.53´E
36˚41.87´N,141˚11.43´E 36˚40.51´N,141˚10.19´E
36˚40.11´N,141˚11.09´E 36˚41.48´N,141˚12.34´E
36˚40.31´N,141˚12.41´E 36˚41.70´N,141˚13.62´E
36˚39.69´N,141˚13.53´E 36˚40.97´N,141˚14.99´E
36˚40.28´N,141˚21.56´E 36˚41.31´N,141˚22.19´E
36˚41.06´N,141˚22.04´E
36˚40.37´N,141˚15.62´E 36˚38.96´N,141˚14.48´E
36˚38.97´N,141˚14.25´E
36˚39.47´N,141˚17.31´E 36˚40.89´N,141˚18.32´E
36˚41.55´N,141˚20.06´E 36˚40.16´N,141˚18.98´E
36˚40.79´N,141˚20.80´E 36˚42.31´N,141˚21.62´E
37˚19.91´N,141˚37.66´E
37˚20.32´N,141˚43.15´E 37˚22.02´N,141˚43.12´E
37˚18.91´N,141˚45.84´E 37˚17.29´N,141˚45.55´E
37˚18.83´N,141˚47.21´E 37˚20.47´N,141˚47.51´E
37˚18.13´N,141˚49.40´E 37˚16.55´N,141˚48.95´E

Depth (m)Date407 409424 425448 452482 483
514 505
1005 1004
545 561753 758
900 904
424 424
449 449484 480
508 506
1196 1196
900 904
151 150
211 210277 279299 314154 156
311 306352 352
450 457
507 510380 384
661 647748 758904 ND411 411560 557
355 351
387 379
411 411427 418454 448507 5091498 ND
750 750901 901
251 249278 278308 309344 351509 511512 508376 381378 373425 422454 452482 479
255 253352 346411 410450 446
480 480

2005/10/25
2005/10/25
2005/10/25
2005/10/25
2005/10/25
2005/10/26
2005/10/26
2005/10/26
2005/10/26
2005/10/27
2005/10/27
2005/10/27
2005/10/27
2005/10/28
2005/10/28
2005/10/29
2005/10/29
2005/10/29
2005/10/29
2005/10/30
2005/10/30
2005/10/31
2005/10/31
2005/10/31
2005/11/01
2005/11/02
2005/11/02
2005/11/02
2005/11/03
2005/11/03
2005/11/04
2005/11/04
2005/11/04
2005/11/09
2005/11/09
2005/11/09
2005/11/10
2005/11/10
2005/11/10
2005/11/11
2005/11/11
2005/11/11
2005/11/11
2005/11/11
2005/11/11
2005/11/12
2005/11/12
2005/11/13
2005/11/13
2005/11/13
2005/11/14
2005/11/14
2005/11/14
2005/11/14
2005/11/14

42

Minoru Imajima

Table 1. (Continued)
StationWA05-FG280
WA05-FG310
WA05-FG380
WA05-FG425
WA05-FG510D
WA05-EF480
WA05-EF510
WA05-EF250
WA05-EF250D
WA05-EF425
WA05-EF450D
WA05-DE250D
WA05-DE380D
WA05-DE510
WA06-C550
WA06-C750
WA06-C900D
WA06-A150
WA06-A150D
WA06-A250D
WA06-A450
WA06-A510
WA06-A350
WA06-A410
WA06-A650
WA06-A1200D
WA06-A1500D
WA06-A900
WA06-B450
WA06-B750D
WA06-B150
WA06-B310D
WA06-B350
WA06-C350D
WA06-D1500D
WA06-D450D
WA06-D750
WA06-D550
WA06-D210D
WA06-G150
WA06-G210
WA06-G280
WA06-G310
WA06-G350
WA06-G380
WA06-G425
WA06-G450
WA06-G480
WA06-G510
WA06-G550
WA06-G650
WA06-F150
WA06-F210
WA06-F250
WA06-F310

Position
37˚20.52´N,141˚39.15´E 37˚22.08´N,141˚39.19´E
37˚21.46´N,141˚41.16´E 37˚19.69´N,141˚41.20´E
37˚19.45´N,141˚44.62´E 37˚21.11´N,141˚44.79´E
37˚19.63´N,141˚46.54´E 37˚17.92´N,141˚46.24´E
37˚16.92´N,141˚50.04´E
38˚00.42´N,142˚05.19´E 37˚58.86´N,142˚04.06´E
38˚00.75´N,142˚05.72´E 38˚01.70´N,142˚06.34´E
37˚59.82´N,141˚50.54´E 38˚01.03´N,141˚51.69´E
37˚58.74´N,141˚49.30´E
37˚44.32´N,141˚54.82´E 38˚01.25´N,142˚03.82´E
38˚02.18´N,142˚04.82´E
38˚40.61´N,141˚55.34´E
38˚39.10´N,142˚02.22´E
38˚38.99´N,142˚07.27´E 38˚37.87´N,142˚07.20´E
39˚33.82´N,142˚18.45´E 39˚34.98´N,142˚18.54´E
39˚33.45´N,142˚22.26´E 39˚34.17´N,142˚22.50´E
39˚35.82´N,142˚32.11´E 39˚36.06´N,142˚32.30´E
40˚47.59´N,141˚49.39´E 40˚46.86´N,141˚51.13´E
40˚46.53´N,141˚51.87´E 40˚46.47´N,141˚52.17´E
40˚51.40´N,141˚50.94´E 40˚51.26´N,141˚51.14´E
40˚58.29´N,141˚45.93´E 40˚58.90´N,141˚45.38´E
41˚00.54´N,141˚46.18´E 41˚00.04´N,141˚46.93´E
40˚55.31´N,141˚42.87´E 40˚55.30´N,141˚43.83´E
40˚57.29´N,141˚43.51´E 40˚57.83´N,141˚43.04´E
41˚04.81´N,141˚48.98´E 41˚04.59´N,141˚49.15´E
40˚59.33´N,142˚12.73´E 40˚58.92´N,142˚12.86´E
40˚52.00´N,142˚33.35´E 40˚51.64´N,142˚33.78´E
41˚09.47´N,141˚53.69´E 41˚09.23´N,141˚53.75´E
40˚14.89´N,142˚15.33´E 40˚13.28´N,142˚16.08´E
40˚21.81´N,142˚20.22´E 40˚22.12´N,142˚20.02´E
40˚13.62´N,142˚07.10´E 40˚15.10´N,142˚06.62´E
40˚09.85´N,142˚13.24´E 40˚10.04´N,142˚13.21´E
40˚06.78´N,142˚14.98´E 40˚08.31´N,142˚14.64´E
39˚48.86´N,142˚17.11´E 39˚49.03´N,142˚17.23´E
39˚18.74´N,142˚40.18´E 39˚17.99´N,142˚40.08´E
39˚02.37´N,142˚10.53´E 39˚02.69´N,142˚10.57´E
38˚58.52´N,142˚16.33´E 38˚59.34´N,142˚16.47´E
39˚05.28´N,142˚12.76´E 39˚04.07´N,142˚12.68´E
38˚56.42´N,141˚59.28´E 38˚56.24´N,141˚59.17´E
36˚59.76´N,141˚17.40´E 37˚01.43´N,141˚17.85´E
36˚57.00´N,141˚22.70´E 36˚58.40´N,141˚23.78´E
36˚55.51´N,141˚24.91´E 36˚54.01´N,141˚24.15´E
36˚56.22´N,141˚26.93´E 36˚54.75´N,141˚26.52´E
36˚56.17´N,141˚30.80´E 36˚57.90´N,141˚31.47´E
36˚53.36´N,141˚27.44´E 36˚54.52´N,141˚28.91´E
36˚53.19´N,141˚29.24´E 36˚52.05´N,141˚27.59´E
36˚51.52´N,141˚28.64´E 36˚52.70´N,141˚30.04´E
36˚51.23´N,141˚29.16´E 36˚50.01´N,141˚27.74´E
36˚51.37´N,141˚30.11´E 36˚52.13´N,141˚31.16´E
36˚58.14´N,141˚37.96´E 36˚59.18´N,141˚38.79´E
36˚50.20´N,141˚34.27´E 36˚50.92´N,141˚35.18´E
37˚35.25´N,141˚33.18´E 37˚36.74´N,141˚33.89´E
37˚36.89´N,141˚35.86´E 37˚38.24´N,141˚36.13´E
37˚38.14´N,141˚39.05´E 37˚36.39´N,141˚38.84´E
37˚36.77´N,141˚43.66´E 37˚36.08´N,141˚43.61´E

Depth (m)Date

276 279
311 312383 383
426 426
516 515487 486
505 514252 251259 253433 418452 454249 249375 ND511 511
ND 558750 749891 893
155 149
146 147
267 266
466 474
511 510360 364
409 421663 6621202 1201
1513 1512
883 882
461 475
750 747
153 151
305 305
348 353357 364
1521 1518
460 460
749 745548 551
213 214151 150
210 208276 279
301 315373 355384 377428 420454 454481 483
508 508
558 554648 648
150 165
213 213257 256
313 309

2005/11/15
2005/11/15
2005/11/15
2005/11/15
2005/11/15
2005/11/16
2005/11/16
2005/11/17
2005/11/17
2005/11/18
2005/11/18
2005/11/19
2005/11/19
2005/11/21
2006/10/05
2006/10/05
2006/10/05
2006/10/09
2006/10/09
2006/10/10
2006/10/10
2006/10/10
2006/10/11
2006/10/11
2006/10/11
2006/10/12
2006/10/12
2006/10/12
2006/10/13
2006/10/13
2006/10/14
2006/10/14
2006/10/15
2006/10/15
2006/10/17
2006/10/17
2006/10/17
2006/10/18
2006/10/19
2006/10/26
2006/10/26
2006/10/26
2006/10/26
2006/10/27
2006/10/27
2006/10/27
2006/10/27
2006/10/28
2006/10/28
2006/10/28
2006/10/28
2006/10/29
2006/10/29
2006/10/29
2006/10/29

43

Deep-sea Benthic Polychaetes off Northern Honshu

Table 1. (Continued)
StationWA06-F350
WA06-F380
WA06-F410
WA06-F425
WA06-F450
WA06-F480
WA06-F510
WA06-F550
WA06-F650D
WA06-F1500D
WA06-F1500D(II)WA06-F750
WA06-F900
WA06-E1200D
WA06-E750
WA06-E900
WA06-E410
WA06-E450
WA06-E480
WA06-E510
WA06-E510D
WA06-E550
WA06-E310
WA06-E380
WA06-E425
WA06-E150
WA06-E210
WA06-FG425
WA06-FG450
WA06-FG480
WA06-G1200D
WA06-G900D
WA06-H250D
WA06-H310
WA06-H210
WA06-H450
WA06-H480
WA06-H550
WA06-H650
WA06-H750
WA06-G750
WA06-H1500D
WA06-GH380
WA06-GH450
WA06-GH510
WA06-GH425
WA06-GH480
WA06-GH480D
WA06-FG310
WA06-FG350
WA06-FG350D
WA06-FG380
WA06-EF280
WA06-EF425
WA06-EF425D

Position
37˚37.56´N,141˚47.32´E 37˚39.19´N,141˚47.42´E
37˚38.47´N,141˚50.54´E 37˚40.07´N,141˚50.61´E
37˚42.96´N,141˚53.93´E 37˚44.58´N,141˚53.56´E
37˚44.01´N,141˚54.82´E 37˚42.28´N,141˚55.05´E
37˚43.55´N,141˚56.56´E 37˚45.20´N,141˚56.40´E
37˚41.70´N,141˚58.98´E 37˚39.88´N,141˚59.02´E
37˚38.58´N,142˚01.11´E 37˚39.78´N,142˚01.38´E
37˚42.06´N,142˚04.07´E 37˚40.91´N,142˚04.73´E
37˚44.92´N,142˚08.51´E 37˚45.23´N,142˚08.37´E
37˚34.64´N,142˚33.47´E 37˚35.02´N,142˚33.50´E
37˚38.92´N,142˚34.09´E 37˚39.39´N,142˚34.32´E
37˚47.30´N,142˚12.20´E 37˚48.09´N,142˚11.96´E
37˚46.22´N,142˚18.93´E 37˚45.73´N,142˚19.10´E
38˚19.25´N,142˚31.74´E 38˚19.36´N,142˚31.65´E
38˚23.14´N,142˚14.52´E 38˚22.43´N,142˚14.10´E
38˚29.76´N,142˚21.63´E 38˚29.14´N,142˚21.50´E
38˚23.67´N,142˚02.55´E 38˚22.14´N,142˚03.27´E
38˚23.51´N,142˚03.99´E 38˚25.24´N,142˚03.61´E
38˚22.73´N,142˚05.21´E 38˚21.16´N,142˚05.89´E
38˚22.63´N,142˚06.29´E 38˚23.89´N,142˚05.73´E
38˚23.81´N,142˚05.57´E 38˚24.11´N,142˚05.42´E
38˚23.42´N,142˚07.32´E 38˚22.30´N,142˚07.32´E
38˚23.15´N,141˚58.17´E 38˚21.48´N,141˚58.44´E
38˚23.40´N,142˚01.57´E 38˚21.94´N,142˚02.33´E
38˚24.23´N,142˚02.95´E 38˚24.92´N,142˚02.74´E
38˚20.03´N,141˚44.46´E 38˚18.24´N,141˚44.07´E
38˚22.23´N,141˚51.30´E 38˚23.99´N,141˚51.55´E
37˚17.80´N,141˚46.21´E 37˚19.53´N,141˚46.55´E
37˚18.81´N,141˚47.19´E 37˚20.50´N,141˚47.50´E
37˚18.18´N,141˚49.47´E 37˚16.45´N,141˚48.83´E
36˚52.56´N,141˚48.58´E 36˚52.25´N,141˚48.23´E
36˚47.53´N,141˚39.39´E 36˚47.32´N,141˚39.14´E
36˚30.85´N,140˚59.58´E 36˚31.09´N,140˚59.78´E
36˚30.57´N,141˚00.56´E 36˚29.07´N,140˚59.52´E
36˚30.01´N,140˚58.35´E 36˚31.42´N,140˚59.00´E
36˚30.44´N,141˚03.63´E 36˚29.04´N,141˚02.32´E
36˚32.12´N,141˚05.99´E 36˚32.74´N,141˚06.90´E
36˚31.77´N,141˚08.69´E 36˚32.62´N,141˚09.78´E
36˚30.98´N,141˚11.69´E 36˚31.68´N,141˚12.69´E
36˚34.07´N,141˚20.52´E 36˚34.59´N,141˚21.26´E
36˚46.26´N,141˚35.48´E 36˚46.41´N,141˚35.68´E
36˚36.45´N,141˚36.21´E 36˚36.66´N,141˚36.14´E
36˚40.37´N,141˚15.64´E 36˚38.90´N,141˚14.50´E
36˚41.63´N,141˚20.13´E 36˚40.25´N,141˚19.03´E
36˚40.26´N,141˚21.57´E 36˚41.23´N,141˚22.16´E
36˚39.39´N,141˚17.30´E 36˚40.87´N,141˚18.31´E
36˚40.68´N,141˚20.77´E‒36˚42.18´N,141˚21.55´E
36˚40.01´N,141˚20.25´E‒36˚39.82´N,141˚19.98´E
37˚21.64´N,141˚41.16´E‒37˚19.80´N,141˚41.20´E
37˚22.16´N,141˚43.12´E‒37˚20.39´N,141˚43.12´E
37˚22.78´N,141˚43.20´E‒37˚23.20´N,141˚43.22´E
37˚19.53´N,141˚44.62´E‒37˚21.14´N,141˚44.77´E
38˚02.72´N,141˚56.49´E‒38˚04.51´N,141˚55.98´E
38˚00.89´N,142˚03.75´E‒38˚02.72´N,142˚04.13´E
38˚03.26´N,142˚03.97´E‒38˚03.12´N,142˚04.05´E

Depth (m)Date353 350386 379
411 411425 424450 450483 478503 511546 551647 641
1511 1508
1466 1471
749 747904 909
1214 1213
758 756905 908406 409448 451
480 484
514 506503 498
553 545305 309377 382423 423
154 151
209 212426 425449 444480 477
1201 1182
925 920248 248309 310213 193456 447481 480
561 557
659 646736 732
753 7541470 1450
377 381453 450509 510425‒422481‒478483‒478312‒313346‒351346‒346382‒382283‒277414‒431420‒424

2006/10/30
2006/10/30
2006/10/30
2006/10/30
2006/10/30
2006/10/31
2006/10/31
2006/10/31
2006/10/31
2006/11/01
2006/11/01
2006/11/01
2006/11/01
2006/11/02
2006/11/02
2006/11/02
2006/11/03
2006/11/03
2006/11/03
2006/11/03
2006/11/03
2006/11/03
2006/11/04
2006/11/04
2006/11/04
2006/11/05
2006/11/05
2006/11/10
2006/11/10
2006/11/10
2006/11/11
2006/11/11
2006/11/12
2006/11/12
2006/11/13
2006/11/13
2006/11/13
2006/11/14
2006/11/14
2006/11/14
2006/11/15
2006/11/15
2006/11/16
2006/11/16
2006/11/16
2006/11/18
2006/11/18
2006/11/18
2006/11/19
2006/11/19
2006/11/19
2006/11/19
2006/11/21
2006/11/21
2006/11/21

44

Minoru Imajima

Table 1. (Continued)
StationWA06-EF350
WA06-EF480
WA06-EF510
WA06-DE280D
WA06-DE410
WA06-DE450
WA07-D550
WA07-D750
WA07-D900
WA07-A250
WA07-A250D
WA07-A310
WA07-A350
WA07-A410
WA07-A450
WA07-A510
WA07-A550
WA07-A650
WA07-A750
WA07-A1500D
WA07-B750
WA07-B1200
WA07-B1500D
WA07-B450
WA07-B510
WA07-B310
WA07-B350
WA07-B410
WA07-B410D
WA07-B150
WA07-C310
WA07-C410
WA07-C510
WA07-C250
WA07-C350
WA07-C350D
WA07-C1500D
WA07-C550
WA07-C650
WA07-C750
WA07-C900
WA07-C450
WA07-D1500D
WA07-D450
WA07-D510
WA07-D210D
WA07-D310
WA07-D350
SO06-M1-B
SO06-M3-B1
SO06-M3-B2
SO06-M4-B
SO07-C7-B
SO07-K1
SO07-K2

Position
38˚00.20´N,141˚59.86´E‒37˚58.68´N,141˚59.32´E
38˚00.45´N,142˚05.23´E‒37˚59.23´N,142˚04.33´E
38˚00.74´N,142˚05.75´E‒38˚01.68´N,142˚06.61´E
38˚42.91´N,141˚58.34´E‒38˚43.12´N,141˚58.37´E
38˚39.29´N,142˚03.42´E‒38˚40.92´N,142˚03.70´E
38˚37.67´N,142˚04.48´E‒38˚39.15´N,142˚04.74´E
39˚03.74´N,142˚12.75´E‒39˚04.91´N,142˚12.69´E
38˚58.68´N,142˚16.04´E‒38˚59.48´N,142˚16.58´E
39˚05.27´N,142˚20.04´E‒39˚06.00´N,142˚20.07´E
40˚51.81´N,141˚50.57´E‒40˚50.52´N,141˚51.90´E
40˚50.96´N,141˚51.23´E‒40˚50.88´N,141˚51.45´E
40˚49.39´N,141˚55.02´E‒40˚50.59´N,141˚53.54´E
40˚55.26´N,141˚43.22´E‒40˚55.02´N,141˚44.66´E
40˚57.91´N,141˚42.48´E‒40˚57.46´N,141˚43.33´E
40˚58.71´N,141˚45.57´E‒40˚58.03´N,141˚46.08´E
41˚00.58´N,141˚46.13´E‒41˚00.42´N,141˚46.39´E
41˚00.85´N,141˚48.68´E‒41˚00.59´N,141˚48.98´E
41˚04.91´N,141˚48.91´E‒41˚04.05´N,141˚49.20´E
41˚07.61´N,141˚50.02´E‒41˚07.38´N,141˚50.10´E
40˚50.46´N,142˚31.48´E‒40˚50.16´N,142˚31.06´E
40˚19.67´N,142˚21.32´E‒40˚20.14´N,142˚20.88´E
40˚21.61´N,142˚33.94´E‒40˚21.84´N,142˚33.75´E
40˚23.90´N,142˚48.52´E‒40˚23.93´N,142˚48.18´E
40˚13.15´N,142˚15.68´E‒40˚14.74´N,142˚15.35´E
40˚16.03´N,142˚16.01´E‒40˚17.03´N,142˚15.58´E
40˚13.42´N,142˚12.38´E‒40˚11.66´N,142˚12.78´E
40˚06.39´N,142˚15.08´E‒40˚08.17´N,142˚14.64´E
40˚15.44´N,142˚14.12´E‒40˚13.65´N,142˚14.55´E
40˚16.88´N,142˚13.51´E‒40˚17.11´N,142˚13.46´E
40˚14.98´N,142˚06.64´E‒40˚13.31´N,142˚07.38´E
39˚47.25´N,142˚16.41´E‒39˚45.56´N,142˚16.01´E
39˚50.27´N,142˚17.92´E‒39˚48.45´N,142˚17.94´E
39˚52.47´N,142˚19.83´E‒39˚51.20´N,142˚20.02´E
39˚40.02´N,142˚14.27´E‒39˚41.66´N,142˚14.48´E
39˚45.66´N,142˚16.90´E‒39˚47.41´N,142˚16.96´E
39˚44.16´N,142˚16.90´E‒39˚44.40´N,142˚16.86´E
39˚33.44´N,142˚51.27´E‒39˚33.61´N,142˚53.31´E
39˚35.51´N,142˚18.58´E‒39˚34.21´N,142˚18.48´E
39˚34.32´N,142˚20.32´E‒39˚35.48´N,142˚20.30´E
39˚34.05´N,142˚22.48´E‒39˚33.53´N,142˚22.32´E
39˚36.07´N,142˚32.68´E‒39˚35.90´N,142˚32.46´E
39˚42.33´N,142˚18.03´E‒39˚40.62´N,142˚17.72´E
39˚20.17´N,142˚40.07´E‒39˚20.46´N,142˚40.25´E
39˚03.41´N,142˚10.43´E‒39˚01.49´N,142˚10.54´E
39˚04.18´N,142˚11.75´E‒39˚05.33´N,142˚12.01´E
38˚57.42´N,141˚59.65´E‒38˚57.74´N,141˚59.85´E
38˚53.52´N,142˚02.78´E‒38˚54.97´N,142˚03.33´E
38˚55.08´N,142˚05.70´E‒38˚53.52´N,142˚05.16´E
39˚40.4´N,143˚11.3´E
39˚40.8´N,143˚50.2´E
39˚45.7´N,143˚53.4´E
39˚35.2´N,144˚02.4´E
39˚40.0´N,142˚34.0´E‒39˚40.0´N,142˚33.9´E
38˚34.5´N,143˚04.2´E‒38˚33.2´N,143˚04.1´E
38˚31.2´N,143˚33.6´E‒38˚31.1´N,143˚34.2´E

Depth (m)Date356‒357486‒482504‒531284‒285408‒408450‒446556‒545754‒751898‒905273‒258258‒258306‒309360‒359412‒415471‒468510‒512550‒551662‒661748‒7471402‒1377
759‒7491208‒1200
1511‒1514
454‒459510‒509309‒307350‒352420‒412416‒416153‒156318‒294409‒415511‒521254‒252358‒358355‒3541499‒1480
552‒559659‒644748‒749900‒893467‒4581505‒1489
448‒463505‒513213‒213303‒307354‒351
2018
3937
3960
4951
820‒8162043‒2081
2968‒3032

2006/11/22
2006/11/22
2006/11/22
2006/11/23
2006/11/24
2006/11/24
2007/10/05
2007/10/05
2007/10/05
2007/10/06
2007/10/06
2007/10/06
2007/10/07
2007/10/09
2007/10/09
2007/10/09
2007/10/09
2007/10/10
2007/10/10
2007/10/11
2007/10/11
2007/10/12
2007/10/12
2007/10/12
2007/10/12
2007/10/13
2007/10/13
2007/10/13
2007/10/13
2007/10/14
2007/10/14
2007/10/14
2007/10/14
2007/10/15
2007/10/15
2007/10/15
2007/10/16
2007/10/16
2007/10/16
2007/10/16
2007/10/16
2007/10/17
2007/10/17
2007/10/17
2007/10/17
2007/10/18
2007/10/18
2007/10/18
2006/07/16
2006/07/17
2006/07/17
2006/07/17
2007/08/05
2007/08/06
2007/08/06

45

Deep-sea Benthic Polychaetes off Northern Honshu

Table 1. (Continued)
StationSO07-K3
SO07-O4
SO07-O1
SO07-O2
SO07-O3
KT-07-29-M-1
KT-07-29-M-2
KT-07-29-M-3-1
KT-07-29-M-3-2
KT-07-29-H-1
KT-07-29-H-2
KT-07-29-M-3-3

Position38˚33.7´N,143˚41.2´E‒38˚33.2´N,143˚41.5´E
36˚54.0´N,142˚56.6´E‒36˚55.5´N,142˚57.3´E
36˚43.9´N,141˚49.1´E‒36˚44.0´N,141˚48.9´E
36˚48.3´N,142˚22.3´E‒36˚48.7´N,142˚22.1´E
36˚53.4´N,142˚37.0´E‒36˚53.6´N,142˚36.8´E
39˚17.86´N,142˚28.40´E‒39˚16.80´N,142˚27.37´E
39˚16.20´N,142˚41.07´E‒39˚18.62´N,142˚43.73´E
39˚20.04´N,142˚50.99´E‒39˚21.75´N,142˚51.90´E
39˚20.19´N,142˚51.40´E‒39˚19.23´N,142˚49.18´E
40˚48.70´N,142˚00.05´E‒40˚47.37´N,142˚00.49´E
40˚00.00´N,143˚31.37´E‒41˚00.76´N,143˚30.25´E
39˚20.09´N,142˚51.17´E‒39˚19.23´N,142˚49.05´E

Depth (m)Date4105‒4181
5219‒5268
2024‒2020
2991‒2948
4128‒4094
1039‒1041
1528‒1603
1728‒17191737‒1709
497‒4542055‒2032
1733‒1695

2007/08/07
2007/08/07
2007/08/08
2007/08/08
2007/08/08
2007/11/05
2007/11/05
2007/11/05
2007/11/06
2007/11/08
2007/11/08
2007/11/08(Scalibregmatidae) and Anobothrus wakatakamaruae (Ampharetidae) are new to science. Thirtyseven species, Bylgides macrolepida, Eunoe depressa, Gattyana ciliata, Harmothoe grandispina
and Parahalosydna krassini (Polynoidae), Glycera branchiopoda (Glyceridae), Goniada brunnea(Goniadidae), Ceratocephala borealis (Nereididae), Drilonereis filum (Oenonidae), Phylo felix,
Scoloplos (Leodamas) robustus and Scoloplos (Scoloplos) similis (Orbiniidae), Acrocirrus heterochaetus (Acrocirridae), Fauveliopsis glabra (Fauveliopsidae), Asychis auritus (Maldanidae), Ophelina acuminata and Travisia pupa (Opheliidae), Asclerocheilus beringianus (Scalibregmatidae),Myriochele olgae (Oweniidae), Amage delus, Ampharete longipaleolata, Amphicteis mederi,
Amphicteis scaphobranchiata, Anobothrus gracilis, Lysippe labiata, Melinnopsis atlantica,
Paramage scutata, Samythella bathycola and Samythella neglecta (Ampharetidae), Trichobranchus glacialis (Trichobranchidae), Artacama proboscidea, Nicolea zostericola, Scionella vinogradovi and Streblosoma bairdi (Terebellidae), Chone gracilis, Chone magna and Euchone
analis (Sabellidae) are newly added to the Japanese polychaetous fauna. Eighty-six species are
known also from the northwestern Pacific Ocean, including Sea of Okhotsk and Bering Sea. Forty
species are known only from Japan, or endemic (see Table 2). The Fauveliopsidae is newly added
to the Japanese fauna.
Thirty-two indeterminable species could not be identified to species due to fragments, damaged or juvenile individuals. The type specimens and other specimens collected have been deposited at the National Museum of Nature and Science, Tokyo (NSMT).
Description of Species
Order Phyllodocida
Family Aphroditidae Malmgren, 1867
Genus Aphrodita Linnaeus, 1758
Aphrodita aculeata Linnaeus, 1758
Aphrodita aculeata Linnaeus, 1758: 655; Chambers and Muir, 1997: 60 61, fig. 7; Barnich and Fiege, 2000: 132 136, figs.
1 2; Imajima, 2003: 6 10, figs. 3a g, 4a l, 5a h.

Material. NSMT-Pol. 109494, WA05-F900 (2 specimens), NSMT-Pol. 109495, WA05-G450(1), NSMT-Pol. 109496, WA05-GH380D (1), NSMT-Pol. 109497, WA05-GH450 (1), NSMT-Pol.
109498, WA05-H380 (1); NSMT-Pol. 109499, WA06-A1200D (11), NSMT-Pol. 109500,

46

Minoru Imajima

WA06-E900 (1), NSMT-Pol. 109501, WA06-E1200D (1); NSMT-Pol. 110153, WA07-B1200 (4),
NSMT-Pol. 110154, WA07-C1500D (1), NSMT-Pol. 110155, WA07-D900 (1).
Distribution. North Atlantic Ocean; Mediterranean Sea; Indian Sea; Japan.
Aphrodita goolmarris Hutchings and McRae, 1993
Aphrodita goolmarris Hutchings and McRae, 1993: 295 298, figs. 9, 10a f, 11a j, 59c, Tables 1, 3; Imajima, 2003: 10 14, figs.
6a n, 7a q.

Material. NSMT-Pol. 109502, WA05-F900 (1); NSMT-Pol. 109503, WA06-E380 (1), NSMT-Pol. 109504, WA06-FG380 (1).
Distribution. Australia; Japan.
Aphrodita negligens Moore, 1905
Aphrodita negligens Moore, 1905: 526 529, pl. 34, figs. 1 2, pl. 35, fig. 31; Pettibone, 1953: 70 72, pl. 34, figs. 308 316, pl.
35, figs. 317 324; Imajima and Hartman, 1964: 14; Imajima, 2003: 17 20, figs. 10a i, 11a n.

Material. NSMT-Pol. 109505, WA06-A150D (1), NSMT-Pol. 109506 , WA06-EF350 (1).
Distribution. Western Canada; Japan.
Aphrodita nipponensis Imajima, 2003
Aphrodita nipponensis Imajima, 2003: 20 23, figs. 12a o, 13a m.

Material. NSMT-Pol. 109507, WA05-E900 (2), NSMT-Pol. 109510, WA05-EF450D (1),
NSMT-Pol. 109508, WA05-F425 (4), NSMT-Pol. 109509, WA05-H900D (1); NSMT-Pol. 109511,
WA06-B450 (1), NSMT-Pol. 109512, WA06-E310 (1), NSMT-Pol. 109513, WA06-E425 (1), NSMT-Pol. 109514, WA06-E510D (1), NSMT-Pol. 109516, WA06-EF350 (1), NSMT-Pol. 109515,
WA06-EF480 (1), NSMT-Pol. 109517, WA06-F425 (1).
Distribution. Japan.
Aphrodita sibogae (Horst, 1916)
Aphroditella sibogae Horst, 1916: 66.
Aphrodita sibogae: Hutchings and McRae, 1993: 307, fig. 60, tab. 1; Imajima, 1997a: 152 153, fig. 2a t; Imajima, 2003:
23 26.

Material. NSMT-Pol. 109468, WA05-DE380D (8), NSMT-Pol. 109463, WA05-E450 (1),
NSMT-Pol. 109467, WA05-E480 (2), NSMT-Pol. 109469, WA05-EF250 (1), NSMT-Pol. 109470,
WA05-EF250D (5), NSMT-Pol. 109471, WA05-EF480 (3), NSMT-Pol. 109472, WA05-F380 (1),
NSMT-Pol. 109487, WA05-F450 (3), NSMT-Pol. 109466, WA05-F480 (1), NSMT-Pol. 109473,
WA05-FG350 (1), NSMT-Pol. 109464, WA05-FG450 (2), NSMT-Pol. 109465, WA05-G510 (1),
NSMT-Pol. 109474, WA05-GH380D (1); NSMT-Pol. 109482, WA06-A250D (1), NSMT-Pol.
109475, WA06-A410 (3), NSMT-Pol. 109483, WA06-A450 (4), NSMT-Pol. 109476, WA06-A510(5), NSMT-Pol. 109484, WA06-B450 (4), NSMT-Pol. 109486, WA06-D210D (2), NSMT-Pol.
109477, WA06-D450D (3), NSMT-Pol. 109478, WA06-DE410 (2), NSMT-Pol. 109479, WA06E550 (2), NSMT-Pol. 109485, WA06-E1200D (1), NSMT-Pol. 109480, WA06-EF510 (1), NSMTPol. 109481, WA06-F410 (1); NSMT-Pol. 110143, WA07-A250D (2), NSMT-Pol. 110142,
WA07-A310 (2), NSMT-Pol. 110144, WA07-A350 (4), NSMT-Pol. 110185, WA07-A510 (2),Deep-sea Benthic Polychaetes off Northern Honshu

47

NSMT-Pol. 110145, WA07-B310 (1), NSMT-Pol. 110146, WA07-B350 (1), NSMT-Pol. 110186,
WA07-B410D (45), NSMT-Pol. 110187, WA07-B450 (3), NSMT-Pol. 110140, WA07-B510D(2), NSMT-Pol. 110147, WA07-C250 (1), NSMT-Pol. 110148, WA07-C310 (1), NSMT-Pol.
110149, WA07-C350 (2), NSMT-Pol. 110188, WA07-C350D (21), NSMT-Pol. 110150, WA07C410 (3), NSMT-Pol. 110141, WA07-C450 (3), NSMT-Pol. 110151, WA07-C510 (1), NSMT-Pol.
110152, WA07-D210D (1), NSMT-Pol. 110189, WA07-D550 (1). NSMT-Pol. 110614, KT0729-H-1 (15).
Distribution. West of Salawatti, Indonesian Archipelago; Japan.
Aphrodita talpa Quatrefages, 1865
Aphrodite talpa Quatrefages, 1865: 196 197, pl. 6, figs. 2 4.
Aphrodita talpa: Fauvel, 1925: 140 144, fig. 4; Monro, 1936: 82 83, fig. 3; Hutchings and McRae, 1993: 307 311, figs. 23a f,
24a k, 59c, tab. 1; Imajima, 2007: 25 26, fig. 9.

Material. NSMT-Pol. 109488, WA05-E410 (1), NSMT-Pol. 109489, WA05-F425 (4), NSMT-Pol. 109490, WA05-G510 (1); NSMT-Pol. 109491, WA06-E510D (1), NSMT-Pol. 109492,
WA06-EF280 (1), NSMT-Pol. 109493, WA06-FG450 (1).
Distribution. Australia; New Zealand; Far Eastern seas; Bering Sea; Japan.
Genus Laetmonice Kinberg, 1856
Laetmonice japonica McIntosh, 1885
Laetmonice japonica McIntosh, 1885: 50 51, pl. 8, fig. 1, pl. 4A, fig. 13, pl. 5A, figs. 9 10; Imajima, 2003: 26 29, figs. 14a h,
15a k; Imajima, 2005: 55.
Laetmatonice japonica: Moore, 1903: 420; Izuka, 1912: 80 82, pl. 1, fig. 4, pl. 9, figs. 14 15; Fauvel, 1936a: 48; Uschakov
and Wu, 1962a: 112.

Material. NSMT-Pol. 109442, WA05-G310 (1), NSMT-Pol. 109446, WA05-G425 (2), NSMT-Pol. 109443, WA05-G450 (6), NSMT-Pol. 109440, WA05-GH350 (6), NSMT-Pol. 109441,
WA05-GH380 (7), NSMT-Pol. 109447, WA05-GH425 (1), NSMT-Pol. 109444, WA05-GH450(5), NSMT-Pol. 109448, WA05-GH480 (16), NSMT-Pol. 109445, WA05-H380 (3); NSMT-Pol.
109450, WA06-G350 (3), NSMT-Pol. 109451, WA06-G425 (2), NSMT-Pol. 109453, WA06-G450(1), NSMT-Pol. 109452, WA06-G480 (1), NSMT-Pol. 109767, WA06-G425 (1), NSMT-Pol.
109449, WA06-GH480 (4); NSMT-Pol. 110125, WA07-B1200 (5). NSMT-Pol. 110570, SO06M3-B2 (4).
Distribution. Yellow Sea; Japan.
Laetmonice producta Grube, 1877
Laetmonice producta Grube, 1877: 512 513; McIntosh, 1885: 39 44, pl. 6, figs. 1 2, pl. 4A, figs. 1 8; Horst, 1917: 54 56, pl.
13, figs. 1 3; Uschakov, 1965: 105; Hutchings and McRae, 1993: 333 336, figs. 45a f, 46a j, 59E, tab. 6, 8 9; Imajima,
2003: 30 31, fig. 16a s.

Material. NSMT-Pol. 109410, WA05-G150 (1), NSMT-Pol. 109411, WA05-G210 (2), NSMT-Pol. 109412, WA05-H150 (1); NSMT-Pol. 109458, WA06-A150 (8), NSMT-Pol. 109454,
WA06-A150D (2), NSMT-Pol. 109462, WA06-B150 (5), NSMT-Pol. 109459, WA06-E150 (3),
NSMT-Pol. 109460, WA06-E210 (1), NSMT-Pol. 109461, WA06-F150 (2), NSMT-Pol. 109457,
WA06-G150 (3), NSMT-Pol. 109455, WA06-G210 (1), NSMT-Pol. 109456, WA06-H210 (3);
NSMT-Pol. 110126, WA07-B150 (2), NSMT-Pol. 110127, WA07-C510 (1).

48

Minoru Imajima

Distribution. Australia; off Kerguelen; Japan.
Laetmonice sp.
Material. NSMT-Pol. 110890, SO07-O4 (2).
Family Polynoidae Malmgren, 1867
Subfamily Harmothoinae Horst, 1917
Genus Bylgides Chamberlin, 1919
Bylgides macrolepida (Moore, 1905)
Antinoe macrolepida Moore, 1905: 538, pl. 35, figs. 21 23; Berkeley and Berkeley, 1948: 14, fig. 15; Hartman, 1948a: 6.
Antinoella macrolepida: Uschakov, 1958: 78, fig. 1A E; Hartman, 1968: 41, figs. 1 3.
Bylgides macrolepida: Uschakov, 1982: 157 158, pl. 57, 1 6.

Material. NSMT-Pol. 110791, WA06-H480 (1); NSMT-Pol. 110792, WA07-A310 (1), NSMT-Pol. 110793, WA07-A350 (3), NSMT-Pol. 110794, WA07-B310 (1), NSMT-Pol. 110795,
WA07-B410 (1), NSMT-Pol. 110796, WA07-B450 (1), NSMT-Pol. 110797, WA07-C310 (1), NSMT-Pol. 110798, WA07-C350D (7), NSMT-Pol. 110799, WA07-C410 (1), NSMT-Pol. 110800,
WA07-D550 (1).
Description. Largest specimen 35 mm long, 17 mm wide including setae for 39 segments.
Prostomium short, broad, deeply cleft; lobes broadly rounded anteriorly. Anterior eyes very large,
occupying almost all anterior part of prostomium, posterior eyes very small at posterior end of
lobe. Ceratophore of median antenna inserted in anterior notch, style long, extending distally nearly to ends of palps. Lateral antennae small, inserted ventrally on underside of prostomium.
Elytra 15 pairs, subcircular to reniform, covered with small, slightly curved, conical spines
and a few slender clavate cilia. Parapodia biramous; notopodia small, neuropodia prominent,
obliquely truncate, terminate in a slender acicular process. Notosetae few, each thick, nearlystraight, with rows of spinelets. Neurosetae very numerous, long and slender; each slightly curved
distally and distal region with rows of fine hairs throughout its length and with a long terminal
bristle.
The genus and species are newly added to the Japanese polychaetous fauna.
Distribution. Southern Sakhalin; American coast from Alaska to California; Monterey Bay;
Japan.
Genus Eunoe Malmgren, 1865
Eunoe barbata Moore, 1910
Eunoe barbata Moore, 1910: 334 338, pl. 28, figs. 1 6; Imajima and Hartman, 1964: 29 30, pl. 2, figs. a f.

Material. NSMT-Pol. 109518, WA05-EF250 (4), NSMT-Pol. 109519, WA05-EF425 (1),
NSMT-Pol. 109520, WA05-F380 (1), NSMT-Pol. 109521, WA05-FG310 (1), NSMT-Pol. 109522,
WA05-FG380 (1), NSMT-Pol. 109523, WA05-FG480 (1), NSMT-Pol. 109524, WA05-G310 (8),NSMT-Pol. 109525, WA05-GH310 (6), NSMT-Pol. 109526, WA05-GH350 (2), NSMT-Pol.
109527, WA05-GH510 (1); NSMT-Pol. 109528, WA06-A250D (2), NSMT-Pol. 109529, WA06A410 (1), NSMT-Pol. 109530, WA06-A510 (1), NSMT-Pol. 199531, WA06-D450D (1), NSMTPol. 109532, WA06-D550 (1), NSMT-Pol. 109533, WA06-DE410 (1), NSMT-Pol. 109534, WA06DE450 (1), NSMT-Pol. 109535, WA06-E210 (2), NSMT-Pol. 109536, WA06-E380 (1), NSMT-Pol.
110313, WA06-F210 (1), NSMT-Pol. 109537, WA06-F250 (1), NSMT-Pol. 109538, WA06-F350

Deep-sea Benthic Polychaetes off Northern Honshu

49(3), NSMT-Pol. 110314, WA06-F380 (1), NSMT-Pol. 110315, WA06-F410 (1), NSMT-Pol.
109539, WA06-F425 (1), NSMT-Pol. 109540, WA06-FG310 (1), NSMT-Pol. 110316, WA06FG350 (1), NSMT-Pol. 110317, WA06-FG450 (1), NSMT-Pol. 109541, WA06-FG480 (1), NSMT-Pol. 110318, WA06-G280 (1), NSMT-Pol. 110319, WA06-G310 (2), NSMT-Pol. 109542,
WA06-G380 (3); NSMT-Pol. 110066, WA07-A310 (2), NSMT-Pol. 110067, WA07-A350 (1),
NSMT-Pol. 110068, WA07-A510 (1), NSMT-Pol. 110069, WA07-B410 (3), NSMT-Pol. 110128,
WA07-B450 (1), NSMT-Pol. 110070, WA07-B510 (1), NSMT-Pol. 110129, WA07-C310 (1), NSMT-Pol. 110130, WA07-C410 (1), NSMT-Pol. 110131, WA07-C450 (5), NSMT-Pol. 110132,
WA07-D310 (1).
Distribution. California; Alaska; Japan.
Eunoe depressa Moore, 1905
(Fig. 2A K)
Eunoe depressa Moore, 1905: 536 538, pl. 34, figs. 17 18, pl. 35, figs. 19 20; Uschakov, 1965: 128, fig. 33G I.

Material. NSMT-Pol. 110326, WA06-E310 (1), NSMT-Pol. 110327, WA06-E750 (5), NSMT-Pol. 110328, WA06-F350 (1), NSMT-Pol. 110329, WA06-F425 (1), NSMT-Pol. 110330,
WA06-F480 (1), NSMT-Pol. 110331, WA06-F510 (2), NSMT-Pol. 110332, WA06-G280 (2), NSMT-Pol. 110333, WA06-G425 (1), NSMT-Pol. 110334, WA06-H450 (1).
Description. Largest entire specimen 42 mm long, 15 mm wide including setae for 40 segments. Body dosally flattened, tapering slightly posteriorly. Dorsum with 2 transverse ciliatedbands per segment.
Prostomium bilobed, wider than long, with distinct cephalic peaks. Two pairs of eyes, anterior pair circular, lying dorsolaterally at widest part of prostomium, posterior pair lying dorsallynear posterior margin of prostomium, closer to midline. Ceratophore of median antenna large,
cylindrical, inserted in anterior notch, style long, slender, gradually tapering to filiform tip. Lateralantennae with small ceratophores, inserted ventrally on underside of prostomium; styles short,
slender, about half as long as median antenna. Palps thick, gently tapering to tips (Fig. 2A).
Elytra 15 pairs, on segments 2, 4, 5, 7, alternate segments to 21, 23, 26, 29 and 32, covering
dorsum completely and overlapping medially and posteriorly. Elytra reniform, without fringe papillae (Fig. 2B); greater part of dorsal surface covered with numerous minute conical microtubercles (Fig. 2C, E) and sparse conical macrotubercles (Fig. 2C D), and clavate papillae scattered on
surface.
Tentacular segment not visible dorsally; tentaculophores lateral to prostomium, with 4 5 peristomial setae on each inner side; dorsal and ventral tentacular cirri tapering gradually to subterminal inflation and filiform tips. Segment 2 with first pair of elytra, biramous parapodia, and papillated buccal cirri considerably longer than following ventral cirri.
Parapodia biramous, similar along length of body. Notopodium smaller than neuropodium,
conical with long acicular lobe. Neuropodium with subtriangular presetal acicular lobe with supraacicular digitiform process from which aciculum projects; postsetal lobe shorter, distally triangular. Dorsal cirri with large, cylindrical cirrophores on posterior sides of notopodia; styles long, tapering to slight subterminal inflation and filiform tip, with few papillae. Ventral cirri shorter thanneuropodium, cirriform, sparsely covered with minute papillae (Fig. 2F). Nephridial papillae short,from 5th parapodium onwards.
Noto- and neurosetae with entire tips. Notosetae thicker than neurosetae, slightly curved;
outer ones in palisade, short, with spinous rows of fine serrations along convex edge (Fig. 2G);
inner ones much longer, with numerous spinous rows of serrations, below bare, tapered tips (Fig.

50

Minoru Imajima

Fig. 2. Eunoe depressa Moore —A, anterior end, dorsal view, × 17; B, median elytron, × 13: C, macro- and microtubercles on elytral surface, × 50; D, macrotubercle, × 130; E, microtubercle, × 130; F, right cirrigerous
parapodium from segment 16, anterior view, × 13; G, short, outer notoseta, × 63; H, long, inner notoseta, × 63,
with detail of median part, × 160; I, upper neuroseta, × 102, with detail of tip, × 206; J, lower neuroseta, × 102;
K, distal region of same, × 206.

2H). Neurosetae arranged in transverse rows; upper ones with spinous rows of strong serrations
below distally bent entire tips (Fig. 2I); lower ones shorter, with few spinous rows of serrations(Fig. 2J K). Pygidium with pair of long anal cirri on short cirrophores.
The species is newly added to the Japanese polychaetous fauna.
Distribution. Gulf of Alaska; northern part of Sea of Japan; Bering Sea; Sea of Okhotsk; Japan.

Deep-sea Benthic Polychaetes off Northern Honshu

51

Eunoe shirikishinai Imajima and Hartman, 1964
Eunoe shirikishinai Imajima and Hartman, 1964: 30 31, pl. 3, figs. a d.

Material. NSMT-Pol. 110133, WA07-B150 (1).
Distribution. Japan.
Genus Gattyana McIntosh, 1900
Gattyana ciliata Moore, 1902
(Figs. 3A J, 4A F)
Gattyana ciliata Moore, 1902: 263 266, pl. 13, figs. 14 19, pl. 14, fig. 20; Annenkova, 1937: 148, pl. 2, figs. 9, 11, pl. 4, fig.
33; Uschakov, 1965: 124, fig. 32A C.

Material. NSMT-Pol. 109543, WA05-DE250D (31), NSMT-Pol. 109544, WA05-DE380D(22), NSMT-Pol. 109545, WA05-E425 (1), NSMT-Pol. 109546, WA05-EF250D (7), NSMT-Pol.
109547, WA05-F350 (1), NSMT-Pol. 109548, WA05-F410 (2), NSMT-Pol. 109549, WA05-F450(2), NSMT-Pol. 109550, WA05-FG250D (1), NSMT-Pol. 109551, WA05-FG350 (2), NSMT-Pol.
109552, WA05-FG410 (1), NSMT-Pol. 109553, WA05-FG425 (1), NSMT-Pol. 109554, WA05FG450 (2), NSMT-Pol. 109555, WA05-GH380 (2), NSMT-Pol. 109556, WA05-GH380D (1),
NSMT-Pol. 109557, WA05-GH425 (1), NSMT-Pol. 109558, WA05-GH480 (1), NSMT-Pol.
109559, WA05-H380 (1); NSMT-Pol. 109560, WA06-A250D (7), NSMT-Pol. 109561, WA06B310D (7), NSMT-Pol. 109562, WA06-C350D (10), NSMT-Pol. 109563, WA06-D210D (1), NSMT-Pol. 109564, WA06-EF425D (1), NSMT-Pol. 109565, WA06-FG350D (3); NSMT-Pol.
110134, WA07-A250D (9), NSMT-Pol. 110135, WA07-B410D (1), NSMT-Pol. 110136, WA07C350 (1), NSMT-Pol. 110233, WA07-C350D (23). NSMT-Pol. 110615, KT07-29-H-1 (1).
Description. Largest entire specimen 40 mm long, 16 mm wide including setae for 36 segments. Body dorsally flattened, tapering slightly posteriorly. Dorsum with 2 transverse ciliatedbands per segment.
Prostomium bilobed, wider than long, with distinct cephalic peaks. Two pairs of eyes, anterior pair circular, lying dorsolaterally under cephalic peaks of prostomium; posterior pair lying
dorsally, near posterior edge of prostomium. Ceratophore of median antenna large, cylindrical,
inserted in anterior notch, with cylindrical papillate style with subterminal enlargement and filamentous tip. Lateral antennae with small ceratophores, inserted ventrally on underside of prostomium; styles short, slender, about half as long as median antenna. Palps thick, gently tapering totips, papillated (Fig. 3A).
Elytra 15 pairs, on segments 2, 4, 5, 7, alternate segments to 21, 23, 26, 29 and 32. Elytra
large, covering dorsum completely overlapping medially and posteriorly; first pair oval, subsequent pairs subreniform. Elytra of first pair (Fig. 3B) with fringe of slender, long papillae along
anterior border (Fig. 3C); median part of dorsal surface of elytron with numerous microtubercles
with chitinous quadrilobate tips extremely variable in size and shape (Fig. 3D F). Macrotubercles
mainly occurring on posterior half of elytra and along anterior margin (Fig. 3G I). Following elytra
large reniform, with fringe of slender papillae along outer and posterior borders and scattered on
surface. Greater part of dorsal surface of each elytron covered with microtubercles ranging fromsmall, blunt knobs near anterior margins to larger, denticulate cones. Macrotubercles large, cylindrical with minute denticles on tips, along mainly inner and rear borders, and other macrotubercles
crowding on anterior outer areas of each elytron; each large, cylindrical with some quadrifid tips
(Fig. 3J).
Tentacular segment not visible dorsally; tentaculophores lateral to prostomium, with

52

Minoru Imajima

Fig. 3. Gattyana ciliata Moore. —A, anterior end, dorsal view, × 9; B, right 1st elytron, × 11; C, marginal papillae
from 1st elytron, × 40; D F, microtubercles on elytron, × 224; G I, macrotubercles, × 54; J, median elytron, ×
15, with detail of papillae and macrotubercles, × 72.

1 2 peristomial setae on each inner side; dorsal and ventral tentacular cirri tapering gradually tosubterminal inflation and filiform tip, covered with slender, clavate papillae. Segment 2 with firstpair of elytra, biramous parapodia, and ventral cirri much longer than following ventral cirri.
Parapodia biramous, similar along length of body. Notopodium smaller than neuropodium,
conical, with short acicular lobe. Neuropodium with longer, subtriangular presetal acicular lobe
with supra-acicular digitiform process, and shorter, rounded postsetal lobe. Dorsal cirri with large,

Deep-sea Benthic Polychaetes off Northern Honshu

53

Fig. 4. Gattyana ciliata Moore. —A, median parapodium, anterior view, × 11; B, same, posterior view, distal partsof dorsal cirrus, notosetae and neurosetae omitted, × 11; C, short, outer notoseta from median parapodium, ×120, with detail of part, × 338; D, long, inner notoseta from same, × 67, with detail of part, × 338; E, upperneuroseta from same, × 108; F, lower neuroseta from same, × 170.

cylindrical cirrophores on posterior sides of notopodia; styles long, tapering to slight subterminal
inflation and filiform tip, with slender, long papillae. Ventral cirri shorter than neuropodium, cirriform, sparsely covered with minute papillae. Nephridial papillae short, from 5th parapodium
onwards (Fig. 4A B).
Notosetae of one kind longer, fine-tipped distally flexible with many more rows of spines

54

Minoru Imajima(Fig. 4C D). Neurosetae larger than notosetae, all unidentate, slightly arched to terminal hook.
Neurosetae of supra-acicular group longer and thicker than subacicular ones, with about 10 spinousrows below long unidentate tips (Fig. 4E); neurosetae of subacicular group, with short spinous
regions of 4 to 5 transverse rows of spines (Fig. 4F). Pygidium with pair of long anal cirri on shortcirrophores.
The species is newly added to the Japanese polychaetous fauna.
Distribution. Chuckchee Sea from Bering Sea; Japan.
Genus Harmothoe Kinberg, 1856
Harmothoe aspera (Hansen, 1878)
Harmothoe aspera: Ditlevsen, 1917: 35, pl. 2, fig. 7; Annenkova, 1937: 152, pl. 1, fig. 3; Uschakov, 1955: 157, fig. 36; Imajima and Hartman, 1964: 32 33, pl. 4, figs. a h.

Material. NSMT-Pol. 110137, WA07-B510 (3).
Distribution. North Sea; Arctic Ocean; Sea of Okhotsk; Japan.
Harmothoe extenuata (Grube, 1840)
Polynoe extenuata Grube, 1840: 86.
Harmothoe extenuata : Ehlers, 1913: 446 447; Pettibone, 1953: 31; Imajima, 1961: 83; Imajima and Hartman, 1964: 33 34,
pl. 5, figs. a e; Imajima, 1997b: 20 21; Imajima, 2006: 325.

Material. NSMT-Pol. 109566, WA05-GH510D (1).
Distribution. Circumpolar. Widespread in the Arctic, Mediterranean Sea, California, Japan.
Harmothoe grandispina Annenkova, 1937
(Fig. 5A I)
Harmothoe impar grandispina Annenkova, 1937: 152, pl. 1, fig. 2; Uschakov, 1965: 140, fig. 41F.

Material. NSMT-Pol. 110345, WA05-FG280 (2), NSMT-Pol. 110346, WA05-FG380 (1).
Description. All specimens somewhat damaged. Anterior fragment consisting of 8 segments
9 mm long and 8 mm wide including parapodia.
Prostomium bilobed, wider than long, with distinct cephalic peaks. Two pairs of eyes, anterior pair circular, lying dorsolaterally at widest part of prostomium, posterior pair lying dorsallynear posterior margin of prostomium, closer to midline. Ceratophore of median antenna large,
cylindrical, inserted in anterior notch, style missing. Lateral antennae with small ceratophores,
inserted ventrally on underside of prostomium; styles short, about two-third as long as prostomium(Fig. 5A). Palps missing. Pharynx with 9 pairs of marginal soft papillae and 2 pairs of chitinous
jaws.
Only four pairs of elytra, on segments 2, 4, 5 and 7, overlapping medially and posteriorly.
Elytra of first pair oval, with fringe of short papillae along outer-lateral border, dorsal surface covered with numerous conical microtubercles. Macrotubercles large horn-like, confined to single
row near external border (Fig. 5B). Following elytra large reniform; dorsal surface of each elytron
covered with conical microtubercles, and about 10 horn-like macrotubercles arranged along rearborders; marginal fringe of slender papillae inserted between 2 macrotubercles (Fig. 5C).
Tentacular segment not visible dorsally; tentaculophores lateral to prostomium, with 1 peristomial seta on each inner side; right dorsal tentacular cirrus tapering gradually to filiform tip (Fig.

Deep-sea Benthic Polychaetes off Northern Honshu

Fig. 5. Harmothoe grandispina Annenkova. —A, anterior end, dorsal view, × 17; B, left 1st elytron, × 28; C, left
elytron from segment 5, × 23, with detail of parts, × 112; D, right parapodium, anterior view, × 19; E, outer
notoseta, × 124, with detail of part, × 250; F, inner notoseta, × 124; G, detail of part of inner notoseta, × 250;
H, upper neuroseta, × 124, with detail of tip, × 320; I, lower neuroseta, × 160.

55

56

Minoru Imajima

5A); left dorsal and ventral tentacular cirri missing. Segment 2 with first pair of elytra, biramous
parapodia, and ventral cirri much longer than following ventral cirri.
Parapodia biramous; notopodium smaller than neuropodium, conical, with short acicularlobe. Neuropodium with longer, subtriangular presetal acicular lobe with supra-acicular digitiform
process, and shorter, rounded postsetal lobe. Dorsal cirri with large, cylindrical cirrophores onposterior sides of notopodia; styles long, tapering to slight subterminal inflation and filiform tip.
Ventral cirri shorter than neuropodium, cirriform, sparsely covered with minute papillae (Fig.
5D).
Notosetae stouter than neurosetae; outer ones in palisade, short, with close-set spinous rows
along convex edge (Fig. 5E); inner ones much larger, with distinct numerous spinous rows below
bare, tapered tips (Fig. 5F G). Neurosetae long, straight; upper ones with longest spinous regionswith many alternating rows of strong serrations below fine, bidentate tips (Fig. 5H); lower ones
with few rows of fine serrations below unidentate tips (Fig. 5I).
The species is newly added to the Japanese polychaetous fauna.
Distribution. Sea of Japan; Japan.
Harmothoe imbricata (Linnaeus, 1767)
Aphrodita imbricata Linnaeus, 1767: 1084.
Harmothoe imbricata: Tebble and Chambers, 1982: 30 31, figs. 7a, 9a b; Imajima, 1997b: 29 31, fig. 14a l.

Material. NSMT-Pol. 110559, WA07-C410 (1).
Distribution. Great Britain; Arctic and north Pacific oceans; Indian Ocean; Japan.
Harmothoe impar (Johnston, 1839)
Polynoe impar Johnston, 1839: 436.
Harmothoe impar: Ditlevsen, 1917: 12, pl. 2, fig. 16, pl. 3, fig. 11; Uschakov, 1955: 157, fig. 41A E; Imajima and Hartman,
1964: 34, pl. 6, figs. a g.

Material. NSMT-Pol. 110344, WA05-FG380 (2), NSMT-Pol. 109567, WA05-FG410 (1);
NSMT-Pol. 110138, WA07-A550 (1), NSMT-Pol. 110139, WA07-B750 (2). NSMT-Pol. 110616,
KT07-29-M-2 (1), NSMT-Pol. 110617, KT07-29-M-3-2 (1).
Distribution. North Atlantic; Mediterranean Sea; Arctic Ocean; southern California; Sea of
Okhotsk; Japan.
Subfamily Lepidastheniinae Pettibone, 1989
Genus Lepidasthenia Malmgren, 1867
Lepidasthenia interrupta (Marenzeller, 1902)
Halosydna interrupta Marenzeller, 1902: 570, pl. 1, fig. 2.
Polynoe semierma Moore, 1903: 402 403, pl. 23, figs. 2 3; Izuka, 1912: 30 31.
Lepidasthenia interrupta: Seidler, 1924: 163 164; Imajima and Hartman, 1964: 22; Imajima, 1997b: 57 59, fig. 28a k.

Material. NSMT-Pol. 109734, WA05-EF450D (1), NSMT-Pol. 109766, WA05-GH510D (1),
NSMT-Pol. 109786, WA05-H380 (1).
Distribution. Japan.

Deep-sea Benthic Polychaetes off Northern Honshu

57

Lepidasthenia izukai Imajima and Hartman, 1964
Polynoe longissima Izuka, 1912: 34 36, pl. 1, fig. 1, pl. 4, figs. 1 5.
Lepidasthenia izukai Imajima and Hartman, 1964: 22 23; Imajima, 1997b: 54 55, fig. 27a n.

Material. NSMT-Pol. 109869, WA05-EF450D (1), NSMT-Pol. 109568, WA05-GH380D (2).
Distribution. Japan; Yellow Sea.
Lepidasthenia magnacornuta (Moore, 1903)
Hylosynda magnacornuta Moore, 1903: 419 420, pl. 23, fig. 18.
Polynoe magnacornuta: Izuka, 1912: 40.
Lepidasthenia magnacornuta: Seidler, 1924: 165 167; Imajima and Hartman, 1964: 23.

Material. NSMT-Pol. 109569, WA05-FG410 (1), NSMT-Pol. 109570, WA05-G425 (1).
Distribution. Japan.
Subfamily Lepidonotinae Willey, 1902
Genus Lepidonotus Leach, 1816
Lepidonotus glaber Imajima, 1997
Lepidonotus glaber Imajima, 1997b: 100 103, figs. 47a g, 48a e, 49a h; Imajima, 2006: 327.

Material. NSMT-Pol. 110500, WA07-C310 (1).
Distribution. Japan.
Genus Parahalosydna Horst, 1915
Parahalosydna krassini (Annenkova, 1952)(Fig. 6A I)
Lepidasthenia (?) krassini Annenkova, 1952: 148, fig. 1.
Parahalosydna krassini: Uschakov, 1965: 113, fig. 23A D.

Material. NSMT-Pol. 110340, WA06-EF425D (12).
Description. All specimens divided into anterior and posterior parts; largest anterior body 13
mm long, about 10 mm wide including parapodia for 17 setigers. Styles of median and lateral antennae, tentacular and dorsal cirri, with brownish pigmentation. Body dorsoventrally flattened,
without color markings.
Prostomium bilobed, wider than long, without cephalic peaks. Two pairs of black eyes inquadrate arrangement, anterior pair slightly lying dorsolaterally. Ceratophore of median antenna,large, cylindrical, inserted in anterior notch; style smooth, cylindrical basally, tapering to filiform
tip. Lateral antennae inserted terminally on distal ends of prostomium, on same level as medianantenna; styles short, about half as long as style of median antenna, with filiform tips. Palps short,stout basally, gently tapering to tips. Tentacular segment not visible dorsally. Tentaculophores lateral to prostomium, 2 pairs of dorsal and ventral tentacular cirri longer than style of median antenna (Fig. 6A). Pharynx with 9 pairs of marginal soft papillae and 2 pairs of chitinous jaws (Fig.
6B).
Elytra 15 pairs, on segments 2, 4, 5, 7, and alternate segments to 29. Elytra covering dorsum
entirely. Elytra delicate, translucent, without fringe of papillae; surface of elytra with many conicaland digitate microtubercles (Fig. 6C D).

58

Minoru Imajima

Fig. 6. Parahalosydna krassini (Annenkova). —A, anterior end, dorsal view, × 18; B, marginal papillae of pharynx,
lateral view, × 12; C, median elytron, × 11; D, microtubercles on elytron, × 170; E, left cirrigerous parapodium
from segment 12, anterior view, × 23; F, same, posterior view, distal parts of neurosetae omitted, × 23; G,
notoseta, × 266; H, upper neuroseta, × 266; I, lower neuroseta, × 266.

Second segment with first pair of elytra, biramous parapodia, and smooth ventral buccal cirri
longer than following ventral cirri. Parapodia biramous and similar along length of body. Notopodium very small, conical, on anterior face of neuropodium (Fig. 6E). Neuropodium with subtriangular presetal acicular lobe; postsetal lobe shorter than presetal one, rounded distally (Fig. 6F).
Dorsal cirrophores large and cylindrical, styles long, tapering gradually to filiform tips (Fig. 6E F).

Deep-sea Benthic Polychaetes off Northern Honshu

59

Ventral cirri short, smooth, with slender tips (Fig. 6F). Nephridial papillae from segment 8 onwards. Upper three to four notosetae short and slender, tapering to rounded tips, with many spinousrows (Fig. 6G). Neurosetae stout, upper ones slender, with long region of fine serrations below
rounded tips (Fig. 6H); lower ones with few rows of fine serrations below rounded tips (Fig. 6I).
Pygidium with pair of long anal cirri.
The species is newly added to the Japanese polychaetous fauna.
Distribution. Sea of Okhotsk; Chuckchee Sea; Japan.
Genus Weberia Horst, 1915
Weberia abyssicola sp. nov.
(Figs. 7A E, 8A-D)
Type material. Holotype—NSMT-Pol. H 480, Stn. KT07-29-M-3-2, off Miyako, 39˚20.19´N,
142˚51.40´E 39˚19.23´N, 142˚49.18´E, 1737 1709 m, Nov. 6, 2007.
Description. Body of holotype somewhat incomplete, for elytra as well as most dorsal cirrilacking, flattened, fusiform, 10 mm long, 3.2 mm wide including setae, with 25 segments. Dorsum
with large tubercles at base of dorsal cirrus and on median line of each segment (Fig. 7A B).
Prostomium oval, with prolonged stout ceratophores of lateral antennae, nearly two-third aslong as prostomium, styles short, with papilliform tips. Median prostomial antenna with massive
ceratophore from anterior half of prostomium, style broken off. Palps stumpy, conical. No eyes
visible. Tentacular segment not visible dorsally, with stout tentaculophores lateral to prostomium,with 2 pairs of dorsal and ventral tentacular cirri (Fig. 7A).
Elytra 12 pairs, on segments 2, 4, 5, 7, 9, 11, 13, 15, 17, 19, 21 and 23, but almost not preserved; anterior elytron small, elliptical, widely exposing dorsum. Median elytron larger than anterior one rounded, margin and surface covered with small conical tubercles and bearing 1 3 large
warts (Fig. 7B D).
Parapodia biramous, similar along length of body except first parapodium (Fig. 8A). Notopodium smaller than neuropodium, digitiform, on anterodorsal side of neuropodium, with aciculum,
without notosetae (Fig. 8B). Neuropodium with large, rounded presetal lobe and unequally bilobed
postsetal lobe; upper part short, rounded and lower part horn-like, with cylindrical papillae arranged in 2 rows along under side (Fig. 8B C); ventral cirri absent except setiger 1. Dorsal cirriwith prominent cirrophores connected by a ridge with dorsal tubercles; styles cylindrical bearing
micropapillae, tapering to digitate tips (Fig. 8B). Neurosetae all unidentate simple setae 3 4 perfascicle, with many rows of minute spines (Fig. 8D). Aciculum numbering 1 in a parapodium.
Pygidium with a pair of short anal cirri (Fig. 7E).
Remarks. Weberia abyssicola can be distinguished from known species, W. pustulata Horst,
1915 from Flores Sea (694 794 m), W. nodulosa (Ditlevsen, 1917) from south of Iceland (1089fms.) and W. nodulosa pacifica (Uschakov, 1950) from Sea of Okhotsk (1366 m), in the feature of
the characteristic neuropodium.
The genus is reported as a part of the Japanese polychaetous fauna for the first time.
Etymology. The species is named after the habitat of the specimen.
Distribution. Japan (1737 1709 m depth).

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Minoru Imajima

Fig. 7. Weberia abyssicola sp. nov. —A, anterior end, dorsal view, × 22; B, median part of body, dorsal view, × 22;
C, elytron on segment 13, × 45, with detail of elytral margin with conical tubercles, × 122; D, large wart on
elytron, × 122; E, posterior end, dorsal view, × 22.

Family Acoetidae Kinberg, 1858
Genus Acoetes Audouin and Milne-Edwards, 1832
Acoetes jogasimae (Izuka, 1912)
Panthalis jogasimae Izuka, 1912: 68 71, pl. 2, fig. 6, pl. 8, figs. 1 6; Imajima and Hartman, 1964: 41; Wu, Sun and Chen,
1980: 114.
Acoetes jogasimae: Pettibone, 1989: 98; Imajima, 1997b: 122 126, figs. 60a i, 61a o, 62a j.

Deep-sea Benthic Polychaetes off Northern Honshu

61

Fig. 8. Weberia abyssicola sp. nov. —A, right elytragerous parapodium from segment 2, posterior view, × 48; B,
right cirrigerous parapodium from segment 14, same view, × 43; C, neuropodium from same parapodium,same view, × 113; D, neuroseta from same parapodium, × 240.

Material. NSMT-Pol. 110347, WA06-H250D (3).
Distribution. Japan; South China Sea.
Family Pholoidae Kinberg, 1858
Genus Pholoe Johnston, 1839
Pholoe sp.
Material. NSMT-Pol. 109905, WA05-DE250D (3); NSMT-Pol. 110272, WA06-DE280D(12).
Family Sigalionidae Malmgren, 1867
Genus Labioleanira Pettibone, 1992
Labioleanira sp.
Material. NSMT-Pol. 110864, SO07-C7-B (4).

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Minoru Imajima

Genus Leanira Kinberg, 1855
Leanira sp.
Material. NSMT-Pol. 110865, SO07-O3 (15).
Genus Neoleanira Pettibone, 1970
Neoleanira areolata (McIntosh, 1885)
Leanira areolata McIntosh, 1885: 151 153, pl. 21, fig. 3, pl. 25, figs. 8 9, pl. 13A, fig. 1.
Sthenolepis areolata: Moore, 1910: 391; Izuka, 1912: 89 90, pl. 10, fig. 8; Imajima and Hartman, 1964: 42 43.
Neoleanira areolata: Pettibone, 1970: 372 376, figs. 5 6; Imajima, 2003: 56 59, figs. 32a h, 33a d, 34a i; Imajima, 2005:
62.

Material. NSMT-Pol. 109571, WA05-DE380D (8), NSMT-Pol. 109572, WA05-E1000D (1),
NSMT-Pol. 109573, WA05-EF450D (6), NSMT-Pol. 109574, WA05-F900 (1), NSMT-Pol.
109575, WA05-FG450 (1), NSMT-Pol. 109576, WA05-G210 (1), NSMT-Pol. 109577, WA05GH350 (1), NSMT-Pol. 109578, WA05-GH380D (2), NSMT-Pol. 109579, WA05-GH425 (1),
NSMT-Pol. 109580, WA05-GH480 (1), NSMT-Pol. 109581, WA05-GH510 (1), NSMT-Pol.
109582, WA05-H310 (1); NSMT-Pol. 109583, WA06-A450 (1), NSMT-Pol. 109584, WA06-A510(5), NSMT-Pol. 109585, WA06-EF425D (2), NSMT-Pol. 109586, WA06-F480 (1), NSMT-Pol.
109587, WA06-F900 (1), NSMT-Pol. 109588, WA06-G550 (1), NSMT-Pol. 109589, WA06G900D (1), NSMT-Pol. 109590, WA06-GH480D (2); NSMT-Pol. 110072, WA07-A410 (1), NSMT-Pol. 110073, WA07-A450 (7), NSMT-Pol. 110074, WA07-B410 (1), NSMT-Pol. 110075,
WA07-B410D (4), NSMT-Pol. 110076, WA07-D900 (1). NSMT-Pol. 110891, SO07-C7-B (1).
Distribution. Sea of Okhotsk; Bering Sea; off Washington to southern California; Japan.
Genus Sigalion Audouin and Milne-Edwards, 1830
Sigalion orientalis sp. nov.
(Figs. 9A D, 10A D, 11A H)
Type material. Holotype—NSMT-Pol. H 481, Stn. WA06-D210D, off Sanriku, 38˚56.42´N,
141˚59.28´E 38˚56.24´N, 141˚59.17´E, 213 214 m, Oct. 19, 2006. Paratypes—NSMT-Pol. P 482,
Stn. WA06-D210D (1), same locality as holotype; NSMT-Pol. P 483, Stn. WA06-A150D, off Sanriku, 146 147 m, Oct. 9, 2006 (1).
Description. All specimens missing posterior ends; holotype 20 mm long, 5 mm wide including parapodia for 28 setigers.
Prostomium rectangular, situated dorsally on segments 1 and 2, with 2 small digitiform lateral antennae on anterior margin, median antenna absent. Two pairs of small black subdermal eyesin rectangular arrangement present on anterior half of prostomium (Fig. 9A). Palps long, smoothand tapered, emerging anteriorly from basal regions of first parapodia. Pair of nuchal organs between prostomium and elytrophores of segment 2.
Parapodia of tentacular segment uniramous, anteriorly directed and medially fused, with pairof tentacular cirri; dorsal tentacular cirri slightly longer than ventral tentacular cirri (Fig. 9 B). Allother parapodia biramous. Segment 2 with small elytrophores, notopodia smaller than neuropodia;
neuropodia elongate, with triangular presetal lobe and long tapered ventral cirri extending beyond
tip of neuropodium (Fig. 9C). Parapodia of segment 3 fused dorsally with segment 2, with long
dorsal cirrus (Fig. 9D). Notopodia with elongate presetal lobe, neuropodia distally expanded withprojecting acicular lobes; superior neuropodial postsetal lobe rudimentary and inferior neuropodial

Deep-sea Benthic Polychaetes off Northern Honshu

63

Fig. 9. Sigalion orientalis sp. nov. —A, anterior end, dorsal view, × 16; B, right tentacular segment, outer view, ×
23; C, right parapodium of segment 2, posterior view, × 26; D, right parapodium of segment 3, same view, ×
26.

postsetal lobe conspicuous throughout. Large ciliated ctenidial pads, 3 per parapodium on segment
4 thereafter, occur between elytrophores and notopodia. Solitary ctenidia present on under faces ofparapodia of segments 3 5, and also present on posterior faces of parapodia from segment 4 throughall segments. Cirriform branchiae with ciliated inner margins present from parapodia of segment 5
and on all parapodia (Fig. 10A). Ventral cirri with distinct cirrophores. More posterior parapodiawith notopodia and neuropodia of approximately equal length, each with single aciculum. Notopodia club-shaped, with presetal distal stylode from segment 4. Neuropodia distally expanded withprojecting acicular lobes and more or less flattened anterior and posterior faces (Fig. 10B).
Elytra on setigers 2, 4, 5, 7, then alternate setigers to 27, and thereafter on all setigers, completely covering dorsum. First pair of elytra oval, remainder subrectangular (Fig. 10C). Outer lateral margin of each elytron with large 13 17 fringe papillae. Each fringe papilla pinnate, with up 7
slender pinnules either side. Fringe papillae with usually 3 shorter, single robust basally inserted
and some pinnules occurring on surface of elytra (Fig. 10D). Solitary short papilla occurring on
posterior margin of elytra (Fig. 10C).
Notopodia with simple distally bifurcate spinose capillaries (Fig. 11A). Neurosetae of segment 2 all slender bidentate compound multiarticulate falcigers with 6 15 articles. Neurosetae ofsegment 3 similar, but with few articles. Superior neurosetae in 2 oblique rows running posteriorly

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Minoru Imajima

Fig. 10. Sigalion orientalis sp. nov. —A, right parapodium of segment 5, posterior view, × 26; B, left parapodiumof segment 28, anterior view, × 26; C, left elytron from segment 28, × 30; D, elytron fringe papillae from same,× 150.

above acicular lobe. Upper group with 2 3 simple spinose setae, beginning from segment 4,
spinules surrounding shaft (Fig. 11B). Lower group with compound multiarticulate falcigers with
10 11 articles. Setal shafts of upper falcigers coarsely spinose (Fig. 11C), but those of median andposterior setae smooth. Terminal article of multiarticulate blades with small irregularly distributed
teeth on inner margins (Fig. 11D). Inferior neurosetae arranged in dense bundle below acicularlobe and include 4 or 5 robust compound falcigers with blades of 2 5 articles in upper posterior
position (Fig. 11E F). Remaining inferior neurosetae compound multiarticulate falcigers, more
slender than superior counterparts; distal article of blades with small teeth on inner margins. Shaftof all inferior neurosetae with smooth subrostral regions (Fig. 11G H).
Remarks. Sigalion orientalis resembles S. edwardsi Kinberg, 1856 of syntypes from Argentina examined by Mackie and Chambers (1990) in that, the presence of branchiae on all elytrophores and dorsal tubercles from setiger 5 and the number of elytral fringe papillae (up to 7).
However, S. orientalis differs from S. edwardsi in that: (1) the median antenna is lacking rather
than present, (2) the simple spinose neurosetae are present from setiger 4 rather than from setiger
6, (3) the robust compound multiarticulate falcigers with coarsely spinose subrostral regions

Deep-sea Benthic Polychaetes off Northern Honshu

65

Fig. 11. Sigalion orientalis sp. nov. —A, notoseta, × 375, with detail of distal and median regions, × 755; B, simple
spinose seta, × 375; C, upper compound multiarticulate neuroseta with spinosed shaft, × 190, with detail of
distal region, × 755 and shaft, × 530; D, same with smooth shaft, × 190; E, upper inferior compound neuroseta with blade of 2 articles, × 242, with detail of distal region, × 480; F, same with blade of 5 articles, × 190;
G H, lower inferior compound multiarticulate neurosetae, × 190, with detail of distal region, × 755.

absent rather than present, (4) compound falcigers with single articule blade among the superior
neurosetae absent rather than present.
Sigalion orientalis can be distinguished from S. mathildae Audouin and Milne-Edwards (inCuvier, 1830) in the form of the elytral fringe papillae and the presence of branchiae from setiger
5. S. mathildae is commonly found in intertidal shallow sublittoral (to 25 m) sands throughout
Europe from the Mediterranean to Scotland. Sigalion orientalis differs markedly from S. shimo-

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Minoru Imajima

daensis Imajima (2006) in the presence of branchiae and setal structures.
Etymology. The species name is after Asian district where the specimens were collected.
Distribution. Japan (146 214 m depth).
Family Phyllodocidae Williams, 1852
Subfamily Eteoninae Bergström, 1914
Genus Eulalia Savigny, 1818
Eulalia bilineata (Johnston, 1840)
Eulalia bilineata: Malmgren, 1865: 99, pl. 13, fig. 26; Fauvel, 1923: 162 163, fig. 58a e; Imajima and Hartman, 1964: 61 62,
pl. 13, figs. a d; Imajima, 1988: 125; Imajima, 1992: 127; Imajima, 1994: 108; Imajima, 1997a: 166; Imajima, 2001a:
55.

Material. NSMT-Pol. 110563, WA06-B750D (1).
Distribution. North Sea; Atlantic Ocean; Mediterranean Sea; Pacific Ocean of western Canada; Japan.
Eulalia sp.
Material. NSMT-Pol. 110348, WA06-GH480D (3).
Genus Mysta Malmgren, 1865
Mysta sp.
Material. NSMT-Pol. 110349, WA06-GH480D (1).
Subfamily Phyllodocinae Örsted, 1843
Genus Paranaitis Southern, 1914
Paranaitis polynoides (Moore, 1909)
Anaitis polynoides Moore, 1909: 339 342, pl. 16, figs. 19 21.
Phyllodoce (Paranaitis) polynoides: Berkeley and Berkeley, 1948: 44, figs. 62 63; Banse and Hobson, 1974: 44, fig. 10j.
Paranaitis polynoides: Hartman, 1936: 117, 119; Hartman, 1968: 291, figs. 1 3; Hartman and Reish, 1950: 12; Uschakov,
1972: 141 142, pl. 7, figs. 1 4; Blake, 1994a: 164 165, fig. 4.22; Imajima, 2003: 98 99, fig. 60a i.

Material. NSMT-Pol. 110363, WA07-C350D (1).
Distribution. Western Canada to central and southern California; Gulf of Mexico; Sea of Japan; Kurile Islands; Japan.
Genus Phyllodoce Savigny, 1818
Phyllodoce groenlandica (Oersted, 1843)
Phyllodoce groenlandica: Bergström, 1914: 141 143, fig. 42; Fauvel, 1923: 153 154, fig. 54f i.
Phyllodoce (Anaitides) groenlandica: Berkeley and Berkeley, 1948: 46, fig. 66; Uschakov, 1950: 170; Uschakov, 1972:
133 134, fig. 5, 1 4.
Anaitides groenlandica: Hartman, 1948a: 19; Imajima, 1963: 181, pl. 12, fig. 10.

Material. NSMT-Pol. 110350, WA05-E1000D (6), NSMT-Pol. 110351, WA05-FG510D (2),
NSMT-Pol. 110352, WA05-GH425 (1), NSMT-Pol. 110353, WA05-GH510D (5), NSMT-Pol.
110354, WA05-H510 (3), NSMT-Pol. 110355, WA05-H900D (2); NSMT-Pol. 110356, WA06-

Deep-sea Benthic Polychaetes off Northern Honshu

67

F1500D II (2), NSMT-Pol. 110357, WA06-G900D (3), NSMT-Pol. 110358, WA06-GH480D (4);
NSMT-Pol. 110359, WA07-A650 (1), NSMT-Pol. 110360, WA07-D210D (1), NSMT-Pol. 110361,
WA07-D900 (18), NSMT-Pol. 110362, WA07-D1500D (1).
Distribution. Greenland; Arctic Ocean; Sea of Japan; west coast of North America; Bering
Sea; Sea of Okhotsk; Japan.
Phyllodoce lineate tosaensis Imajima, 2001
Phyllodoce lineate tosaensis Imajima, 2001a: 56 57, fig. 15.

Material. NSMT-Pol. 110275, WA06-B750D (1), NSMT-Pol. 110290, WA06-D210D (7).
Distribution. Japan.
Phyllodoce madeirensis Langerhans, 1880
Phyllodoce (Anaitis) madeirensis Langerhans, 1880: 307 308, pl. 17, fig. 44a b.
Phyllodoce (Anaitides) madeirensis: Fauvel, 1923: 150 151, fig. 53d h; Rullier, 1965: 21; Day, 1967a: 145, fig. 5.2.d g;
Uschakov, 1972: 138, pl. 6, figs. 7 8.
Phyllodoce madeirensis: Fauvel, 1914: 111 113, pl. 6, figs. 5 13; Imajima, 2003: 103 107, fig. 63a h; Imajima, 2005: 78.

Material. NSMT-Pol. 110778, WA05-G210D (5); NSMT-Pol. 110779, WA06-DE280D (1),
NSMT-Pol. 110780, WA06-E1200D (13), NSMT-Pol. 110781, WA06-G1200D (16), NSMT-Pol.
110783, WA06-GH480D (1), NSMT-Pol. 110782, WA06-H1500D (1).
Distribution. North Atlanti; Gulf of Mexico; Caribbean Sea; Gulf of Guinea; Japan.
Family Glyceridae Grube, 1850
Genus Glycera Savigny, 1818
Glycera branchiopoda Moore, 1911
(Fig. 12A M)
Glycera branchiopoda Moore, 1911: 302 304, pl. 20, figs. 155 156, pl. 21, figs. 157 159; Böggemann, 2002: 41 42, figs.
25 27.

Material. NSMT-Pol. 110320, WA06-B750D (3), NSMT-Pol. 110321, WA06-D450D (1),
NSMT-Pol. 110322, WA06-E510D (5), NSMT-Pol. 110323, WA06-F650D (3), NSMT-Pol.
110324, WA06-FG350D (6); NSMT-Pol. 110325, WA07-A650 (2).
Description. Complete largest specimen 45 mm long, about 4 mm wide including parapodia
for 110 setigers. Mid-body segments triannulate; two anterior annuli of about same length, posterior annulus slightly longer. Conical prostomium consisting of 9 rings; terminal annulus bearing 2
pairs of slender antennae (Fig. 12A). Proboscis with two types of papillae; numerous digitiform
papillae with straight, longitudinal ridges (Fig. 12B C), and isolated, broader, oval to globular
papillae with ridges (Fig. 12D). Aileron with inner ramus united to outer ramus by a triangularinter-ramal plate (Fig. 12E).
First 2 parapodia uniramous, following parapodia biramous. Two slender digitiform presetal
lobes: in anterior parapodia both lobes of about same length (Fig. 12F G), in mid-body notopodial
lobe distinctly longer and wider than neuropodial lobe (Fig. 12H L), in posterior parapodia both
lobes becoming slightly slimmer and notopodial lobe shorter than neuropodial one (Fig. 12M).
One postsetal lobe short, rounded, sometimes blunt triangular (Fig. 12G, I, K, M). Dorsal cirri
distinctly visible from 3rd parapodia, short digitiform, inserted on body wall far above parapodialbase. Ventral cirri slender digitiform; in anterior parapodia about as long as presetal lobes (Fig.

68

Minoru Imajima

Fig. 12. Glycera branchiopoda Moore. —A, anterior end, dorsal view, × 32; B D, proboscidial papillae, × 300; E,
aileron, × 53; F, left parapodium from setiger 20, anterior view, × 38; G, same, posterior view, × 38; H, leftparapodium from setiger 40, anterior view, × 38; I, same, posterior view, × 38; J, left parapodium from setiger
60, anterior view, × 38; K, same, posterior view, × 38; L, left parapodium from setiger 87, anterior view, × 38;
M, left parapodium from setiger 108, posterior view, × 38.

12F); in following parapodia distinctly longer and wider than notopodial presetal lobes (Fig.
12H M); situated medio-ventrally on parapodia. Branchiae absent. Notopodia with simple capillaries, neuropodia with spinigerous compound setae.
The species is newly added to the Japanese polychaetous fauna.

Deep-sea Benthic Polychaetes off Northern Honshu

69

Distribution. Mexico; California; Puget Sound; Gulf of Alaska; Japan.
Glycera capitata Oersted, 1842
Glycera capitata: O’Connor, 1987: 183 184, fig. 13; Böggemann, 2002: 34 37, figs. 16 18; Imajima, 2007: 223, fig. 66.

Material. NSMT-Pol. 110094, WA07-C350D (1), NSMT-Pol. 110095, WA07-D210D (1).
Distribution. North Atlantic Ocean; boreal and arctic seas; Alaska to California; Japan.
Glycera neorobusta sp. nov.
(Fig. 13A N)
Type material. Holotype—NSMT-Pol. H 484, Stn. WA06-A1500D, off Sanriku, 40 52.00´N,
142 33.35´E 40 51.64´N, 142 33.78´E, 1513 1512 m, Oct. 12, 2006.
Description. Holotype missing posterior end 88 mm long, 13 mm wide including parapodia
for 133 setigers. Mid-body segments biannulate; anterior annulus about as long as posterior one.
Conical prostomium consisting of 9 rings (Fig. 13A). Proboscis with two types of papillae: numerous conical papillae with 5 7 ridges (Fig. 13B E); isolated, broader, oval to globular papillae with
6 ridges (Fig. 13F); ridges more U-shaped. Aileron with triangular base (Fig. 13G).
First two parapodia uniramous. Following anterior parapodia with 2 triangular presetal lobesof about same length, and 1 postsetal lobe large, rounded (Fig. 13H). From about setiger 10 postsetal lobe slightly incised, and appearing 2 presetal and 2 postsetal lobes. Two triangular presetal
lobes of anterior region about same length, longer than postsetal lobes with pointed end distally(Fig. 13I J); postsetal lobes low, triangular, notopodial lobe shorter than neuropodial lobe (Fig.
13I).
In median parapodia presetal lobes becoming mammiform, wider basally and distally pointed(Fig. 13K); postsetal lobes rounded to blunt triangular, slightly shorter than presetal lobe (Fig.
13L). In more posterior parapodia pre- and postsetal lobes becoming mammiform with pointed
tips (Fig. 13M N), presetal lobes slightly longer than postsetal lobes. Dorsal cirri from 3rd parapodia conical, inserted on body wall slightly above parapodial base. Ventral cirri triangular, shorter
than postsetal lobes, situated medio-ventrally on parapodia. Branchiae non-retractile, blister-like,starting from about 100th parapodia, situated dorsally on parapodial bases (Fig. 13M N). Notosetae thin capillaries with finely serrated edge; neurosetae compound spinigers with finely serrated
blades.
Remarks. Glycera neorobusta is closely allied to G. robusta Ehlers, 1868 and G. pseudorobusta Böggemann and Fiege, 2001 in having the conical proboscidial papillae and the blister-likebranchiae dorsally of parapodial bases. However, G. neorobusta is distinguished from the each ofboth species in the parapodia with the mammiform pre- and postsetal lobes and the proboscidial
papillae with U-shaped ridges.
Etymology. This species is named for its great similarity to Glycera robusta Ehlers.
Distribution. Japan (1513 1512 m depth).
Glycera okai sp. nov.
(Fig. 14A J)
Glycera sp. C, Imajima, 2007: 246, fig. 82.

Type material. Holotype—NSMT-Pol. H 485, Stn. WA06-DE280D, off Sanriku, 38˚42.91´N,

70Minoru Imajima

Fig. 13. Glycera neorobusta sp. nov. —A, anterior end, dorsal view, × 14; B F, proboscidial papillae, B D, × 400, E F, ×360; G, aileron, × 20; H, right parapodium from setiger 6, posterior view, × 19; I, right parapodium from setiger 20,
anterior view, × 16; J, same, posterior view, × 16; K, right parapodium from setiger 50, anterior view, × 16; L, same,
posterior view, × 16; M, right parapodium from setiger 106, anterior view, × 16; N, same, posterior view, × 16.

141˚58.34´E 38˚43.12´N, 141˚58.37´E, 284 285 m, Nov. 23, 2006. Paratype—NSMT-Pol. P 486,
Stn. WA07-A1500D, off Sanriku, 1402 1377 m, Oct. 11, 2007 (1).
Description. Holotype of largest, complete specimen 123 mm long, 8 mm wide including
parapodia for 214 setigers. Mid-body segments biannulate, anterior annulus slightly longer thanposterior one.
Conical prostomium consisting of 9 rings; terminal annulus bearing 2 pairs of slenderDeep-sea Benthic Polychaetes off Northern Honshu

71

Fig. 14. Glycera okai sp. nov. —A, anterior end, dorsal view, × 22; B D, proboscidial papillae, × 657; E, aileron, ×
65; F, right parapodium from setiger 10, posterior view, × 26; G, same from setiger 20, same view, × 26; H,
same from setiger 50, same view, × 26; I, same from posterior setiger, same view, × 26; J, same from nearposterior end, same view, × 26.

antennae (Fig. 14A). Proboscis with two types of papillae: numerous conical papillae with 6 7
ridges (Fig. 14B C); isolated, broader, globular papillae with 6 ridges (Fig. 14D); ridges more Ushaped. Aileron with inner ramus united to outer ramus by a triangular inter-ramal plate (Fig.
14E).
First two parapodia uniramous. Following anterior parapodia with 2 triangular presetal lobesof about same length, and notopodial postsetal lobes low, hill-like and neuropodial lobes rounded(Fig. 14F). In following parapodia neuropodial presetal lobes becoming longer than notopodial
lobes and slightly slimmer posteriorly (Fig. 14G J). Two shorter postsetal lobes distinctly triangular, always notopodial lobes shorter than neuropodial lobe (Fig. 14G J). Dorsal cirri distinctly
visible from 3rd parapodia, conical, inserted on body wall slightly above parapodial base. Ventral
cirri triangular to digitiform, shorter than postsetal lobes, but more posterior cirri slender and elongated, about as long as neuropodial presetal lobe (Fig. 14J). Branchiae absent.
Remarks. Glycera okai is closely allied to G. knoxi Kirkegaard, 1995 from New Zealand(55 2677 m), in the structures of pre- and postsetal lobes of parapodia. However, G. okai has only
conical and globular papillae with 6 7 U-shaped ridges, whereas G. knoxi has numerous conicalpapillae with 4 6 U-shaped ridges and isolated, broader, oval to globular papillae without ridges.
Etymology. The species name is after the Director Kenji Oka, of the Marine Biological Re-

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Minoru Imajima

search Institute of Japan Co. Ltd., who always support my taxonomic study on the Japanese polychaetes.
Distribution. Japan (284 1402 m depth).
Glycera semibranchiopoda sp. nov.
(Fig. 15A M)
Type material. Holotype—NSMT-Pol. H 487, Stn. WA 05-FG510D, off Sanriku, 37˚16.92´N,
141˚50.04´E, 516 515 m, Nov. 15, 2005. Paratypes—NSMT-Pol. P 488, Stn. WA05- FG510D,
same locality as holotype (2); NSMT-Pol. 489, Stn. WA05-DE380D, off Sanriku, 375 m, Nov. 19,
2005 (3); NSMT-Pol. P 490, Stn. WA05-FG250D, off Sanriku, 255 253 m, Nov. 14, 2005 (3);
NSMT-Pol. P 491, Stn. WA05-GH510D, off Kashima Sea, 512 508 m, Nov. 11, 2005 (1); NSMTPol. P 492, Stn. WA05-H900D, off Kashima Sea, 904 m, Nov. 2, 2005 (2); NSMT-Pol. P 493, Stn.
WA06-E510D, off Sanriku, 503 498 m, Nov. 3, 2006 (3); NSMT-Pol. P 494, Stn. WA06-GH480D,
off Kashima Sea, 483 478 m, Nov. 18, 2006 (3).
Description. All specimens missing posterior end, holotype of largest one 40 mm long, about
4 mm wide including parapodia for 87 setigers. Mid-body segments triannulate; all annuli of aboutsame length. Conical prostomium consisting of 9 rings; terminal annulus bearing 2 pairs of slenderantennae (Fig. 15A). Proboscis with two types of papillae; numerous digitiform or conical papillae
with straight, longitudinal ridges (Fig. 15B D), and isolated, broader, oval to globular papillae withridges (Fig. 15E). Aileron with inner ramus united to outer ramus by a triangular inter-ramal plate(Fig. 15F).
First 2 parapodia uniramous, following parapodia biramous. Two presetal lobes digitiform,
notopodial presetal lobe slightly shorter than neuropodial one throughout all parapodia (Fig.
15G L). One postsetal lobe short, rounded with a distal small notch (Fig. 15J K, M). Dorsal cirri
distinctly visible from 3rd parapodia, short digitiform, inserted on body wall far above parapodialbase. Ventral cirri digitiform; in anterior parapodia about as long as postsetal lobes (Fig. 15J); in
following parapodia distinctly longer and wider than anterior ones (Fig. 15K, M); situated medioventrally on parapodia. Branchiae absent. Notopodia with simple capillaries, neuropodia withspinigerous compound setae.
Remarks. Glycera semibranchiopoda shows some similarities to G. branchiopoda Moore,
1911 from California. Both species have the same types of proboscidial papillae, ailerons, digitiform presetal lobes and ventral cirri. However, Glycera semibranchiopoda can be distinguishedfrom G. branchiopoda in having short notopodial presetal lobes than neuropodial presetal lobes
throughout all parapodia, the notopodial presetal lobes of G. branchiopoda are elongated and longer than the neuropodial ones. Glycera semibranchiopoda and G. guatemalensis Böggemann and
Fiege, 2001 from Guatemala show almost identical proboscidial papillae and parapodial structures. However, the ailerons of G. guatemalensis have deeply incised bases rather than pointed
triangular bases.
Etymology. The species is named in having allied characteristics of G. branchiopoda.
Distribution. Japan (253 904 m depth).
Glycera tesselata Grube, 1863
Glycera tesselata Grube, 1863: 41 42, pl. 4, fig. 4; Gallardo, 1968: 70 71, pl. 21, figs. 1 6; Imajima, 2003: 117, fig. 69h l;
Imajima, 2005: 81.

Material. NSMT-Pol. 110096, WA07-B150 (1), NSMT-Pol. 110097, WA07-D210D (1), NS-Deep-sea Benthic Polychaetes off Northern Honshu

73

Fig. 15. Glycera semibranchiopoda sp. nov. —A, anterior end, dorsal view, × 32; B E, proboscidial papillae, × 300;
F, aileron, × 53; G, left parapodium from setiger 5, anterior view, × 42; H, same, posterior view, × 42; I, leftparapodium from setiger 20, anterior view, × 42; J, same, posterior view, × 42; K, left parapodium from setiger
50, same view, × 42; L, left parapodium from setiger 80, anterior view, × 42; M, same, posterior view, × 42.

MT-Pol. 110098, WA07-D900 (1).
Distribution. Mediterranean Sea; North Atlantic Ocean; western Canada to California; IndoPacific areas; Japan.

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Glycera sp.
Material. NSMT-Pol. 110866, SO07-C7-B (1).
Family Goniadidae Kinberg, 1866
Genus Glycinde Müller, 1858
Glycinde sp.
Material. NSMT-Pol. 110867, SO07-O2 (1).
Genus Goniada Audouin and Milne-Edwards, 1833
Goniada brunnea Treadwell, 1906
(Fig. 16A M)
Goniada brunnea Treadwell, 1906: 1174, figs. 67 70; Pettibone, 1963: 228, figs. 57a b; Hilbig, 1994a: 222 224, fig. 7.3;
Böggemann, 2005: 86 89, figs. 45 46.

Material. NSMT-Pol. 110029, WA05-FG510D (1), NSMT-Pol. 110030, WA05-GH510D (4);
NSMT-Pol. 110031, WA06-G900D (1), NSMT-Pol. 110032, WA06-H250D (3); NSMT-Pol.
110099, WA07-A1500D (2), NSMT-Pol. 110100, WA07-C1500D (1), NSMT-Pol. 110101, WA07D1500D (1).
Description. Largest specimen missing posterior part 73 mm long, about 3 mm wide including parapodia for 165 setigers. Segments in both anterior and posterior regions uniannulate.
Prostomium broadly triangular, depressed, consisting of 9 rings; frontal margin of distal annulus broad and truncated, with 2 pairs of small conical antennae; dorsal ones inserted slightly
more proximal than ventral ones. Basal ring fused with peristomium, much longer than preceding
annuli, without eyes (Fig. 16A). Proboscis covered with small, heart-shaped proboscideal papillae
with short stalks (Fig. 16B). Terminal jaws surrounded by 18 soft papillae (Fig. 16C). Two macrognaths 4 to 5 fangs (Fig. 16D E); dorsal arc with 4 smaller, simple inverted Y-shaped micrognaths (Fig. 16F), ventral arc with 7 H- and W-shaped compound micrognaths (Fig. 16G H). Proboscideal base armed with 15 to 17 broadly V-shaped chevrons on each side; largest ones in middleof each group (Fig. 16I).
First parapodia with only one neuropodial pre- and postsetal lobes (Fig. 16J); second, lower
presetal lobe developed from setiger 2 6; both presetal lobes digitiform, upper one slightly broader than lower one; conical to triangular postsetal lobe always distinctly shorter. Dorsal cirri foliaceous with slightly constricted base; ventral cirri long and conical (Fig. 16K). From setiger 44 45
each parapodium biramous with conical to triangular notopodial presetal lobes and shorter, rounded to slightly conical postsetal lobes; parapodia enlarged and with noto- and neuropodia clearly
separated. Neuropodial presetal lobes in median and posterior setigers each triangular, and widely
separated distally; neuropodial postsetal lobes broad and triangular, shorter than presetal lobes.
Ventral cirri becoming more conical, distinctly shorter than neuropodial postsetal lobes (Fig. 16L).
Notosetae capillary. Neurosetae compound spinigers with blades of different lengths (Fig. 16 M).
The species is newly added to the Japanese polychaetous fauna.
Distribution. North Atlantic; Indo-Pacific; southwest, central and northeast Pacific; Japan.
Goniada foliacea Moore, 1903
Goniada (Leonnatus) foliacea Moore, 1903: 457 460, pl. 26, figs. 75 76; Böggemann, 2005: 80 84, figs. 41 42.

Deep-sea Benthic Polychaetes off Northern Honshu

75

Fig. 16. Goniada brunnea Treadwell. —A, anterior end, dorsal view, × 21; B, proboscidial papilla, × 680; C, dissected proboscis, showing soft papillae, macrognaths and micrognaths, × 24; D, E, macrognaths, × 83; F, micrognath in dorsal arc, × 287; G H, micrognaths in ventral arc, × 145; I, chevrons, × 45; J, left parapodium
from setiger 1, posterior view, × 75; K, left parapodium from setiger 20, same view, × 70; L, left parapodium
from setiger 73, same view, × 57; M, part of compound spiniger, × 792.
Goniada annulata Moore, 1905: 549 553, pl. 36, figs. 45 48; Uschakov, 1955: 173, fig. 48F I; Hilbig, 1994a: 220 222, fig.
7.2; Imajima, 1997a: 169 170, fig. 8a j; Imajima, 2003: 118.

Material. NSMT-Pol. 110021, WA05-DE380D (4), NSMT-Pol. 110022, WA05-E1000D (1),
NSMT-Pol. 110023, WA05-EF450D (1), NSMT-Pol. 110024, WA05-FG450 (1), NSMT-Pol.
110025, WA05-FG510D (4), NSMT-Pol. 110026, WA05-G1500D (1), NSMT-Pol. 110027, WA05-

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GH380D (1), NSMT-Pol. 110028, WA05-GH510D (1); NSMT-Pol. 109661, WA06-D450D (1),
NSMT-Pol. 109662, WA06-E510D (15), NSMT-Pol. 109663, WA06-E1200D (2), NSMT-Pol.
109664, WA06-EF425D (3), NSMT-Pol. 109665, WA06-F1500D II (2), NSMT-Pol. 109666,
WA06-FG350D (2), NSMT-Pol. 109667, WA06-GH480D (11), NSMT-Pol. 109668, WA06H250D (1); NSMT-Pol. 110089, WA07-A450 (1), NSMT-Pol. 110090, WA07-A650 (1), NSMTPol. 110091, WA07-B410D (4), NSMT-Pol. 110092, WA07-C350D (8), NSMT-Pol. 110093,
WA07-D900 (1).
Distribution. Northwest Atlantic; east Indian Ocean; Indo Pacific; Alaska to western Mexico;
Sea of Okhotsk; Japan.
Goniada maculata Oersted, 1843
Goniada maculata: Okuda, 1939: 233 234, textfig. 8; Gardiner, 1976: 167 169, fig. 19c f; Imajima, 1997a: 169; Böggemann,
1998: 97, 99; Böggemann, 2005: 104 114, figs. 57 58.

Material. NSMT-Pol. 110033, WA05-DE250D (7), NSMT-Pol. 110034, WA05-EF480 (1),
NSMT-Pol. 110035, WA05-FG250D (40), NSMT-Pol. 110036, WA05-GH510D (1); NSMT-Pol.
110037, WA06-D210D (1), NSMT-Pol. 110039, WA06-DE280D (47), NSMT-Pol. 110040, WA06E510D (1), NSMT-Pol. 110041, WA06-GH480D (1); NSMT-Pol. 110102, WA07-B410D (7),
NSMT-Pol. 110103, WA07-D210D (8).
Distribution. Western Europe; Gulf of Iran; Atlantic of northeastern North America; Alaska;
Yellow Sea; Japan.
Goniada vorax (Kinberg, 1866)
Leonnatus vorax Kinberg, 1866: 247.
Goniada distorta Moore, 1903: 461 464, pl. 26, fig. 77.
Goniada sagamiana Imajima, 2003: 121 125, figs. 72 74.
Goniada vorax: Böggemann, 2005: 126 129, figs. 67 68.

Material. NSMT-Pol. 110042, WA05-FG510D (1).
Distribution. West Atlantic; Caribbean Sea, northeast Atlantic; Mediterranean Sea; Indian
Ocean; northwest and southwest Pacific; Japan.
Family Sphaerodoridae Malmgren, 1867
Genus Clavodorum Hartman and Fauchald, 1971
Clavodorum sp.
Material. NSMT-Pol. 110828, SO07-C7-B (62), NSMT-Pol. 110829, SO07-K1 (1), NSMTPol. 110830, SO07-K2 (2). NSMT-Pol. 110608, KT07-29-H-1 (1), NSMT-Pol. 110609, KT0729-M-1 (16).
Genus Ephesiella Chamberlin, 1919
Ephesiella brevicapitis (Moore, 1909)
Sphaerodorum brevicapitis Moore, 1909: 335 336, pl. 15, figs. 13 14; Hartman, 1968: 607 608, figs. 1 2.
Ephesiella brevicapitis: Fauchald, 1972a: 96, pl. 19, figs. a f; Kudenov, 1994: 233 234, fig. 8.1; Imajima, 2001b: 164, fig.
77.

Material. NSMT-Pol. 109651, WA06-DE280D (1). NSMT-Pol. 110826, SO07-C7-B (3),Deep-sea Benthic Polychaetes off Northern Honshu

77

NSMT-Pol. 110827, SO07-K3 (8).
Distribution. Central California; western Mexico; Japan.
Genus Sphaerodoropsis Hartman and Fauchald, 1971
Sphaerodoropsis biserialis (Berkeley and Berkeley, 1944)
Sphaerodorum biserialis Berkeley and Berkeley, 1944: 3, figs. 1 3.
Sphaerodoridium biserialis: Lützen, 1961: 415; Imajima, 1969: 154 155, fig. 3a d.
Sphaerodoropsis biserialis: Fauchald, 1974: 272, fig. 3.17; Kudenov, 1994: 234 236, fig. 8.2; Imajima, 2001b: 167, fig. 80.

Material. NSMT-Pol. 109652, WA06-DE280D (5), NSMT-Pol. 109653, WA06-GH480D (1).
NSMT-Pol. 110604, KT07-29-H-1 (1), NSMT-Pol. 110854, KT07-29-M-1 (2), NSMT-Pol.
110607, KT07-29-M-3-2 (2).
Distribution. Canada; Alaska south to California and western Mexico; Japan.
Sphaerodoropsis minuta (Webster and Benedict, 1887)
Ephesia minuta Webster and Benedict, 1887: 728 729, pl. 4, figs. 64 66.
Sphaerodorum minutum: Fauvel, 1923: 380 381, fig. 149a c; Uschakov, 1955: 222, fig. 70.
Sphaerodoridium minutum: Hartman, 1965: 94; Imajima, 1969: 153 154, fig. 2a d.
Sphaerodoropsis minuta: Fauchald, 1974: 275, fig. 3.7.

Material. NSMT-Pol. 110605, KT07-29-H-1 (9).
Distribution. Spitzbergen; Arctic and Atlantic oceans; Pacific Ocean from western Canada to
California; Alaska; Bering Sea; Japan.
Genus Sphaerodorum Oersted, 1843
Sphaerodorum gracilis (Rathke, 1843)
Ephesia gracilis: Fauvel, 1923: 377 378, fig. 148a f.
Sphaerodorum gracilis: Pettibone, 1963: 207 208, fig. 52a c; Imajima, 1969: 152 153, fig. 1a c; Fauchald, 1974: 278 279,
fig. 4.1 5.

Material. NSMT-Pol. 109654, WA05-DE250D (15), NSMT-Pol. 109655, WA05-G280 (1);
NSMT-Pol. 109656, WA06-A450 (1), NSMT-Pol. 109657, WA06-D450D (1), NSMT-Pol. 109658,
WA06-G900D (1); NSMT-Pol. 110381, WA07-A450 (1), NSMT-Pol. 110382, WA07-B410D (3),
NSMT-Pol. 110383, WA07-C350D (4), NSMT-Pol. 110384, WA07-D210D (2). NSMT-Pol.
110606, KT07-29-H-1 (19).
Distribution. Atlantic Ocean off Europe and America; Japan.
Family Hesionidae Sars, 1862
Genus Podarkeopsis Laubier, 1961
Podarkeopsis glabra (Hartman, 1961)
Oxydromus arenicolus glabrus Hartman, 1961: 68.
Podarkeopsis glabra: Hilbig, 1994b: 261 263, fig. 9.9; Imajima, 2007: 449, fig. 142.

Material. NSMT-Pol. 109593, WA05-FG250D (1).
Distribution. California to Alaska; Japan.

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Family Pilargidae St. Joseph, 1899
Genus Ancistrosyllis McIntosh, 1879
Ancistrosyllis groenlandica McIntosh, 1879
Ancistrosyllis groenlandica McIntosh, 1879: 502, pl. 65, figs. 3, 20; Imajima, 1987: 153 155, fig. 2a k; Imajima, 2001b: 180,
fig. 83.

Material. NSMT-Pol. 109594, WA05-FG510D (1), NSMT-Pol. 109595, WA05-GH510D (2);
NSMT-Pol. 109596, WA06-E1200D (1), NSMT-Pol. 110365, WA06-GH480D (1), NSMT-Pol.
109597, WA06-H1500D (1); NSMT-Pol. 110366, WA07-A1500D (1), NSMT-Pol. 110367, WA07B1500D (1), NSMT-Pol. 110368, WA07-C1500D (1).
Distribution. West Greenland; off northeastern South America; Mediterranean Sea; Japan.
Genus Cabira Webster, 1879
Cabira sp.
Material. NSMT-Pol. 110599, KT07-29-H-2 (1).
Genus Sigambra Müller, 1858
Sigambra bassi (Hartman, 1947)
Ancistrosyllis bassi Hartman, 1947a: 501 504, pl. 61, figs. 1 7.
Sigambra bassi: Pettibone, 1966: 186, fig. 16a f; Hartman, 1968: 389, figs. 1 5; Salazar- Vallejo, 1986: 200; Blake, 1994b:
287 288, fig. 10.7; Licher and Westheide, 1997: 6 10, figs. 1 2; Imajima, 2001b: 184, fig. 87.

Material. NSMT-Pol. 109598, WA05-FG250D (1).
Distribution. Gulf of Mexico; NW Atlantic Ocean; NE Pacific Ocean; Japan.
Sigambra bidentata Britaev and Saphronova, 1981
Sigambra bidentata Britaev and Saphronova, 1981: 1315 1316, textfig. 1; Salazar-Vallejo, 1990: 511; Imajima, 2001b: 185,
fig. 88.

Material. NSMT-Pol. 109599, WA05-G1500D (1); NSMT-Pol. 110364, WA06-H1500D (1)Distribution. Sea of Japan; Japan.
Sigambra spp.
Material. NSMT-Pol. 110831, SO07-C7-B (1), NSMT-Pol. 110832, SO07-K3 (1), NSMTPol. 110833, SO07-O2 (2). NSMT-Pol. 110600, KT07-29-H-2 (1).
All specimens are missing characteristic posterior ends.
Family Syllidae Williams, 1851
Subfamily Autolytinae Rioja, 1925
Genus Autolytus Grube, 1850
Autolytus sp.
Material. NSMT-Pol. 110369, WA05-DE380D (4).

Deep-sea Benthic Polychaetes off Northern Honshu

79

Subfamily Eusyllinae Rioja, 1925
Genus Eusyllis Malmgren, 1867
Eusyllis blomstrandi Malmgren, 1867
Eusyllis blomstrandi: Fauvel, 1923: 293 294, fig. 112h m; Pettibone, 1954: 260 261, fig. 28g i; Imajima, 1966a: 92 94, textfig. 29a h; Imajima, 1996: 50, fig. 36.

Material. NSMT-Pol. 110370, WA05-DE380D (4), NSMT-Pol. 110371, WA05-EF250D (1);
NSMT-Pol. 110372, WA06-A250D (1), NSMT-Pol. 110373, WA06-GH480D (2); NSMT-Pol.
110374, WA07-B410D (1), NSMT-Pol. 110375, WA07-D210D (1).
Distribution. Spitsbergen; Iceland; Mediterranean Sea; Alaska; Massachusetts; Japan.
Subfamily Exogoninae Rioja, 1925
Genus Exogone Oersted, 1845
Exogone breviantennata Hartmann-Schröder, 1959
Exogone breviantennata Hartmann-Schröder, 1959: 125 127, figs. 75 78; Imajima, 1996: 9, fig. 3; Imajima, 2001a: 59.
Exogone verugera: Imajima, 1966b: 399, textfig. 3a h.

Material. NSMT-Pol. 109600, WA05-DE380D (1); NSMT-Pol. 109601, WA06-A450 (3).
Distribution. Mediterranean Sea; Mexico; South Africa; Australia; Japan.
Exogone sp.
Material. NSMT-Pol. 110574, SO06-M1-B (30).
Genus Sphaerosyllis Claparède, 1863
Sphaerosyllis erinaceus Claparède, 1863
Sphaerosyllis erinaceus Claparède, 1863: 45 46, pl. 13, fig. 38; Pettibone, 1963: 135 136, fig. 35a; Imajima, 1966b: 402 404,
textfig. 5a g; Imajima, 2003: 150.

Material. NSMT-Pol. 109602, WA06-A450 (5). NSMT-Pol. 110855, KT07-29-H-1 (1).
Distribution. Mediterranean Sea; Atlantic Ocean; Yellow Sea; Japan.
Sphaerosyllis sp.
Material. NSMT-Pol. 110575, SO06-M1-B (10).
Subfamily Syllinae Rioja, 1925
Genus Syllis Savigny, 1818
Syllis spongiphila Verrill, 1885
Syllis spongiphila Verrill, 1885: 435; Hartman, 1944a: 339, pl. 24, fig. 10; Imajima, 1966c: 250 251, textfig. 49l s; Imajima,
2003: 157 158; Imajima, 2005: 83.

Material. NSMT-Pol. 109603, WA05-G425 (1), NSMT-Pol. 109604, WA05-GH380 (1); NSMT-Pol. 110571, WA06-E380 (1), NSMT-Pol. 109605, WA06-GH425 (4).
Distribution. Massachusettes; Falkland Islands; western Canada; Japan.

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Genus Typosyllis Langerhans, 1879
Typosyllis alternata (Moore, 1908)
Syllis alternata Moore, 1908: 323 325, figs. a f.
Typosyllis alternata: Hartman, 1948a: 21; Imajima, 1966d: 273 275, textfig. 58a l; Imajima, 2005: 84.

Material. NSMT-Pol. 109606, WA05-DE380D (1), NSMT-Pol. 109607, WA05-G310 (1),
NSMT-Pol. 109608, WA05-GH380D (1); NSMT-Pol. 110572, WA06-DE280D (1), NSMT-Pol.
109609, WA06-G900D (1), NSMT-Pol. 109610, WA06-GH480D (8).
Distribution. Alaska; California; Vancouver Island; Japan.
Typosyllis spp.
Material. NSMT-Pol. 110376, WA05-DE250D (2), NSMT-Pol. 110377, WA05-EF250D (1);
NSMT-Pol. 110378, WA06-B310D (1); NSMT-Pol. 110379, WA07-B410D (1).
Family Nereididae Johnston, 1865
Genus Ceratocephala Malmgren, 1867
Ceratocephala borealis Wesenberg-Lund, 1950
(Figs. 17A I, 18A F)
Ceratocephala borealis Wesenberg-Lund, 1950: 18 19, chart 25, pl. 5, figs. 19 21, pl. 6, figs. 23 25.

Material. NSMT-Pol. 110385, WA05-DE380D (2), NSMT-Pol. 110386, WA05-E1000D (3),
NSMT-Pol. 110387, WA05-EF250D (4), NSMT-Pol. 110388, WA05-G1500D (1); NSMT-Pol.
110389, WA06-B750D (2), NSMT-Pol. 110390, WA06-E1200D (1), NSMT-Pol. 110391, WA06H250D (1).
Description. All specimens missing posterior ends; largest one 30 mm long, 6 mm wide atanterior body including parapodia for 32 setigers. Body slender, slightly flattened, opaque white inalcohol.
Prostomium wider than long, with 2 thick palps and 2 more slender frontal antennae fused inpairs at bases. Palps with stout ceratophores and short palpostyles. Styles of antennae extending tolevel of palps. Eyes not recognized. Four pairs of tentacular cirri with short, thick cirratophores;
longest one extending to setiger 4, smooth, tapering towards tips (Fig. 17A B). Proboscis withtypical two regions in extrusion, with curved jaws, transparent yellowish-brown with 5 coarse
teeth (Fig. 17C). Proboscis with soft papillae only on oral ring as follows: area V with three large
coniform papillae (Fig. 17D), areas VII VIII with seven papillae of same size (Fig. 17E).
First 2 parapodial segments uniramous; dorsal cirri of usual cirriform type, with rather stout,cylindrical cirrophores; neuropodia with median presetal ligule, ventral ligule and postsetal acicular lobe; neurosetae surrounding ligule semilunaryly as a thin membrane; aciculum black, distally
tapering in curved tip; ventral cirri simple (Fig. 17F G). Following parapodia all biramous, notoand neuropodia deeply separated from each other. Notopodia with large, conspicuous, slender
digitate presetal ligule and short, triangular to digitate postsetal lobes. Neuropodia larger than notopodia, with digitate presetal ligules and short ventral ligule on ventral edge. Ventral cirri bifurcated, upper one longer than lower one; two cirri arissing from common base (Fig. 17H I). From
11th parapodia dorsal cirri transformed into long sausage-shaped cirrophore with slender dorsal
cirrus at tip (Fig. 18A).
Setae entirely composite, spinigerous; all notosetae in supra-acicular bundle slightly

Deep-sea Benthic Polychaetes off Northern Honshu

81

Fig. 17. Ceratocephala borealis Wesenberg-Lund. —A, anterior end, dorsal view, × 13; B, same, ventral view, ×
13; C, jaw, × 22; D, papillae on area V of proboscis, × 30; E, papillae on areas VII VIII of proboscis, × 30; F,
1st right parapodium, anterior view, × 34; G, 1st left parapodium, posterior view, dorsal and ventral cirri lacking, × 34; H, 3rd right parapodium, anterior view, × 30; I, same, posterior view, × 30.

heterogomph spinigers with long appendages with minutely serrated margin (Fig. 18B C); those in
infra-acicular bundle with short appendages. Neurosetae in supra-acicular bundle slightly heterogomph spinigers with long appendages with coarsely serrated margin (Fig. 18D); those in infraacicular bundle homogomph spinigers with short appendages with minutely serrated margin (Fig.
18E F).
The species is newly added to the Japanese polychaetous fauna.

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Fig. 18. Ceratocephala borealis Wesenberg-Lund. —A, 12th right parapodium, anterior view, × 32; B, notoseta insupra-acicular bundle, × 153; C, setal shaft of same, × 428; D, neuroseta in supra-acicular bundle, × 153, withdetail of setal shaft, × 428; E, neuroseta in infra-acicular bundle, × 153; F, setal shaft of same, × 428.

Distribution. West of Greenland (599 1096 m depths); Japan.
Genus Nereis Linnaeus, 1758
Nereis abyssa sp. nov.
(Figs. 19A F, 20A J)
Type material. Holotype—NSMT-Pol. H 495, Stn. WA05-DE510, off Sanriku, 38˚38.99´N,
142˚07.27´E 38˚37.87´N, 142˚07.20´E, 511 511 m, Nov. 21, 2005. Paratypes—NSMT-Pol. P 496,
Stn. WA05-GH380, off Kashima Sea, 376 381 m, Nov. 12, 2005 (3); NSMT-Pol. P 497, Stn.
WA06-A510, off Sanriku, 511 510 m, Oct. 10, 2006 (1); NSMT-Pol. P 498, Stn. WA06-D550, off
Sanriku, 548 551 m, Oct. 18, 2006 (5); NSMT-Pol. P 499, Stn. WA06-E410, off Sanriku, 408 408
m, Nov. 24, 2006 (2).
Other material. NSMT-Pol. 110393, WA05-E510 (1), NSMT-Pol. 110394, WA05-F510 (3),
NSMT-Pol. 110395, WA05-F900 (1), NSMT-Pol. 110396, WA05-FG425 (4), NSMT-Pol. 110397,
WA05-FG480 (3), NSMT-Pol. 110398, WA05-G550(1), NSMT-Pol. 110399, WA05-G900 (2),
NSMT-Pol. 110400, WA05-GH250 (2), NSMT-Pol. 110401, WA05-GH280 (6), NSMT-Pol.
110402, WA05-GH380 (16), NSMT-Pol. 110403, WA05-GH425 (1), NSMT-Pol. 110404, WA05GH450 (1), NSMT-Pol. 110405, WA05-GH480 (1), NSMT-Pol. 110406, WA05-H450 (2), NSMTPol. 110407, WA05-H510 (1), NSMT-Pol. 110408, WA05-H900D (1); NSMT-Pol. 110410,

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Fig. 19. Nereis abyssa sp. nov. —A, anterior end, dorsal view, × 9; B, jaw, × 18; C, paragnaths in areas I VIII, × 36;
D, right parapodium from setiger 2, posterior view, × 23; E, same from setiger 6, same view, × 23; F, same,
anterior view, × 23.

WA06-B450 (1), NSMT-Pol. 110411, WA06-C750 (2), NSMT-Pol. 110413, WA06-DE450 (7),
NSMT-Pol. 110414, WA06-E150 (1), NSMT-Pol. 110416, WA06-E550 (1), NSMT-Pol. 110417,
WA06-E750 (1), NSMT-Pol. 110418, WA06-E900 (1), NSMT-Pol. 110419, WA06-F250 (3), NSMT-Pol. 110420, WA06-F510 (3), NSMT-Pol. 110421, WA06-F550 (2), NSMT-Pol. 110422,
WA06-F900 (1), NSMT-Pol. 110423, WA06-FG480 (14), NSMT-Pol. 110415, WA06-G480 (1),
NSMT-Pol. 110424, WA06-GH425 (5), NSMT-Pol. 110425, WA06-GH510 (4); NSMT-Pol.
110392, WA07-A410 (4), NSMT-Pol. 110409, WA07-B150 (1), NSMT-Pol. 110412, WA07-B310(1), NSMT-Pol. 110426, WA07-B410 (1), NSMT-Pol. 110427, WA07-B510 (1), NSMT-Pol.
110428, WA07-C410 (2), NSMT-Pol. 110429, WA07-C510 (12), NSMT-Pol. 110430, WA07C550 (3), NSMT-Pol. 110431, WA07-C650 (4), NSMT-Pol. 110432, WA07-C1500D (3), NSMTPol. 110433, WA07-D510 (1), NSMT-Pol. 110434, WA07-D550 (3).
Description. Holotype in median size 65 mm long, 6 mm wide including parapodia for 82

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Fig. 20. Nereis abyssa sp. nov. —A, right parapodium from setiger 26, posterior view, × 23; B, same from setiger 30
in large paratype, same view, × 18; C, same from setiger 50, same view, × 23; D, same from setiger 70, same view,
× 23; E, notopodial homogomph spiniger from setiger 26, × 196, with detail of distal part of shaft, × 612; F, notopodial homogomph falciger from setiger 50, × 612; G, notopodial homogomph falciger from posterior setiger,
× 612; H, neuropodial heterogomph falciger in upper bundle from same, × 612; I, neuropodial heterogomph
spiniger in lower bundle from same, × 612; J, neuropodial heterogomph falciger in lower bundle from same, ×612.

setigerous segments. Body robust, flattened, color yellowish brown in spirit, without pigmented
pattern.
Prostomium hexagonal, with straight, narrow, entire anterior margin. Frontal antennae longer
than half length of prostomium; eyes absent, but 2 pairs of sunken depressions representing positions of missing eyes present. Palps thick, each with globular palpostyle. Peristomium about twice

Deep-sea Benthic Polychaetes off Northern Honshu

85

as long as first setiger; four pairs tentacular cirri, longest extending to setiger 5 (Fig. 19A).
Pharynx with slender, blackish brown jaws curving more towards tips, with 8 teeth reduced
to low undulations distally (Fig. 19B). Paragnaths conical, dark brown, arranged as follows: Area
I = 4; II = 18 (right) 20 (left) in 3 oblique arcs; III = 47 in 3 transverse rows; IV = 35 36 in
oblique crescent; V = 0; VI = 6 (right) 10 (left) in small oval patch; VII-VIII = 65 in 2 3 irregularrows, some of the larger cones forming single row anteriorly (Fig. 19C).
First 2 pairs of parapodia uniramous, with slender dorsal cirrus and acutely conical notopodial ligule; neuropodium similar to those of following setigers (Fig. 19D). Parapodia biramousfrom setiger 3 posteriorly. Dorsal cirrus about 1.5 times length of upper notopodial ligule in anterior setigers, elongating to 2 2.5 times length of ligule posteriorly. Upper and lower notopodial
ligules acutely conical, upper ligules slightly longer than lower ligules in anterior region (Figs.
19E F; 20A B), and upper ligules with thickened glandular border (Fig. 20A B). Neuropodial acicular lobes subtriangular; neuropodial ligules subequal in size and shape to notopodial lower
ligules. Ventral cirri slender, about as long as neuropodial ligules. Parapodia of middle and posterior regions grandually modified; notopodium becoming slightly elevated posteriorly (Fig. 20C D);
upper and lower notopodial ligules similar in length.
Notopodia with homogomph spinigers with minute serrations along cutting margin (Fig. 20E)
in anterior parapodia, thereafter a single homogomph falciger appearing from setigers 30 38, short
blade with slender teeth along cutting margin (Fig. 20F). More posterior notopodia with only 1 2
homogomph falcigers with short rod-like blade with few serrations along cutting margin (Fig.
20G). Neuropodia with homogomph spinigers and heterogomph falcigers (Fig. 20H) in upper
bundle, and heterogomph spinigers (Fig. 20I) and heterogomph falcigers (Fig. 20J) in lower bundle. Acicula black, occurring singly in each noto- and neuropodium. Pygidium with a pair of anal
cirri.
Remarks. Nereis abyssa resembles N. anoculis Hartman, 1960 from Tanner Basin (1414 1486m) and San Nicolas Basin (1670 m) off southern California, in the features of the parapodia prolonged in their basal parts. However, N. abyssa have many paragnaths on areas VII-VIII instead ofa single row of 4 cones.
Etymology. The species is named after the habitat in deep seas.
Distribution. Japan (151 1499 m).
Genus Rullierinereis Pettibone, 1971
Rullierinereis profunda sp. nov.
(Figs. 21A E, 22A M)
Type material. Holotype—NSMT-Pol. H 500, Stn. SO06-M1-B, off Miyako, 39˚40.4´N,
143˚11.3´E, 2018 m, Jul. 16, 2006. Paratypes—NSMT-Pol. P 501, Stn. SO06-M1-B, same locality
as holotype (13).
Description. Holotype of largest, complete specimen, 16 mm long, 1 mm wide including
parapodia for 76 setigerous segments. Color in spirit uniformly tan.
Prostomium pentagonal and slightly longer than wide, with short tapered frontal antennae
inserted close together. Eyes not visible. Palps thick and elbowed, palpostyles directed medially(Fig. 21A B). Achaetous tentacular segment about twice length of following segment. Four pairsof slender tentacular cirri, longest one about as long as length of tentacular segment, and shortest
one about half as long as longest one. Pharyngeal jaws translucent yellow, distally curved, eachone bearing 7 cutting teeth (Fig. 21C); paragnaths or papillae not observed.
First 2 parapodial segments uniramous (Fig. 21D). Dorsal cirri subulate, subequal in length

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Fig. 21. Rullierinereis profunda sp. nov.—A, anterior end, dorsal view, × 45; B, same, ventral view, × 48; C, pharyngeal jaw, × 107; D, left parapodium from setiger 1, anterior view, × 130; E, same from setiger 5, same view, ×
130.

of notopodial ligules; neuropodial acicular lobes subtriangular; neuropodial ligules subequal insize and shape to notopodial ligules. Ventral cirri short, subulate. Upper bundle of neurosetae homogomph spinigers and heterogomph falcigers; lower bundle of neurosetae heterogomph spinigers and heterogomph falcigers. Following parapodia of anterior region (Fig. 21E) with upper andlower notopodial ligules large, subulate and 3 homogomph spinigers (Fig. 22A). Upper bundles ofneurosetae with homogomph spinigers and heterogomph falcigers with moderately long appendages (Fig. 22B), lower bundles with heterogomph spinigers and heterogomph falcigers with somewhat shorter appendages (Fig. 22C). In parapodia of middle regions (Fig. 22D), upper notopodial
ligules gradually reduced to small ligules than lower ligules posteriorly, with notopodial homogomph spinigers (Fig. 22E) and stout homogomph falcigers with short, rod-like appendage withminute serrations along cutting edge (Fig. 22F). Upper bundle of neurosetae homogomph spinigers and heterogomph falcigers; low bundle heterogomph spnigers and heterogomph falcigers(Fig. 22G). Posterior parapodia slightly modified (Fig. 22H), notopodia with single stout homogomph falciger (Fig. 22I) with short, rod-like appendage with 5 coarse teeth along cutting edge.
Neuropodia with homogomph spinigers with long appendages (Fig. 22J) and heterogomph falcigers in upper acicular positions, and heterogomph spinigers with rather short appendages (Fig.
22K) and heterogomph falcigers (Fig. 22L) in lower acicular positions.

Deep-sea Benthic Polychaetes off Northern Honshu

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Fig. 22. Rullierinereis profunda sp. nov.—A, notopodial homogomph spiniger from setiger 5, × 846; B, neuropodial
heterogomph falciger in upper bundle from same, × 846; C, same in lower bundle from same, × 846; D, leftparapodium from median setiger, anterior view, × 130; E, notopodial homogomph spiniger from same, × 846; F,
notopodial homogomph falciger from same, × 846; G, neuropodial heterogomph falciger in lower bundle fromsame, × 846; H, left parapodium from posterior setiger, anterior view, × 130; I, notopodial homogomph falciger
from same, × 864; J, neuropodial homogomph spiniger in upper bundle from same, × 580; K, heterogomph spiniger in lower bundle from same, × 580; L, heterogomph falciger in same, × 864; M, posterior end, dorsal view, ×48.

Pygidium with ventral slender anal cirri and dorsal anus (Fig. 22M).
Remarks. This species is referred to Rullierinereis Pettibone (1971) because the pharynx
lacks paragnaths and papillae, and the notopodia have homogomph spinigers and homogomph
falcigers in posterior setigers. Rullierinereis profunda can be distinguished from known eight

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species, Rullierinereis mexicana (Treadwell, 1942), R. bahamensis (Hartmann-Schröder, 1958), R.
zebra (Rullier, 1963), R. uncinata (Hartman, 1965), R. misakiensis (Imajima and Hayashi, 1969),
R. gallardoi (Pettibone, 1971), R. hancocki (Fauchald, 1977) and R. anoculata (Cantone, 1982), in
the structures of the characteristic prostomium and palps, and of the appendages of notopodial
homogomph falcigers.
Etymology. The species is named after the habitat of the specimens.
Distribution. Japan (2018 m depth).
Family Nephtyidae Grube, 1850
Genus Aglaophamus Kinberg, 1866
Aglaophamus japonicus Imajima and Takeda, 1985
Aglaophamus japonicus Imajima and Takeda, 1985: 73 75, fig. 8a l; Imajima, 2006: 357.

Material. NSMT-Pol. 109611, WA05-FG250D (2).
Distribution. Japan.
Aglaophamus malmgreni (Théel, 1879)
Nephthys malmgreni Théel, 1879: 26, pl. 1, fig. 17, pl. 2, fig. 17.
Aglaophamus malmgreni: Pettibone, 1956: 557; Imajima and Takeda, 1985: 68 70, fig. 6a n.

Material. NSMT-Pol. 109612, WA05-DE380D (1), NSMT-Pol. 109613, WA05-FG250D (5),NSMT-Pol. 109614, WA05-GH510D (2); NSMT-Pol. 109784, WA06-DE280D (4), NSMT-Pol.
109615, WA06-E510D (1), NSMT-Pol. 109785, WA06-GH480D (4), NSMT-Pol. 109616, WA06H250D (1). NSMT-Pol. 110812, SO07-C7-B (2), NSMT-Pol. 110813, SO07-K1 (10), NSMT-Pol.
110814, SO07-K2 (1), NSMT-Pol. 110815, SO07-O1 (3). NSMT-Pol. 110618, KT07-29-H-2 (6),
NSMT-Pol. 110619, KT07-29-M-1 (1), NSMT-Pol. 110620, KT07-29-M-2 (3), NSMT-Pol.
110621, KT07-29-M-3-2 (28), NSMT-Pol. 110622, KT07-29-M-3-3 (37).
Distribution. Arctic Ocean; Bering Sea; north Japan Sea; Japan.
Aglaophamus sinensis (Fauvel, 1932)
Nephthys sinensis Fauvel, 1932a: 536 537, fig. 1a c.
Aglaophamus sinensis: Hartman, 1950: 117; Fauchald, 1968: 12 13, figs. 16 18; Imajima and Takeda, 1985: 65 68, figs. 4a i,
5a d.

Material. NSMT-Pol. 109617, WA06-A150D (5).
Distribution. Yellow Sea; Viet Nam; Japan.
Aglaophamus spp.
Material. NSMT-Pol. 110451, WA05-EF450D (1), NSMT-Pol. 110452, WA05-G280 (4);
NSMT-Pol. 110459, WA06-D210D (1), NSMT-Pol. 110453, WA06-F650D (1); NSMT-Pol.
110454, WA07-B410D (3), NSMT-Pol. 110455, WA07-B1200 (6), NSMT-Pol. 110456, WA07C900 (4), NSMT-Pol. 110457, WA07-C1500D (1), NSMT-Pol. 110458, WA07-D210D (3). NSMT-Pol. 110623, KT07-29-M-3-3 (3).

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Genus Nephtys Cuvier, 1817
Nephtys caeca (Fabricius, 1780)
Nephtys caeca: Verrill, 1881: 294 296, 307, 314; Berkeley and Berkeley, 1948: 54, figs. 80 81; Imajima, 1961: 88 89, fig. 4;
Imajima and Takeda, 1985: 63 67, figs. 12a m, 14.

Material. NSMT-Pol. 110071, WA07-D210D (1).
Distribution. North Atlantic, Pacific and Arctic oceans; California; Yellow Sea; Japan.
Nephtys longosetosa Oersted, 1843
Nephthys longisetosa: Verrill, 1881: 295, 319; Okuda, 1939: 231 232.
Nephtys longosetosa: Hartman, 1944a: 339, pl. 15, fig. 7; Pettibone, 1963: 204 205, fig. 47a; Imajima and Takeda, 1987:
60 61, figs. 10a i, 14.

Material. NSMT-Pol. 109618, WA05-DE250D (1), NSMT-Pol. 109619, WA05-EF250D (1),
NSMT-Pol. 109620, WA05-FG250D (15); NSMT-Pol. 109621, WA06-C900D (1), NSMT-Pol.
109622, WA06-D210D (2), NSMT-Pol. 109623, WA06-E510D (2), NSMT-Pol. 109624, WA06EF425D (3).
Distribution. Northern Atlantic Ocean; Alaska to California; Bering Sea; Sea of Okhotsk;
Yellow Sea; Japan.
Nephtys oligobranchia Southern, 1921
Nephthys oligobranchia Southern, 1921: 610 611, pl. 24, fig. 12A C; Fauvel, 1932b: 119; Okuda, 1943: 100 102, figs. 1 3;
Uschakov and Wu, 1962b: 25 26, pl. 3, I; Uschakov and Wu, 1979: 55 57, fig. 17, I; Lee and Jae, 1983: 23 24, fig. 2,
pl. II, E H.
Nephtys oligobranchia: Imajima and Takeda, 1987: 47 50, figs. 4a l, 6.

Material. NSMT-Pol. 110624, KT07-29-H-1 (18), NSMT-Pol. 110625, KT07-29-M-1 (22),
NSMT-Pol. 110626, KT07-29-M-2 (1).
Distribution. Indian Ocean; Yellow Sea; Japan.
Nephtys paradoxa Malm, 1874
Nephthys paradoxa: Fauvel, 1914: 199.
Nephtys paradoxa: Hartman, 1944a: 335, 339, pl. 15, fig. 6; Fauchald, 1968: 13 15, figs. 1A, 2B, 3C; Imajima and Takeda,
1987: 50 52, figs. 5a i, 6.

Material. NSMT-Pol. 109787, WA05-DE380D (1), NSMT-Pol. 109788, WA05-GH380D(2); NSMT-Pol. 109789, WA06-A1500D (1), NSMT-Pol. 109790, WA06-D1500D (2), NSMTPol. 109791, WA06-GH480D (8), NSMT-Pol. 109792, WA06-H250D (5), NSMT-Pol. 109793,
WA06-H1500D (1); NSMT-Pol. 110077, WA07-A450 (4), NSMT-Pol. 110078, WA07-C1500D(1), NSMT-Pol. 110079, WA07-D1500 (1). NSMT-Pol. 110629, KT07-29- M-3-1 (1)Distribution. North Atlantic Ocean; Bering Strait; Greenland; Japan.
Nephtys punctata Hartman, 1938
Nephtys punctata Hartman, 1938: 155 156, fig. 67; Berkeley and Berkeley, 1942: 193; Imajima and Takeda, 1987: 61 63,
figs. 11a m, 14.

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Material. NSMT-Pol. 109625, WA05-DE250D (1), NSMT-Pol. 109626, WA05-DE380D (1),
NSMT-Pol. 109627, WA05-E1000D (1), NSMT-Pol. 109628, WA05-EF250D (8), NSMT-Pol.
109629, WA05-EF450D (5), NSMT-Pol. 109630, WA05-FG250D (3), NSMT-Pol. 109631, WA05FG510D (1), NSMT-Pol. 109632, WA05-G1500D (1), NSMT-Pol. 109633, WA05-H900D (2);
NSMT-Pol. 109634, WA06-A1200D (5), NSMT-Pol. 109794, WA06-D210D (1), NSMT-Pol.
109635, WA06-DE280D (4), NSMT-Pol. 109636, WA06-EF425 (5), NSMT-Pol. 109637, WA06F650D (1), NSMT-Pol. 109638, WA06-F750 (4), NSMT-Pol. 109639, WA06-F1500D II (3), NSMT-Pol. 109795, WA06-FG350D (1), NSMT-Pol. 109640, WA06-G900D (6), NSMT-Pol. 109641,
WA06-G1200D (4), NSMT-Pol. 109642, WA06- H1500D (3); NSMT-Pol. 110080, WA07-A250D(3), NSMT-Pol. 110081, WA07-A650 (2), NSMT-Pol. 110082, WA07-A1500D (1), NSMT-Pol.
110083, WA07-B410D (15), NSMT-Pol. 110084, WA07-B1500D (3), NSMT-Pol. 110085, WA07C350D (2), NSMT-Pol. 110086, WA07-C1500D (4), NSMT-Pol. 110087, WA07-D210D (6), NSMT-Pol. 110088, WA07-D900 (3). NSMT-Pol. 110816, SO07-C7-B (2). NSMT-Pol. 110627,
KT07-29-H-2 (1), NSMT-Pol. 110628, KT07-29-M-1 (1), NSMT-Pol. 110629, KT07-29-M-3-1(4), NSMT-Pol. 110630, KT07-29-M-3-2 (4).
Distribution. Alaska to California; Kamchatka; Japan.
Family Paralacydoniidae Pettibone, 1963
Genus Paralacydonia Fauvel, 1913
Paralacydonia paradoxa Fauvel, 1913
Paralacydonia paradoxa Fauvel, 1913: 54, fig. 55; Blake, 1994c: 364 365, fig. 14.1; Imajima, 2003: 180 182, fig. 90a j.

Material. NSMT-Pol. 109643, WA05-EF250D (1), NSMT-Pol. 109644, WA05-G280 (2),
NSMT-Pol. 109645, WA05-G1500D (3), NSMT-Pol. 109646, WA05-GH510D (3); NSMT-Pol.
109647, WA06-E510D (1), NSMT-Pol. 109648, WA06-E1200D (1), NSMT-Pol. 109649, WA06G1200D (1), NSMT-Pol. 109650, WA06-GH480D (9); NSMT-Pol. 110380 WA07-C1500D (3).
NSMT-Pol. 110601, KT07-29-H-2 (7), NSMT-Pol. 110602, KT07-29-M-3-1 (1), NSMT-Pol.
110611, KT07-29-M-3-2 (2), NSMT-Pol. 110603, KT07-29-M-3-3 (1).
Distribution. Mediterranean Sea; South Africa; off Massachusetts; Yellow Sea; off southern
California; Japan.
Order Amphinomida
Family Euphrosinidae Williams, 1851
Genus Euphrosine Lamarck, 1818
Euphrosine digitalis sp. nov.
(Fig. 23A G)
Euphrosine borealis: Imajima, 2007: 472, fig. 144 (not Oersted, 1843).

Type material. Holotype—NSMT-Pol. H 502, Stn. WA06-G425, off Kashima Sea, 36˚53.19´N,
141˚29.24´E 36˚52.05´N, 141˚27.59´E, 428 420 m, Oct. 27, 2006. Paratypes—NSMT-Pol. P 503,
Stn. WA06-G425, same locality as holotype (2); NSMT-Pol. P 504, Stn. WA05- GH450, off Kashima Sea, 454 452 m, Nov. 13, 2005 (2).
Description. Holotype 18 mm long, about 7 mm wide excluding setae at median region for 28
setigers. Body whitish yellow in spirit. Median dorsal field narrow, about one sixth body width.
Prostomium well developed, with pair of large dorsal eyes lateral to median antenna (Fig.
23A); ventral eyes inserted just anterior to palpal antennae (Fig. 23B). Median antenna short, aboutDeep-sea Benthic Polychaetes off Northern Honshu

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Fig. 23. Euphrosine digitalis sp. nov. —A, prostomium and caruncle, dorsal view, × 20; B, prostomium and setiger
1, anteroventral view, × 23; C, right setiger 14, posterior view, × 23; D, left setiger 9 omitted setae, same view,
× 23; E, slender notoseta, × 300; F, ringent notoseta, × 300; G, neuroseta, × 300.

one sixth length of caruncle, tapering distal style. Palpal antennae short, digitiform. Caruncle appearing trilobed dorsoventrally, all lobes connected by thin ventral membrane; dorsal lobe extending to setiger 4. Palps large, elliptical, paired, separated by mid-ventral groove (Fig. 23B).
Parapodia biramous, with parapodial cirri from setiger 1, with spreading notosetal and whorllike neurosetal fascicles. Notopodia not raised above dorsum. Neuropodia resembling flat circularpad (Fig. 23C D). Notosetae arrayed in 3 tiers: simple, slender notosetae (Fig. 23E) in anterior andposterior tiers; ringent notosetae (Fig. 23F) present in middle tier with simple notosetae. Neurosetae all bifurcate, each with long prong distally recurved, falcate, having a short prong (Fig. 23G).
Branchiae present from setiger 1, simple, cirriform, numbering 7 pairs per segment wherebest developed in midbody segments (Fig. 23C D), slightly longer than dorsal, lateral and ventral
cirri, but gill 1 of 2 or 3 parapodia in middle body having bifurcate filaments (Fig. 23C). Lateral
cirrus arising between gill 3 and gill 4, digitiform, shorter than branchiae (Fig. 23C D).
Remarks. Euphrosine digitalis allied to E. longesetosa Horst, 1903, E. pilosa Horst, 1903 andE. armata Kudenov, 1993 in having entire cirriform branchiae. E. digitalis differs from E. longesetosa and E. armata in having lateral cirri inserted between gill 3 and gill 4. E. digitalis resembles
E. pilosa in having lateral cirri inserted between gill 3 and gill 4 and smooth notosetae. However,E. digitalis can be distinguished from E. pilosa in that, (1) the lateral lobes of the caruncle fused to
one another rather than the lateral lobes separated from the median lobe, and free from the body

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wall (Horst, 1912, pl. 1, fig. 1), (2) the notosetae in the anterior and posterior tiers are simple,
slender, without short prong rather than having a developed, large short prong and (3) the specimens are benthic in depths of 420 m rather than pelagic worms.
Etymology. The species is named in having slender digitate branchiae.
Distribution. Japan (420 454 m depth).
Euphrosine pseudonotialis sp. nov.
(Fig. 24A G)
Type material. Holotype—NSMT-Pol. H 505, Stn. WA06-G425, off Kashima Sea, 36˚53.19´N,
141˚29.24´E 36˚52.05´N, 141˚27.59´E, 428 420 m, Oct. 27, 2006.
Description. Holotype 15 mm long, about 8 mm wide at median region for 27 setigers. Bodyin spirit whitish yellow, lacking pigmentation. Dorsomedian field narrow, about one fifth bodyFig. 24. Euphrosine pseudonotialis sp. nov. —A, prostomium and caruncle, dorsal view, × 16; B, prostomium andsetigers 1 2, anteroventral view, × 29; C, right setiger 13, posterior view, × 24; D, notoseta of anterior tier, ×306; E, ringent notoseta, × 270; F, simple notoseta, × 306; G, superior neuroseta, × 270.

Deep-sea Benthic Polychaetes off Northern Honshu

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width.
Prostomium well developed, with pair of small dosal eyes located upper-lateral to medianantenna (Fig. 24A); ventral eyes inserted just anterior to palpal antennae. Median antenna short,about one seventh length of caruncle, distal style cirriform.fl Palpal antennae short, digitiform(Fig. 24B). Caruncle appearing bilobed dorsoventrally, dorsal lobe extending to setiger 4, ventral
lobe to setiger 3. Palps large, elliptical, paired, separated by mid-ventral groove leading directly tomouth (Fig. 24B).
Parapodia biramous, with parapodial cirri from setiger 1, with spreading notosetal and whorllike neurosetal fascicles separated by narrow interramal gap occupied by branchiae 8; all withdorsal, lateral and ventral cirri from setiger 1 to end of body. Notopodia not raised above dorsum.
Neuropodia resembling flat circular pad (Fig. 24C). Notosetae shorter than neurosetae, of two basic kinds, arrayed in 3 tiers: bifurcate notosetae in anterior and posterior tiers distally calcified,inner surface of long prongs serrated with characteristic scalelike denticles and short prong extremely minute (Fig. 24D); ringent notosetae (Fig. 24E) present in middle tier associated with longsimple notosetae (Fig. 24F). Neurosetae all bifurcate, slenderer than notosetae, generally smooth,long prong of superior bristles sometimes serrated inconspicuously on inner surface (Fig. 24G).
Branchiae present from setiger 1, numbering 9 pairs per segment where best developed in
midbody segments, having 6 bifurcate and 3 simple nonbranching cirriform (Fig. 24C). Branchiae
about one half the length of longest notosetae. Dorsal and lateral cirri similar in size and shape;
ventral cirri slightly longer. Lateral cirrus arising between gill 3 and gill 4, digitiform, shorter thanbranchiae.
Remarks. Euphrosine pseudonotialis is most closely allied to E. notialis Ehlers, 1900 from
Strait of Magellan. Both species have same types of the body and branchiae. However, E. pseudonotialis can be distinguished from E. notialis in that, (1) the lateral cirri are inserted between thirdand fourth branchial pairs rather than second and third branchial pairs, (2) the bifurcate notosetae
have extremely minute short prong rather than considerable size. E. pseudonotialis resembles E.
bicirrata Moore, 1905 from northeast Pacific in having the branchiae as 2 simple cirriform filaments per gill and occasionally one. However, E. pseudonotialis can be distinguished in the structures of the median antenna, notosetae and neurosetae.
Etymology. The species is named in having allied characteristics of E. notialis.
Distribution. Japan (420 428 m depth).
Order Eunicida
Family Onuphidae Kinberg, 1865
Subfamily Hyalinoeciinae Paxton, 1986
Genus Anchinothria Paxton, 1986
Anchinothria macrobranchiata (McIntosh, 1885)
Nothria macrobranchiata McIntosh, 1885: 320 322, pl. 41, figs. 6 7, textfigs.77 78; Moore, 1903: 445.
Onuphis macrobranchiata: Izuka, 1912: 100 101.
Paranorthia macrobranchiata: Imajima and Hartman, 1964: 248.
Anchinothria macrobranchiata: Paxton, 1986: 28; Imajima, 1999: 13 16, figs. 7a g, 8a l.

Material. NSMT-Pol. 110801, SO07-O2 (33).
Distribution. Japan.

94

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Genus Nothria Malmgren, 1867
Nothria grossa Imajima, 1989
Nothria grossa Imajima, 1989a: 1 7, figs. 2a y, 3a e; Imajima, 1999: 36 40, figs. 20a i, 21a o.
Nothria abyssala Imajima, 1989a: 6 10, fig. 4a z.

Material. NSMT-Pol. 109684, WA05-DE380D (11), NSMT-Pol. 109685, WA05-E550 (19),
NSMT-Pol. 109686, WA05-F350 (3), NSMT-Pol. 109687, WA05-F450 (1), NSMT-Pol. 109688,
WA05-FG380 (3), NSMT-Pol. 109689, WA05-FG410 (1), NSMT-Pol. 109690, WA05-G410 (1),
NSMT-Pol. 109691, WA05-G425 (4), NSMT-Pol. 109692, WA05-G450 (4), NSMT-Pol. 109693,
WA05-G1500D (5), NSMT-Pol. 109694, WA05-GH380D (4), NSMT-Pol. 109695, WA05-GH450(1), NSMT-Pol. 109696, WA05-H150 (1), NSMT-Pol. 109697, WA05-H380 (6); NSMT-Pol.
109698, WA06-C350D (2), NSMT-Pol. 109699, WA06-E480 (2), NSMT-Pol. 109700, WA06E510 (2), NSMT-Pol. 109701, WA06-E550 (2), NSMT-Pol. 109702, WA06-EF425D (11), NSMTPol. 109703, WA06-EF510 (2), NSMT-Pol. 109704, WA06-F310 (3), NSMT-Pol. 109705, WA06F350 (3), NSMT-Pol. 109706, WA06-F380 (3), NSMT-Pol. 109707, WA06-F410 (3), NSMT-Pol.
109708, WA06-F425 (4), NSMT-Pol. 109709, WA06-F450 (3), NSMT-Pol. 109710, WA06-FG310(1), NSMT-Pol. 109711, WA06-FG350 (7), NSMT-Pol. 109712, WA06-G350 (4), NSMT-Pol.
109713, WA06-G380 (5), NSMT-Pol. 109714, WA06-G425 (2), NSMT-Pol. 109715, WA06-G450(4), NSMT-Pol. 109716, WA06-G480 (1), NSMT-Pol. 109717, WA06-GH380 (48), NSMT-Pol.
109718, WA06-GH425 (60), NSMT-Pol. 109719, WA06-GH450 (41), NSMT-Pol. 109720, WA06GH480D (18), NSMT-Pol. 109721, WA06-H1500D (1); NSMT-Pol. 110106, WA07-A250D (2),
NSMT-Pol. 110107, WA07-C350D (5), NSMT-Pol. 110108, WA07-C410 (11), NSMT-Pol. 110109,
WA07-D310 (1), NSMT-Pol. 110110, WA07-D350 (6), NSMT-Pol. 110111, WA07-D450 (1).
Distribution. Japan.
Subfamily Onuphinae Audouin and Milne-Edwards, 1833
Genus Epidiopatra Augener, 1918
Epidiopatra rugosa Kucheruk, 1979
Epidiopatra rugosa Kucheruk, 1979: 1228 1229, fig. 2; Imajima, 1999: 57 61, figs. 30a g, 31a l, 32a j; Imajima, 2005: 88.

Material. NSMT-Pol. 109670, WA05-DE380D (1), NSMT-Pol. 109671, WA05-E1000D (1),
NSMT-Pol. 109672, WA05-FG510D (2), NSMT-Pol. 109673, WA05-G1500D (3), NSMT-Pol.
109674, WA05-GH510D (1), NSMT-Pol. 109675, WA05-H900D (15); NSMT-Pol. 109676,
WA06-E510D (2), NSMT-Pol. 109677, WA06-F750 (1), NSMT-Pol. 109678, WA06-F1500D (6),
NSMT-Pol. 109679, WA06-F1500D II (17), NSMT-Pol. 109680, WA06-G900D (5), NSMT-Pol.
109681, WA06-G1200D (7), NSMT-Pol. 109682, WA06-GH480D (4), NSMT-Pol. 109683,
WA06-H1500D (4); NSMT-Pol. 110104, WA07-B1500D (5), NSMT-Pol. 110105, WA07-D900(3).
Distribution. Japan.
Genus Onuphis Audouin and Milne-Edwards, 1833
Onuphis geophiliformis (Moore, 1903)
Nothria geophiliformis Moore, 1903: 445 448, pl. 25, figs. 57 59; Imajima and Hartman, 1964: 244 245.
Onuphis geophiliformis: Maekawa and Hayashi, 1989: 72 74, fig. 7a j; Imajima, 1997a: 182 183; Maekawa and Hayashi,
1999: 169 173, fig. 3a j.

Deep-sea Benthic Polychaetes off Northern Honshu

95

Material. NSMT-Pol. 109723, WA06-GH480D (8), NSMT-Pol. 109724, WA06-H250D (2),
NSMT-Pol. 109725, WA06-H1500D (3).
Distribution. Eastern Pacific Ocean; Alaska; southern California; Japan.
Onuphis holobranchiata Marenzeller, 1879
Onuphis holobranchiata Marenzeller, 1879: 132 134, pl. 4, fig. 1; Izuka, 1912: 106 108, pl. 11, figs. 10 12; Imajima, 1986:
94 95, fig. 2a q.

Material. NSMT-Pol. 109726, WA06-GH480D (30), NSMT-Pol. 109727, WA06-H250D(22), NSMT-Pol. 110565, WA06-H1500D (2); NSMT-Pol. 110112, WA07-D210D (2).
Distribution. Japan.
Onuphis imajimai Maekawa and Hayashi, 1989
Onuphis imajimai Maekawa and Hayashi, 1989: 77 79, fig. 10a k; Maekawa and Hayashi, 1999: 178 182, fig. 6a k; Imajima,
2001a: 73.

Material. NSMT-Pol. 109728, WA06-E510D (43), NSMT-Pol. 109768, WA06-E1200D (1),
NSMT-Pol. 109729, WA06-F1500D II (21), NSMT-Pol. 109769, WA06-G900D (1), NSMT-Pol.
109730, WA06-G1200D (4), NSMT-Pol. 109731, WA06-GH480D (346), NSMT-Pol. 109732,
WA06-H480 (36), NSMT-Pol. 109733, WA06-H1500D (12); NSMT-Pol. 110113, WA07-A650(7), NSMT-Pol. 110114, WA07-D1500D (1).
Distribution. Japan.
Onuphis opalina (Verrill, 1873)
Nothria opalina Verrill, 1873: 102.
Onuphis opalina: Fauchald, 1982: 50 51, fig. 14b, tab. 16; Imajima, 1986: 97 99, fig. 3a s; Imajima, 1997a: 182; Maekawa
and Hayashi, 1999: 188 189.

Material. NSMT-Pol. 109735, WA05-EF250D (12), NSMT-Pol. 109736, WA05-FG250D(1), NSMT-Pol. 109737, WA05-FG510D (10), NSMT-Pol. 109738, WA05-G1500D (7), NSMTPol. 109739, WA05-GH510D (64), NSMT-Pol. 109746, WA05-H900D (1); NSMT-Pol. 109740,
WA06-D210D (31), NSMT-Pol. 109741, WA06-F1500D (1), NSMT-Pol. 109742, WA06-F1500D
II (3), NSMT-Pol. 109743, WA06-G1200D (3), NSMT-Pol. 109744, WA06-GH480D (7), NSMTPol. 109745, WA06-H1500D (2); NSMT-Pol. 110115, WA07-D210D (105).
Distribution. Atlantic Ocean; northern Japan.
Onuphis taraba Maekawa and Hayashi, 1989
Onuphis taraba Maekawa and Hayashi, 1989: 79 81, fig. 11a n; Maekawa and Hayashi, 1999: 195 196, fig. 11a j.

Material. NSMT-Pol. 109747, WA06-E510D (5), NSMT-Pol. 109748, WA06-F1500D II (1),
NSMT-Pol. 109749, WA06-GH480D (21), NSMT-Pol. 109750, WA06-H250D (3), NSMT-Pol.
109751, WA06-H1500D (5).
Distribution. Japan.

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Genus Paradiopatra Ehlers, 1887
Paradiopatra crassa Imajima, 1999
Paradiopatra crassa Imajima, 1999:75 77, figs. 41a h, 42a k; Imajima, 2001a: 73.

Material. NSMT-Pol. 110116, WA07-D210D (1).
Distribution. Japan.
Paradiopatra gracilis sp. nov.
(Figs. 25A G, 26A O, 27A E)
Type material. Holotype—NSMT-Pol. H 506, Stn. WA06-F1500D II, off Sanriku, 37˚38.92´N,
142˚34.09´E 37˚39.39´N, 142˚34.32´E, 1466 1471 m, Nov. 1, 2006. Paratypes—NSMT-Pol. P
507, Stn. WA06-F1500D II, same locality as holotype (6); NSMT-Pol. P 508, Stn. WA06-F1500D,
off Sanriku, 1511 1508 m, Nov. 1, 2006 (20); NSMT-Pol. P 509, Stn. WA07-A1500D, off Sanriku,
1402 1377 m, Oct. 11, 2007 (8).
Other material. NSMT-Pol. 110554, WA05-E1000D (10), NSMT-Pol. 110555, WA05H900D (9); NSMT-Pol. 110556, WA06-D210D (2), NSMT-Pol. 110557, WA06-E1200D (18),
NSMT-Pol. 110560, WA06-G900D (576), NSMT-Pol. 110561, WA06-G1200D (72), NSMT-Pol.
110562, WA06-H1500D (3); NSMT-Pol. 110564, WA07-C1500D (3).
Description. Most specimens missing posterior ends; a complete largest specimen 68 mm
long, 1.2 mm wide for 144 setigers. Holotype lacking posterior end 39 mm long, 1.4 mm wide
including parapodia for 66 setigers. Body cylindrical through first 4 setigers, then dorsoventrally
flattened. Color in preserved specimen pale, without pigmentation exept a part of prostomium.
Prostomium suboval, with pair of clavate frontal lips and a brown oval pattern on anterodorsal and posterodorsal parts of prostomium. Ceratophores of antennae arranged in semicircle, with
3 5 proximal rings and a long distal ring. Styles of palps reaching to setiger 2, lateral antennae tosetiger 5, median antenna to setiger 3. Eyes and nuchal grooves not observed. Peristomium slightly shorter than following setigers; peristomial cirri absent (Fig. 25A C).
Anterior 3 pairs of parapodia slightly modified, not prolonged. Parapodia 1 and 2 similar insize and shape, with rounded presetal lobes and spindle-shaped postsetal lobes with broad bases,
as long as ventral cirrus; dorsal cirri digitiform, slightly longer than postsetal lobes (Fig. 25D E).
Parapodia 3 slightly smaller than former, with subulate postsetal lobes; dorsal cirri similar to thoseof first 2 setigers, but ventral cirri followed by transitory cirri (Fig. 25F), replaced by ventral glandular pads from setiger 4 (Fig. 25G).
Branchiae first present from setiger 8 (setigers 8 10 in paratypes), as simple filament (Fig.
26A), bifid from setiger 14 (setigers 12 14 in paratypes) (Fig. 26B), trifid in setigers 21 40 (setigers 22 36 in paratypes) (Fig. 26C), thereafter gradually diminishing to bifid (Fig. 26D) and single
filament posteriorly; last branchia on setiger 53 (setigers 50 56 in paratypes). Abranchial posterior
parapodia much smaller, with short dorsal cirri (Fig. 26E).
First 3 pairs of modified parapodia with single upper simple (Fig. 26F) or limbate setae (Fig.
26G) and 4 bidentate pseudocompound hooks with distinct second tooth; hoods long, acutely distally pointed (Fig. 26H K). Acicula number 2 3 in a fascicle, thick and distally bent, with slender
pointed end (Fig. 26L M). Parapodia 4 with upper limbate setae (Fig. 26N) and lower cultriform
setae (Fig. 26O) replaced by pseudocompound hooks. Following setigers with limbate setae, slender pectinate setae (Fig. 27A) and bidentate subacicular hooks (Fig. 27B); pectinate setae with
about 20 fine teeth present from setiger 9; subacicular hooks first present on setiger 9, usually 2 infascicle.

Deep-sea Benthic Polychaetes off Northern Honshu

97

Fig. 25. Paradiopatra gracilis sp. nov. —A, anterior end, dorsal view, × 25; B, anterior end of paratype, lateralview, × 31; C, prostomium and ceratophores of antennae, frontal view, × 48; D, left parapodium from setiger
1, anterior view, × 75; E, same from setiger 2, same view, × 75; F, same from setiger 3, same view, × 75; G,right parapodium from setiger 4, posterior view, × 75.

Mandibles fragile, united at distal end for short distance (Fig. 27C). Maxillary formula: Mx
I=1+1, Mx II=6­8+8, Mx III=7­8+0, Mx IV=7+7­8, Mx V=1+1 (Fig. 27D).
Pygidium small, bearing 2 pairs of ventrally inserted anal cirri, upper pair longer than lower
pair (Fig. 27E).
Remarks. This species possesses characters that fit neither Paradiopatra Ehlers, 1887 nor

98Minoru Imajima

Fig. 26. Paradiopatra gracilis sp. nov. —A, left parapodium from setiger 9, posterior view, × 70; B, same from setiger 16, same view, × 70; C, same from setiger 30, same view, × 70; D, same from setiger 45, same view, × 70;
E, same from setiger 65, same view, × 70; F, upper simple seta from setiger 1, × 578; G, limbate seta from setiger 3, × 287; H I, bidentate pseudocompound hooks from setiger 1, × 578; J, same from setiger 2, × 578; K,
same from setiger 3, × 578; L M, acicula, × 578; N, limbate seta from setiger 4, × 513; O, lower cultriform seta,
× 513.

Notonuphis Kucheruk, 1978 as they are presently defined. It has pseudocompound hooks withlong, pointed hoods, and 3 anterior pairs of modified parapodia. However, it has no peristomial
cirri, and has branchiae with simple at first, pectinate with 3 filaments where best developed. In
contrast, Paradiopatra has peristomial cirri, and Notonuphis has no branchiae.
According to Dr. Hannelore Paxton (personal communication) the value of peristomial cirriDeep-sea Benthic Polychaetes off Northern Honshu

99

Fig. 27. Paradiopatra gracilis sp. nov. —A, pectinate seta, × 680; B, subacicular hook, × 477; C, mandibles, × 67;
D, maxillae, × 64; E, posterior end, ventral view, × 30.

as a systematic character has been overstated in the past, and Notonuphis, and the new species, arebasically Paradiopatra species without peristomial cirri. Nataliya Budaeva is currently revising
the Paradiopatra complex and will be redefining the genus Paradiopatra in the near future.
Etymology. The specific name, gracilis, is Latin for slender, referring to the relatively slenderappearance of the body of this species.
Distribution. Japan (213 1511 m depth).
Paradiopatra striata (Uschakov, 1950)
Onuphis parva striata Uschakov, 1950: 193, fig. 25; Uschakov, 1955: 234, figs. 74b, 77j.
Paradiopatra parva striata: Paxton, 1986: 38.
Paradiopatra striata: Imajima, 1986: 110 114, fig. 9a t; Imajima, 1999: 84 88; Imajima, 2001a: 74.

Material. NSMT-Pol. 109752, WA05-DE380D (9), NSMT-Pol. 109753, WA05-EF250D(15), NSMT-Pol. 110566, WA05-FG250D (1), NSMT-Pol. 109754, WA05-FG510D (2), NSMTPol. 109755, WA05-G280 (1), NSMT-Pol. 109756, WA05-G1500D (1), NSMT-Pol. 109757,
WA05-GH380D (6); NSMT-Pol. 109758, WA06-A150D (1), NSMT-Pol. 109759, WA06-D210D(2), NSMT-Pol. 109760, WA06-E510D (6), NSMT-Pol. 109761, WA06-FG350D (1), NSMT-Pol.
109762, WA06-G900D (233), NSMT-Pol. 109763, WA06-GH480D (4), NSMT-Pol. 109764,
WA06-H250D (55), NSMT-Pol. 109765, WA06-H1500D (3); NSMT-Pol. 110117, WA07-C1500D(6), NSMT-Pol. 110118, WA07-D210D (62).
Distribution. Sea of Okhotsk; Japan.
Family Eunicidae Savigny, 1818
Genus Eunice Cuvier, 1817
Eunice mucronata Moore, 1903
Eunice mucronata Moore, 1903: 437 440, pl. 25, figs. 42 45; Miura, 1986: 305 308, figs. 30 31; Imajima, 2001a: 75; Imajima, 2007: 354 355, fig. 109.

100

Minoru Imajima

Material. NSMT-Pol. 109770, WA05-FG510D (2), NSMT-Pol. 109771, WA05-G425 (3),
NSMT-Pol. 109772, WA05-GH380 (2), NSMT-Pol. 109773, WA05-GH380D (22), NSMT-Pol.
109774, WA05-GH425 (1), NSMT-Pol. 109775, WA05-H380 (1), NSMT-Pol. 109776, WA05H750 (1); NSMT-Pol. 109777, WA06-A150D (5), NSMT-Pol. 109778, WA06-E1200D (1), NSMT-Pol. 109779, WA06-EF425D (1), NSMT-Pol. 110567, WA06-FG425 (2), NSMT-Pol. 109780,
WA06-G380 (2), NSMT-Pol. 109781, WA06-G650 (1), NSMT-Pol. 109782, WA06- G900D (5),NSMT-Pol. 109783, WA06-GH480D (56).
Distribution. Japan.
Family Lumbrineridae Malmgren, 1867
Genus Augeneria Monro, 1930
Augeneria bidens (Ehlers, 1887)
Lumbriconereis bidens Ehlers, 1887: 103, pl. 31, figs. 7 17.
Augeneria bidens: Orensanz, 1973: 372; Uebelacker, 1984a: 41-17 18, figs. 41-13, 14a h; Imajima, 2001b: 340, fig. 141;
Imajima, 2006: 373.
Augeneria tentaculata Monro, 1930: 140 142, fig. 52a k; Imajima and Higuchi, 1975: 7 8, fig. 1a j.

Material. NSMT-Pol. 109796, WA05-G1500D (1). NSMT-Pol. 110817, SO06-M4-B (1).
NSMT-Pol. 110840, KT07-29-H2 (1).
Distribution. Antarctic area; Argentine; Gulf of Mexico; Japan.
Genus Eranno Kinberg, 1865
Eranno abyssicola (Uschakov, 1950)(Figs. 28A H, 29A K)
Lumbriconereis abyssicola Uschakov, 1950: 195, fig. 27; Uschakov, 1965: 220, fig. 80G K.
Eranno abyssicola: Orensanz, 1990: 78; Frame, 1992: 194 195.

Material. NSMT-Pol. 110301, WA05-E1000D (2), NSMT-Pol. 110302, WA05-FG250D (1),
NSMT-Pol. 110303, WA05-FG450 (1), NSMT-Pol. 110304, WA05-GH510D (2); NSMT-Pol.
110305, WA06-F650D (1), NSMT-Pol. 110306, WA06-F750 (1), NSMT-Pol. 110310, WA06FG350D (1), NSMT-Pol. 110307, WA06-GH480D (1), NSMT-Pol. 110308, WA06-H480 (1),
NSMT-Pol. 110309, WA06-H1500D (1); NSMT-Pol. 110714, WA07-B410D (5), NSMT-Pol.
110715, WA07-D210D (2). NSMT-Pol. 110892, SO07-O1 (1), NSMT-Pol. 110841, SO07-O2(22), NSMT-Pol. 110893, SO07-O3 (1).
Description. Largest specimen missing posterior end 76 mm long, 4 mm wide for 234 setigers. Body cylindrical, without pigments.
Prostomium depressed triangle, shorter than basal wide, with nuchal organs. Peristomium
consisting of 2 apodous and asetigerous rings, ventrally developed into pair of buccal cushions;both segments partly fused ventrally (Fig. 28A B).
First parapodia comparatively large, with low presetal lobe and small flattened, bluntly
rounded postsetal lobe (Fig. 28C). Following about 30 parapodia subequal in size, but postsetal
lobes gradually reduced to small triangular ones (Fig. 28D). In about setiger 30, pre- and postsetal
lobes becoming subequal size; both lobes obliquely truncated (Fig. 28E). Median and posterior
parapodia becoming slender, postsetal lobes distally pointed (Fig. 28F G). In more posterior setigers postsetal lobes digitate, slightly larger than presetal ones (Fig. 28H).
First parapodia with 6 limbate setae (Fig. 29A) in upper bundle, and 7 hooded hooks and 2
limbate setae in lower bundle (Fig. 28C); hooks slender than following ones, with long hood andDeep-sea Benthic Polychaetes off Northern Honshu

101

Fig. 28. Eranno abyssicola (Uschakov). —A, anterior end, dorsal view, × 15; B, same, ventral view, × 15; C, rightparapodium from setiger 1, anterior view, × 66; D, same from setiger 20, same view, × 42; E, same from setiger 30, same view, × 42; F, same from setiger 46, same view, × 42; G, same from setiger 58, same view, × 42;
H, same from setiger 230, same view, × 42.

a fang surmounted by minute 10 teeth in tandem (Fig. 29B). Limbate setae in upper bundle becoming extremely elongated with slender tips; lower limbate setae reduced in number (Fig. 28D), andreplaced by hooded hooks from about setiger 30 (Fig. 28E). Limbate setae with whiplike tips on
setigers 30 60 (Fig. 28F). Hooded hooks on anterior parapodia with long hood (Fig. 29C), hooks
bulbous in median and posterior setigers, with large main fang and 8 11 irregular teeth in tandem(Fig. 29D H). Aciculae yellowish brown, numbering 2 3 in parapodia, with long acute tips (Fig.
29I) protruding through parapodial wall.

102

Minoru Imajima

Fig. 29. Eranno abyssicola (Uschakov). —A, limbate seta from setiger 1, × 145; B, hooded hook from setiger 1, × 287,
with detail of distal end, × 842; C, same from setiger 20, × 287, with detail of distal end, × 842; D, same from setiger
30, × 287; E, distal end of hook from setiger 30, × 578; F, hooded hook from setiger 58, × 287; G, distal part of same
hook, × 578; H, hooded hook from setiger 230, × 513; I, aciculum, × 405; J, mandibles, × 22; K, maxillae, × 22.

Mandibles long and slender, fused for two thirds of their length (Fig. 29J). Maxillary carriers
broad, slightly constricted (Fig. 29K); MI forceps long; MII two thirds the length of MI, with 4 or
5 asymmetrically arranged teeth, distal tooth largest, MII attached to MI by long broad basal support in addition to small connecting membrane; MIII bidentate; MIV large soft plate, tapering tosingle tooth; MV square, weakly sclerotized plates.
The species is newly added to the Japanese polychaetous fauna.
Distribution. Sea of Okhotsk; Japan.
Eranno bifurcata (McIntosh, 1885)
Lumbriconeris bifurcata McIntosh, 1885: 241 243, pl. 36, figs. 10 12, pl. 17, fig. 16, textfigs. 7 8; Izuka, 1912: 142 143;
Uschakov, 1955: 242, fig. 80A F.
Lumbrineris bifurcata: Imajima and Hartman, 1964: 262; Imajima and Higuchi, 1975: 26 28, fig. 10a n; Imajima, 1997a:
185 186.
Lumbrineris ezoensis Uchida, 1968: 599 600, fig. 4.
Eranno bifurcata: Imajima, 2001b: 342, fig. 143.

Material. NSMT-Pol. 110248, WA06-A1200D (11), NSMT-Pol. 110249, WA06-D210D (5),NSMT-Pol. 110250, WA06-D450D (2), NSMT-Pol. 110251, WA06-DE280D (4), NSMT-Pol.

Deep-sea Benthic Polychaetes off Northern Honshu

103110252, WA06-E510D (1), NSMT-Pol. 110253, WA06-EF425D (1); NSMT-Pol. 110716, WA07A450 (1), NSMT-Pol. 110254, WA07-A650 (15), NSMT-Pol. 110717, WA07-C350 (6), NSMTPol. 110718, WA07-D210 (1), NSMT-Pol. 110568, WA07-D900 (1).
Distribution. Pacific coast of North America; Japan.
Eranno tosaensis Imajima, 2001
Eranno tosaensis Imajima, 2001a: 76 78, figs. 20 21.

Material. NSMT-Pol. 110914, WA05-DE250D (1), NSMT-Pol. 110915, WA05-EF250D (1);
NSMT-Pol. 110916, WA06-D450D (1), NSMT-Pol. 110917, WA06-DE280D (1), NSMT-Pol.
110918, WA06-EF425D (1), NSMT-Pol. 110919, WA06-GH480D (1).
Distribution. Japan.
Genus Lumbrineris Blainville, 1828
Lumbrineris inflata Moore, 1911
Lumbrineris inflata Moore, 1911: 289 291, pl. 19, figs. 128 132, pl. 20, figs. 133 134; Berkeley and Berkeley, 1948: 97 98,
figs. 150 152; Day, 1967a: 435 436, fig. 17.16.d h; Imajima and Higuchi, 1975: 20 22, fig. 7a-l; Imajima, 1994: 112;
Imajima, 1997a: 185.
Lumbriconereis inflata: Uschakov and Wu, 1962c: 66.

Material. NSMT-Pol. 110119, WA07-B410D (4). NSMT-Pol. 110842, SO07-C7-B (1).
Distribution. Northeast Pacific; south to Gulf of California; Bering Sea; Yellow Sea; Japan.
Lumbrineris japonica (Marenzeller, 1879)
Lumbriconereis japonica Marenzeller, 1879: 137 138, pl. 5, fig. 3; Izuka, 1912: 139 140, pl. 14, figs. 17 18.
Lumbrineris japonica: Imajima and Hartman, 1964: 263 264; Fauchald, 1970: 91 92, pl. 14, figs. e f: Imajima and Higuchi,
1975: 30 32, fig. 12a n; Imajima, 2001a: 78 79.

Material. NSMT-Pol. 109797, WA05-GH510D (2); NSMT-Pol. 109798, WA06-F650D (1),
NSMT-Pol. 109799, WA06-F750 (1), NSMT-Pol. 109800, WA06-G900D (9): NSMT-Pol. 110120,
WA07-A1500D (1). NSMT-Pol. 110843, SO07-C7-B (7). NSMT-Pol. 110836, KT07-29-H-1 (2),
NSMT-Pol. 110837, KT07-29-H-2 (1), NSMT-Pol. 110835, KT07-29-M-3-2 (2).
Distribution. Indo Pacific areas; Pacific of California south to western Mexico; Japan.
Lumbrineris latreilli (Audouin and Milne-Edwards, 1834)
Lumbriconereis latreilli: Fauvel, 1923: 431, fig. 171m r.
Lumbrineris latreilli: Hartman, 1944b: 158 159, pl. 9, figs. 213 216; Pettibone, 1963: 258 260, fig. 67a c; Fauchald, 1970:
94 97, pl. 15, figs. f h; Hartmann-Schröder, 1971: 253 255, fig. 84a h; Imajima and Higuchi, 1975: 32 36, fig. 13a m;
Imajima, 2001a: 79.
Lumbrinereis latreilli: Berkeley and Berkeley, 1948: 98 99, figs. 154 156.

Material. NSMT-Pol. 109801, WA05-E1000D (1), NSMT-Pol. 109802, WA05-EF250D (6),
NSMT-Pol. 109803, WA05-FG250D (1), NSMT-Pol. 109804, WA05-GH510D (16); NSMT-Pol.
109805, WA06-EF425D (2), NSMT-Pol. 109806, WA06-GH480D (57), NSMT-Pol. 109807,
WA06-H1500D (1); NSMT-Pol. 110121, WA07-B410D (2), NSMT-Pol. 110122, WA07-C1500D(2), NSMT-Pol. 110460, WA07-D210D (1). NSMT-Pol. 110844, SO07-C7-B (24). NSMT-Pol.
110838, KT07-29-M-1 (4), NSMT-Pol. 110839, KT07-29-M-3-1 (1).

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Distribution. Atlantic, Pacific and Indian oceans; Mediterranean Sea; Japan.
Genus Ninoe Kinberg, 1865
Ninoe palmata Moore, 1903
Ninoe palmata Moore, 1903: 456 457, pl. 26, figs. 68 71; Izuka, 1912: 137 139; Uschakov and Wu, 1962a: 120, pl. 4z o;
Imajima and Hartman, 1964: 264; Imajima, 1968: 141; Imajima and Higuchi, 1975: 10 14, fig. 3a m; Imajima, 1997a:
184; Imajima, 2006: 374.

Material. NSMT-Pol. 109808, WA05-FG250D (1); NSMT-Pol. 109809, WA06-E1200D (1).
NSMT-Pol. 110845, SO07-K3 (3). NSMT-Pol. 110846, SO07-O1 (1).
Distribution. Yellow Sea; Japan.
Genus Paraninoe Levenstein, 1977
Paraninoe simpla Moore, 1905
Paraninoe simpla Moore, 1905: 547 549, pl. 35, fig. 30, pl. 36, figs. 39 44; Imajima, 2001b: 399, fig. 160.
Lumbrineris abyssalis Imajima and Higuchi, 1975: 16 20, fig. 6a r.

Material. NSMT-Pol. 109810, WA06-F650D (1); NSMT-Pol. 110123, WA07-A1500D (1),
NSMT-Pol. 110124, WA07-C1500D (2). NSMT-Pol. 110847, SO07-C7-B (1). NSMT-Pol. 110849,
KT07-29-H-2 (1), NSMT-Pol. 110848, KT07-29-M-2 (1).
Distribution. Alaska; Japan.
Genus Scoletoma Blainville, 1828
Scoletoma longifolia (Imajima and Higuchi, 1975)
Lumbrineris longifolia Imajima and Higuchi, 1975: 24 26, fig. 9; Imajima, 1994: 112.
Lumbriconereis debilis Uschakov and Wu, 1962c: 65, pl. 3k m.

Material. NSMT-Pol. 110808, WA06-GH480D (16). NSMT-Pol. 110850, SO07-C7-B (2).
NSMT-Pol. 110851, SO07-K2 (43), NSMT-Pol. 110852, SO07-O3 (1). NSMT-Pol. 110853, KT0729-H-1 (1).
Distribution. Japan; Yellow Sea.
Scoletoma nipponica (Imajima and Higuchi, 1975)
Lumbrineris nipponica Imajima and Higuchi, 1975: 22 24, fig. 8a m; Imajima, 1992: 129; Imajima, 1994: 112.
Scoletoma nipponica: Imajima, 2001b: 363, fig. 164; Imajima, 2006: 375.

Material. NSMT-Pol. 110311, WA07-B410D (1), NSMT-Pol. 110312, WA07-C1500D (2).
Distribution. Japan.
Family Oenonidae Kinberg, 1865
Genus Drilonereis Claparède, 1870
Drilonereis falcata japonica Imajima, 1964
(Figs. 30A G, 31A J)
Drilonereis falcata japonica Imajima, 1964: 244, figs. 21 27.

Material. NSMT-Pol. 110719, WA05-EF250D (1), NSMT-Pol. 110720, WA05-FG510D (1);Deep-sea Benthic Polychaetes off Northern Honshu

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NSMT-Pol. 110721, WA06-D450D (1), NSMT-Pol. 110722, WA06-DE280D (1), NSMT-Pol.
110723, WA06-E480 (1), NSMT-Pol. 110724, WA06-E1200D (1), NSMT-Pol. 110725, WA06GH480D (2), NSMT-Pol. 110726, WA06-H1500D (2); NSMT-Pol. 110727, WA07-A1500D (2),
NSMT-Pol. 110728, WA07-B410D (2), NSMT-Pol. 110729, WA07-C1500D (1).
Description. Largest specimen missing posterior end 140 mm long, 3 mm wide including
parapodia for 194 setigerous segments. Body evenly reddish brown, cylindrical, with conspicuous
parapodia throughout.
Prostomium spatulate, depressed conical, slightly longer than wide, with median dorsal furrow; nuchal organs inconspicuous, without eyes (Fig. 30A B). Peristomium consisting of 2 rings,
each as long as following setigers and first ring slightly wider than prostomial base.
Parapodia conspicuous and of subequal shape throughout length of body, slightly small infirst 2 setigers (Fig. 30C), presetal lobes truncate, postsetal lobes conical, pointing dorsad, both
lobes divergent; notopodial papilla minute (Fig. 30D G). Pharynx armed with mandibles and 5
pairs of maxillae. Mandibles well sclerotized, roughly triangular, about half as long as M I, freefrom one another (Fig. 31A). Maxillae with slender carriers, about 1.5 times as long as M I,

Fig. 30. Drilonereis falcata japonica Imajima. —A, anterior end, dorsal view, × 20; B, same, lateral view, × 20; C,
right parapodium from setiger 1, anterior view, × 67; D, same from setiger 5, anterior view, × 67; E, leftparapodium from setiger 50, same view, × 67; F, same from setiger 100, posterior view, × 67; G, right parapodium from setiger 190, anterior view, × 67.

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Fig. 31. Drilonereis falcata japonica Imajima. —A, mandibles, × 40; B, maxillae, × 40; C, right MII, × 80; D, left
MII, × 80; E, right MIII and MIV, × 80; F, upper limbate seta from anterior parapodium, × 228; G, lower
limbate seta from same, × 228, with detail of part, × 642; H, upper limbate seta from posterior parapodium, ×228; I, lower limbate seta from same, × 228; J, acicular spine, × 338.

anteriorly fused over short distance; ventral unpaired piece short, oval (Fig. 31B). M I strongly
falcate, with 3 or 4 large basal teeth; M II with 8 teeth, subdistal one smallest (Fig. 31C D); M III
with 4 teeth, 1 large and 3 small (Fig. 31E); M IV and M V each with one pointed tooth (Fig. 31B,
E). Limbate setae weakly geniculate, with finely serrated wings, setae in upper bundle longer thanlower ones (Fig. 31F I). Acicular spine (Fig. 31J) first present from setigers 1 3, protruding furthest in posterior setigers, and longer than postsetal lobes in middle and posterior setigers (Fig.
31E G).
Remarks. The subspecies is characterized in the following respects; (1) the pharynx is armed
with 5 pairs of maxillae, and mandibles are present, (2) the parapodia are well developed throughout length of the body, (3) acicular spines are first present from setigers 1 5.
Distribution. Japan.

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Drilonereis filum (Claparède, 1868)(Figs. 32A H, 33A F)
Lumbriconereis filum Claparède, 1868: 454, pl. 9, fig. 1.
Drilonereis filum: Fauvel, 1923: 436, fig. 174a h; Orensanz, 1974: 395 397, fig. 6; Hartman, 1968: 799 800, figs. 1 2.

Material. NSMT-Pol. 110730, WA05-EF250D (1).
Description. Specimen missing posterior end for 81 setigerous segments 53 mm long, 4 mm
wide including parapodia. Body almost cylindrical, with conspicuous parapodia throughout. Colorin spirit brown.
Prostomium spatulate, wider than long, rounded anteriorly, with median dorsal furrow. Nuchal

Fig. 32. Drilonereis filum (Claparède). —A, anterior end, dorsal view, × 15; B, same, lateral view, × 15; C, leftparapodium from setiger 1, anterior view, × 57; D, same from setiger 5, same view, × 57; E, right parapodium
from setiger 11, same view, × 57; F, left parapodium from setiger 40, same view, × 57; G, same from setiger
44, same view, × 57; H, same from setiger 82, same view, × 57.

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Fig. 33. Drilonereis filum (Claparède). —A, mandibles, × 35; B, maxillae, × 28, with detail of left and right maxillae II, × 76; C, right maxillae III V, × 76; D E, limbate setae, × 214; F, acicular spines from setiger 44, ×
236.

organs inconspicuous. Peristomium with 2 rings, the anterior one indistinctly separated from prostomium, about as wide as prostomial base and slightly shorter than second ring and subsequent
setigers (Fig. 32A B).
Parapodia conspicuous and of equal shape throughout length of body, slightly small in few
anterior setigers; presetal lobes truncate, postsetal lobes conical pointing dorsad; notopodial papilla minute (Fig. 32C H).
Pharynx armed with mandibles and 5 pairs of maxillae. Mandibles well sclerotized, roughly
triangular, fused along short distal area (Fig. 33A). Maxillae with long, slender carriers and lanceolate unpaired ventral piece; M I large, strongly falcate, basally smooth; M II with 8 to 10 teeth,
subdistal one smallest; M III with 4 teeth, 1 large and 3 small; M IV and M V each with one
pointed tooth (Fig. 33B C). Setae narrowly limbate with smooth wings, more straight and slender(Fig. 33D E). Acicular spine first present from setiger 10, usually single, sometimes 2 per parapodium (Fig. 33F).
The species is newly added to the Japanese polychaetous fauna.
Distribution. Mediterranean Sea; California; Japan.

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Genus Notocirrus Schmarda, 1861
Notocirrus japonicus (Okuda, 1939)
Arabella geniculata var. japonica Okuda, 1939: 237 238, textfig. 10.
Notocirrus japonicus: Imajima and Hartman, 1964: 266 267.

Material. NSMT-Pol. 109811, WA05-EF250D (1), NSMT-Pol. 109812, WA05-GH510D (1);
NSMT-Pol. 109813, WA06-A150D (1), NSMT-Pol. 109814, WA06-DE280D (1), NSMT-Pol.
109815, WA06-G1200D (1); NSMT-Pol. 110757, WA07-D210D (1).
Distribution. Japan.
Order Orbiniida
Family Orbiniidae Hartman, 1942
Genus Califia Hartman, 1957
Califia calida Hartman, 1957
Orbiniid n. g. and sp. Hartman, 1955: 179.
Califia calida Hartman, 1957: 306 308, pl. 42, figs. 1 3; Hartman, 1969: 17 18; Blake, 1996a: 11 12, fig. 1.3; Imajima,
1997a: 191, fig. 12a h.

Material. NSMT-Pol. 110165, WA05-GH510D (3).
Distribution. San Pedro Basin; California; Japan.
Genus Leitoscoloplos Day, 1977
Leitoscoloplos pugettensis (Pettibone, 1957)
Scoloplos (Scoloplos) pugettensis Pettibone, 1957: 162.
Leitoscoloplos pugettensis: Mackie, 1987: 8 9, fig. 8; Blake, 1996a: 9 10, fig. 1.2; Imajima, 1997a: 188.

Material. NSMT-Pol. 109669, WA06-DE280D (9).
Distribution. Puget Sound; British Columbia; Japan.
Genus Naineris Blainville, 1828
Naineris japonica sp. nov.
(Figs. 34A E, 35A G)
Type material. Holotype—NSMT-Pol. H 510, Stn. WA07-B410D, off Sanriku, 40˚16.88´N,
142˚13.51´E 40˚17.11´N, 142˚13.46´E, 416 416 m, Oct. 13, 2007. Paratypes—NSMT-Pol. P 511,
Stn. WA06-EF425D, off Sanriku, 420 424 m, Nov. 21, 2006 (2); NSMT-Pol. P 512, Stn. WA07C350D, off Sanriku, 355 354 m, Oct. 15, 2007 (2).
Description. All specimens missing posterior ends, holotype of largest one 23 mm long, 3.3mm wide including parapodia for 44 setigerous segments. Body greatly expanded and dorsoventrally flattened in thoracic region, parapodia elevated, forming distinct mid-dorsal trough; thoracic
segments less crowded than those in abdomen. Thorax consisting of peristomium and 17 setigers.
Prostomium broadly spatulate and truncated at its frontal margin; eyes absent; nuchal organs
not evident (Fig. 34A B). Peristomium uniannulate, achaetous, and forming posterior lip of large
mouth (Fig. 34C). Branchiae first present from setiger 5; each branchia long, digitate, pointed apically in thoracic region (Fig. 34E), then becoming flattened, with short conical tips in abdominal
region (Fig. 35B). Thoracic notopodia with long, digitate postsetal lobes. Thoracic neuropodia

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Fig. 34. Naineris japonica sp. nov. —A, anterior end, dorsal view, × 17; B, prostomium and first setiger, same view,
× 25; C, same, ventral view, × 25; D, right thoracic parapodium from setiger 1, anterior view, × 63; E, left
thoracic parapodium from setiger 8, same view, × 40.

with a conical postsetal lobe at middle part of postsetal ridge (Fig. 34D, 35A). Abdominal notopodia slender, distally tapering postsetal lobes; abdominal neuropodia bilobed, with infra-acicularlobe longest (Fig. 35B). Thoracic notosetae camerated capillaries (Fig. 35C); abdominal segmentswith capillaries and furcate setae (Fig. 35G). Thoracic neurosetae include camerated capillaries
and ridged uncini with partial transparent hood (Fig. 35D); those arranged in 4 5 rows with 10 15
in a row at maximum developed (Fig. 34E); uncini in anterior row of fascicle shorter (Fig. 35E)
than posterior ones (Fig. 35F). Uncini in first setiger circularly arranged in a row around neuropodial lobe (Fig. 34D). Abdominal neurosetae include fine, delicate capillaries.
Remarks. Naineris japonica resembles N. grubei (Gravier, 1909) in the shape of the prostomium and having 17 thoracic setigerous segments. However, N. japonica differs from N. grubei as

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Fig. 35. Naineris japonica sp. nov. —A, left thoracic parapodium from setiger 8, posterior view, × 42; B, left abdominal parapodium from setiger 26, anterior view, × 42; C, part of notoseta, × 680; D, neuropodial uncinus
from setiger 1, × 432; E, neuropodial uncinus in anterior row of fascicle, × 370; F, same in posterior row offascicle, × 370; G, furcate seta from abdominal notopodium, × 990.

follows; (1) the branchiae are present from setiger 5 rather than setiger 7, (2) the thoracic neuropodial ridge has a short lobe at its midlength rather than with a short lobe at its superior edge and (3)abdominal notopodia have capillaries and furcate setae rather than without furcate setae.
Etymology. The species is named because it was collected from Japanese waters.
Distribution. Japan (354 424 m depth).
Genus Phylo Kinberg, 1866
Phylo felix Kinberg, 1866
(Fig. 36A I)
Phylo felix Kinberg, 1866: 251 252; Hartman, 1948b: 105 106, pl. 15, fig. 10; Hartman, 1953: 37 38; Hartman, 1957:
262 265, pl. 23, figs.1 7.

Material. NSMT-Pol. 110435, WA05-DE380D (1), NSMT-Pol. 110436, WA05-EF450D (2),
NSMT-Pol. 110437, WA05-GH380D (1); NSMT-Pol. 110438, WA06-EF425D (1), NSMT-Pol.

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110439, WA06-GH480D (1); NSMT-Pol. 110440, WA07-B410D (12), NSMT-Pol. 110441, WA07C350D (10).
Description. All specimens missing posterior ends; largest one 29 mm long, about 3 mm
wide for 65 setigers. Dorsum elevated through 4th setiger and depressed to concave in segments
farther back. Thorax divided into anterior and posterior regions: anterior encompassing setigers
1 10; posterior including setigers 11 17; transitional to abdominal region on setiger 18.

Fig. 36. Phylo felix Kinberg. —A, anterior end, lateral view, × 16; B, left parapodium from setiger 7, anterior view,
× 32; C, same, posterior view, × 32; D, left parapodium from setiger 13, anterior view, × 24; E, thoracic neuropodial uncinus, × 493; F G, modified spines from posterior thoracic segment, × 145; H, right parapodium
from setiger 30, anterior view, × 47; I, furcate seta from abdominal notopodium, × 697.

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Prostomium small, conical, lateral margins with pigment in lateral depressions. Thoracic notopodia with long, simple postsetal lobes and fascicles of pointed setae (Fig. 36A). First 10 thoracic neuropodia with parapodial ridges flanked by postsetal fringe; subpodial or ventral papillae
absent. First neuropodia with 2 or 3 lobes, gradually increasing to 9 to 12 lobes on middle andposterior thoracic segments (Fig. 36A C). Ventral fringe first present on 11th setiger and increasing
to 20 lobes on 15th setiger (Fig. 36D), absent after thoracic segment 20.
Branchiae present from setiger 5, continuing to end of body; branchia with wide base, lingulate, laterally fimbriated. Thoracic notosetae slender camerated capillaries. Thoracic neurosetae of
two kinds; 3 to 5 rows of yellow uncini distally curved and transversely ridged, distally hooded(Fig. 36E); setae in posterior row longer, distally pointed, with transverse series of fine spinelets.
From 11th setiger uncini replaced by dark brown modified spearlike spines (Fig. 36F G),
number 4 to 6 in anterior transverse row. These spines present in setigers 11 to 16. Uppermost
spine projects from neuropodium, more conspicuous than others; posterior pointed setae and inferior uncinal rows remain as in front but more limited in number. Anterior abdominal parapodiawith 2 3 postsetal neuropodial lamellae; remaining parapodia with long, slender notopodial postsetal lobes and shorter neuropodial lobes, interramal cirri and ventral cirrus (Fig. 36H). Interramal
cirrus present as long simple lobe, from second last thoracic or not before first abdominal segment,
continued back through long abdominal region. Abdominal notopodia with 1 3 furcate setae (Fig.
36I) in bundle, accompanied by pointed setae. Abdominal neuropodia with 3 6 slender pointed
setae.
The species is newly added to the Japanese polychaetous fauna.
Distribution. Brazil; Patagonia; California; Mexico; East Falkland Islands; Japan.
Phylo fimbriatus (Moore, 1903)
Aricia fimbriata Moore, 1903: 464 467, pl. 24, figs. 31 35; Okuda, 1937a: 99 100, textfigs. 1 2.
Phylo fimbriatus: Imajima and Hartman, 1964: 276 277; Imajima, 1997a: 189.

Material. NSMT-Pol. 110442, WA05-GH380D (1), NSMT-Pol.0110443, WA05-GH510D(1).
Distribution. Japan.
Phylo nudus (Moore, 1911)
Aricia nuda Moore, 1911: 311 315, pl. 21, figs. 172 176; Fauvel, 1953: 303 304, fig. 157a d.
Phylo nudus: Hartman, 1957: 268; Hartman, 1969: 39, figs. 1 4; Sun and Yang, 1987: 158, fig. 5; Blake, 1996a: 13, fig. 1.4;
Imajima, 1997a: 188 189, fig. 11a l.

Material. NSMT-Pol. 110444, WA05-E1000D (1); NSMT-Pol. 110445, WA06-A1200D (1),
NSMT-Pol. 110446, WA06-D450D (1), NSMT-Pol. 110447, WA06-EF425D (1); NSMT-Pol.
110448, WA07-B410D (2), NSMT-Pol. 110449, WA07-D900 (10).
Distribution. Southern California; Burma; East and South China seas; Japan.
Genus Scoloplos Blainville, 1828
Scoloplos (Leodamas) robustus (Kinberg, 1866) comb. nov.
(Figs. 37A E, 38A H)
Alcandra robusta Kinberg, 1866: 251; Kinberg, 1910: 62 63, pl. 24, fig. 6; Hartman, 1948b: 106; Hartman, 1957: 295, pl. 31,
fig. 5.

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Material. NSMT-Pol. 110804, WA05-GH380D (1).
Description. Body divided into 3 parts, and total consisting of 93 mm long, 5 mm wide for
130 setigers. Prostomium short, bluntly conical. Peristomium broad, achaetous, anteriorly bearing
a pair of dorso-lateral nuchal organs; eyes absent (Fig. 37A).
Thorax with 15 setigers. Thoracic notopodia with lanceolate lobes (Fig. 37B D); neuropodial
lobes digitate (Fig. 37B) or shorter and broader (Fig. 37C D). Postsetal lobes of notopodia gradually increase in size along thorax. No subpodal or stomach papillae.
Branchiae present from setiger 5, initially small, digitate, rapidly increasing in size, becoming
long, slender and triangular. Branchiae laterally ciliated, generally longer than notopodial lamellae.
In abdomen notopodial postsetal lobes abruptly increase in size, becoming more foliaceous(Fig. 37E), then gradually reduced posteriorly (Fig. 38A B). Anterior neuropodia weakly bilobed;
rudimentary inner lobe lost, uniramous condition evident in later setigers (Figs. 37E, 38A B).

Fig. 37. Scoloplos (Leodamas) robustus (Kinberg). —A, anterior end, dorsal view, × 15; B, right parapodium fromsetiger 1, anterior view, × 44; C, left parapodium from setiger 5, same view, × 23; D, left parapodium fromsetiger 15, same view, × 23; E, right parapodium from setiger 25, same view, × 23.

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Fig. 38. Scoloplos (Leodamas) robustus (Kinberg). —A, right parapodium from setiger 40, anterior view, × 26; B,
left parapodium from setiger 70, same view, × 33; C, furcate seta, × 717; D, uncinus in anterior row, × 285; E,
same in posterior row, × 285; F, part of uncinus, × 717; G, protruding aciculum from posterior notopodium, ×224; H, same from posterior neuropodium, × 224.

Subpodal papillae and interramal cirri lacking.
Thoracic notosetae camerated capillaries and 4 5 furcate setae (Fig. 38C); furcate setae present from the first parapodia. Thoracic neurosetae including camerated capillaries and conspicuoustransverse, palisaded rows of curved uncini. Uncini occurring from setiger 1, those in anteriormost
row shorter and more sharply bent near tip (Fig. 38D) than posteriormost ones (Fig. 38E); all conspicuously denticulated along cutting edge (Fig. 38F).
Abdominal notopodia with long camerated capillaries, 8 10 furcate setae and a protruding
aciculum (Figs. 37F, 38B, G); neuropodia with fine, camerated capillaries and a protruding aciculum occurring from posterior setigers (Fig. 38B, H).
Remarks. As mentioned by Hartman (1957), Alcandra robusta can be referred to Scoloplos(Leodamas) Kinberg by the characteristics of the prostomium, branchiae and setae. However, thetype specimen is only thoracic segments, so that the abdominal region is not known.
This species is characterized as follows: the thorax consists of 15 setigers; the branchiae arepresent from the fifth setigerous segment and continue on all other segments; the notopodia of thethorax and abdomen have camerated capillaries and furcate setae; the thoracic neuropodia have

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camerated capillaries and a number of uncini; the posterior noto- and neuropodia have singly occurring curved acicula that extends from the both rami; the subpodal papillae and interramal cirri
are absent.
The species is newly added to the Japanese polychaetous fauna.
Distribution. Off Brazil; Japan.
Scoloplos (Scoloplos) similis Mackie, 1987
(Fig. 39A F)
Scoloplos similis Mackie, 1987: 24, fig. 24.

Material. NSMT-Pol. 110805, WA05-DE380D (3), NSMT-Pol. 109591, WA05-FG250D(18); NSMT-Pol. 109592, WA06-DE280D (5), NSMT-Pol. 110806, WA06-E510D (2), NSMT-Pol.
110807, WA06-GH480D (10).
Description. Largest specimen missing posterior end 11 mm long, about 0.8 mm wide including parapodia for 42 setigers.
Prostomium pointed, longer than wide; eyes absent; paired nuchal slits laterally. Peristomial
ring about as long as setiger 1 (Fig. 39A). Thorax with 14 setigers. Thoracic notopodial postsetal
lobes single, slender triangular; thoracic neuropodial postsetal lobes mammiform to triangular for
first 12 setigers (Fig. 39B), bifurcate thereafter (Fig. 39C).
Branchiae present from setiger 11, initially short and triangular (Fig. 39C), then becoming
larger and erect, subdistally swollen, ciliated along inner margin, approximately same length as
notopodial lobes (Fig. 39D). More posteriorly, branchiae becoming somewhat strap-like, withblunt tips (Fig. 39E).
Abdominal notopodial postsetal lobes single, slender, digitate (Fig. 39D E). Abdominal neuropodia bilobed, inner lobe longer and more robust, with a distinct subpodal notch at insertion ofshort, subpodal flange (Fig. 39D E). Pair of small subpodal papillae present setigers 14 to 20 (Fig.
39D), thereafter only single papilla evident (Fig. 39E). Interramal cirri present on setigers 14 to 23
in cirriform shape (Fig. 39D).
Thoracic notosetae camerated capillaries. Thoracic neuropodia possess a number of camerated capillaries and about 10 serrate uncini with small hood (Fig. 39F); uncini appearing from firstparapodia. Abdominal notosetae all camerated capillaries, forked setae not seen. Abdominal neurosetae 2 3 camerated capillaries.
Remarks. The examined specimens differ from the type specimen from the South Africa insome characteristics in that, the setigerous number of the thoracic region (14 setigers instead of 15)
and the appearance of the branchiae (from setiger 11 instead of 12). It is likely that those are
merely variants of characters.
The species is newly added to the Japanese polychaetous fauna.
Distribution. South Africa; Japan.
Family Paraonidae Cerruti, 1909
Genus Aricidea Webster, 1879
Aricidea (Acmira) simplex Day, 1963
Aricidea suecica simplex Day, 1963a: 364 365, fig. 3a b.
Aricidea (Acmira) simplex: Blake, 1996b: 63 64, fig. 2.18; Imajima, 1997a: 193; Imajima, 2001b: 433, fig. 178; Lovell, 2002:
45 46, fig. 7A C.
Aricidea neosuecica nipponica Imajima, 1973: 263 265, fig. 5a f.

Deep-sea Benthic Polychaetes off Northern Honshu

117

Fig. 39. Scoloplos (Scoloplos) similis Mackie. —A, anterior end, dorsal view, × 34; B, left parapodium from setiger
5, posterior view, × 107; C, right parapodium from setiger 13, same view, × 107; D, right parapodium fromsetiger 17, anterior view, × 107; E, left parapodium from setiger 27, posterior view, × 107; F, serrate uncinus, ×680.

Material. NSMT-Pol. 109816, WA05-DE380D (3); NSMT-Pol. 110495, WA06-GH480D (3).
Distribution. South Africa; Atlantic Ocean; New Zealand; Bering Sea; California; Japan.
Aricidea (Aedicira) belgicae (Fauvel, 1936)
Paraonis belgicae Fauvel, 1936b: 29 31, fig. 3.

118

Minoru Imajima

Aricidea (Aedicira) belgicae: Hartman, 1957: 327; Imajima, 2001b: 435, fig. 180.
Aedicira belgicae: Hartman, 1965: 133 135; Imajima, 1973: 277 279, fig. 13a k.

Material. NSMT-Pol. 109817, WA05-GH510D (1); NSMT-Pol. 110496, WA06-DE280D(11), NSMT-Pol. 110497, WA06-G1200D (1), NSMT-Pol. 110498, WA06-GH480D (2), NSMTPol. 110499, WA06-H1500D (1).
Distribution. Antarctic Ocean; off northeastern South America; South Africa; Japan.
Aricidea (Allia) antennata Annenkova, 1934
Aricidea antennata Annenkova, 1934a: 658, figs. 2 3.
Aricidea uschakovi: Imajima, 1973: 256 258, fig. 1a k; Imajima, 2006: 377.
Aricidea (Allia) antennata: Blake, 1996b: 48 50, fig. 2.10.

Material. NSMT-Pol. 109818, WA05-GH510D (1); NSMT-Pol. 110501, WA06-GH480D(1).
Distribution. North Japan Sea; western Canada; Southern California; Japan.
Aricidea (Allia) sp.
Material. NSMT-Pol. 110612, KT07-29-H-1 (1).
Gens Cirrophorus Ehlers, 1908
Cirrophorus branchiatus Ehlers, 1908
Cirrophorus branchiatus Ehlers, 1908: 124 126, pl. 17, figs. 5 9; Day, 1963b: 423 424, textfig. 9l o; Day, 1967b: 563, fig.
24.3.a e; Imajima, 1973: 274 275, fig. 11a g.

Material. NSMT-Pol. 110613, KT07-29-M-3-3 (2).
Distribution. Southern Africa; British Isles; Japan.
Genus Levinsenia Mesnil, 1897
Levinsenia gracilis (Tauber, 1879)
Aonides gracilis Tauber, 1879: 115.
Levinsenia gracilis: Mesnil and Caullery, 1898: 135 137; Blake, 1996b: 33 34, fig. 2.1; Imajima, 1997a: 193.
Paraonis gracilis minuta Hartmann-Schröder, 1965: 197 198, figs. 181 182; Imajima, 1973: 284 285, fig. 16a f.

Material. NSMT-Pol. 110502, WA06-E1200D (1), NSMT-Pol. 110503, WA06-GH480D (1).
Distribution. Cosmopolitan in continental shelf and slope depths.
Genus Paradoneis Hartman, 1965
Paradoneis lyra (Southern, 1914)
Paraonis (Paraonides) lyra Southern, 1914: 94 95, pls. 9 10, fig. 22a g; Fauvel, 1927: 72 73, fig. 24a f.
Paraonides lyra lyra: Day, 1963b: 425.
Paradoneis lyra: Hartman, 1965: 140; Imajima, 1997a: 193 194; Imajima, 2006: 378.
Paraonides lyra: Imajima, 1973: 287 288, fig. 18a e.

Material. NSMT-Pol. 109819, WA05-DE380D (4), NSMT-Pol. 110505, WA05-GH510D(11); NSMT-Pol. 110504, WA06-GH480D (5).

Deep-sea Benthic Polychaetes off Northern Honshu

119Distribution. Western Ireland; Denmark; Black Sea; South Africa; Pacific of Southern California; Massachusetts; Japan.
Order Spionida
Family Poecilochaetidae Hannerz, 1956
Genus Poecilochaetus Claparède, 1875
Poecilochaetus granulatus Imajima, 1989
Poecilochaetus granulatus Imajima, 1989b: 94 99, figs. 17a e, 18a h, 19a i.

Material. NSMT-Pol. 110541, WA06-GH480D (1).
Distribution. Japan.
Family Spionidae Grube, 1850
Genus Laonice Malmgren, 1867
Laonice cirrata (Sars, 1851)
Nerine cirrata Sars, 1851: 207.
Spionides japonicus: Moore, 1907: 204 206, pl. 16, figs. 31 34.
Laonice cirrata: Fauvel, 1927: 38, fig. 12a e; Okuda, 1937b: 222; Imajima and Hartman, 1964: 281 282; Day, 1967b: 480,
fig. 18.6.h k; Imajima, 2006: 378 379.

Material. NSMT-Pol. 110156, WA05-DE250D (2), NSMT-Pol. 110157, WA05-DE380D(30), NSMT-Pol. 110158, WA05-E1000D (1), NSMT-Pol. 110159, WA05-EF450D (4), NSMTPol. 110160, WA05-FG250D (1), NSMT-Pol. 110161, WA05-FG510D (1); NSMT-Pol. 110487,
WA06-D450D (1), NSMT-Pol. 110488, WA06-DE280D (9), NSMT-Pol. 110489, WA06-E510D(14), NSMT-Pol. 110162, WA06-EF425D (1), NSMT-Pol. 110163, WA06-FG350D (12), NSMTPol. 110164, WA06-GH480D (7); NSMT-Pol. 110492, WA07-A250D (30), NSMT-Pol. 110493,
WA07-A1500D (1), NSMT-Pol. 110494, WA07-B410D (24).
Distribution. Northern Norway; north Atlantic and Pacific oceans; Japan.
Genus Prionospio Malmgren, 1867
Prionospio (Prionospio) bocki Söderström, 1920
Prionospio bocki Söderström, 1920: 234 235, figs. 142 143; Day, 1967b: 490; Maciolek, 1985: 336; Imajima, 1990: 122 124,
figs. 10a c, 11a j.

Material. NSMT-Pol. 110481, WA05-DE380D (2); NSMT-Pol. 110482, WA06-E510D (2),
NSMT-Pol. 110483, WA06-EF425D (1), NSMT-Pol. 110484, WA06-GH480D (2).
Distribution. Madagascar; Japan.
Prionospio (Prionospio) depauperata Imajima, 1990
Prionospio (Prionospio) depauperata Imajima, 1990: 114 118, figs. 6a d, 7a l; Imajima, 1994: 114; Imajima, 1997a: 194.

Material. NSMT-Pol. 110788, WA05-DE250D (5), NSMT-Pol. 110789, WA05-FG250D(198); NSMT-Pol. 110790, WA06-DE280D (39).
Distribution. Japan.

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Prionospio (Prionospio) dubia Day, 1961
Prionospio malmgreni var. dubia Day, 1961: 489 490, fig. 3j n.
Prionospio (Prionospio) dubia: Maciolek, 1985: 336 339, figs. 2 3; Imajima, 1990: 118 121, figs. 8a e, 9a h.

Material. NSMT-Pol. 109870, WA05-DE380D (1), NSMT-Pol. 109871, WA05-EF250D (1);
NSMT-Pol. 109873, WA06-GH480D (7), NSMT-Pol. 109872, WA06-H250D (8); NSMT-Pol.
110485, WA07-D210D (1).
Distribution. South Africa; North Carolina; Virginia; Bay of Biscay; Mediterranean Sea;
Sweden; Surinam; Japan.
Prionospio sp.
Material. NSMT-Pol. 110573, WA06-DE280D (6). NSMT-Pol. 110635, KT07-29-H-1 (1).
Genus Spiophanes Grube, 1860
Spiophanes japonicum Imajima, 1991
Spiophanes japonicum Imajima, 1991: 123 128, figs. 5a h, 6a h, 7a n; Imajima, 1997a: 195.

Material. NSMT-Pol. 109874, WA06-H250D (1).
Distribution. Japan.
Spiophanes kroyeri Grube, 1860
Spiophanes kroyeri Grube, 1860: 88.
Spiophanes kroeyeri: Fauchald, 1972a: 99, fig. 4c d; Light, 1977: 79 80, fig. 5d; Imajima, 1991: 118 123, figs. 2a d, 3a h,
4a o; Imajima, 1997a: 194; Imajima, 2006: 380.

Material. NSMT-Pol. 109875, WA05-E1000D (35), NSMT-Pol. 109876, WA05-GH510D(2), NSMT-Pol. 109877, WA05-H900D (2); NSMT-Pol. 109878, WA06-B750D (3), NSMT-Pol.
109879, WA06-E1200D (1), NSMT-Pol. 109880, WA06-F650D (1), NSMT-Pol. 109881, WA06G1200D (4), NSMT-Pol. 109882, WA06-GH480D (3), NSMT-Pol. 109883, WA06-H250D (1),
NSMT-Pol. 109884, WA06-H1500D (17); NSMT-Pol. 110486, WA07-D900 (1).
Distribution. Greenland; western Norway; Australia; Ross Sea; Japan.
Family Chaetopteridae Malmgren, 1867
Genus Spiochaetopterus Sars, 1853
Spiochaetopterus typicus Sars, 1856
Spiochaetopterus typicus: Uschakov, 1965: 271, fig. 106A D.

Material. NSMT-Pol. 110894, WA06-A450 (23), NSMT-Pol. 110895, WA06-A510 (3), NSMT-Pol. 110896, WA06-DE280D (1), NSMT-Pol. 110897, WA06-E510 (19), NSMT-Pol. 110898,
WA06-EF425D (5), NSMT-Pol. 110899, WA06-F1500D II (1), NSMT-Pol. 110900, WA06FG350D (3), NSMT-Pol. 110901, WA06-G900D (4), NSMT-Pol. 110902, WA06- G1200D (1),
NSMT-Pol. 110903, WA06-GH480D (156), NSMT-Pol. 110904, WA06-H250D (13), NSMT-Pol.
110905, WA06-H1500D (2); NSMT-Pol. 110906, WA07-A450 (21), NSMT-Pol. 110907, WA07A650 (1), NSMT-Pol. 110908, WA07-A1500D (31), NSMT-Pol. 110909, WA07-B310 (1), NSMT-Deep-sea Benthic Polychaetes off Northern Honshu

121Pol. 110910, WA07-B410D (147), NSMT-Pol. 110911, WA07-B1500D (4), NSMT-Pol. 110912,
WA07-D210D (1), NSMT-Pol. 110913, WA07-D1500D (5).
The species is newly added to the Japanese polychaetous fauna.
Distribution. Far Eastern seas; Japan.
Family Cirratulidae Carus, 1863
Genus Aphelochaeta Blake, 1991
Aphelochaeta sp.
Material. NSMT-Pol. 110636, WA05-FG510D (1), NSMT-Pol. 110637, WA05-G1500D (1);
NSMT-Pol. 110638, WA06-F650D (1), NSMT-Pol. 110639, WA06-G900D (1), NSMT-Pol.
110640, WA06-H250D (1). NSMT-Pol. 110641, KT07-29-H-1 (2).
Genus Chaetozone Malmgren, 1867
Chaetozone setosa Malmgren, 1867
Chaetozone setosa Malmgren, 1867a: 96, pl. 14, fig. 84; Uschakov, 1965: 279, fig. 110A F; Blake, 1996c: 274 276, fig. 8.1;
Imajima, 1997a: 197.

Material. NSMT-Pol. 110642, WA05-H250D (3). NSMT-Pol. 110643, KT07-29-H-1 (10).
Distribution. Arctic and subarctic seas; Japan.
Chaetozone spinosa Moore, 1903
Chaetozone spinosa Moore, 1903: 468 470, pl. 26, figs. 73 74; Hartman, 1969: 243 244; Imajima, 1997a: 197.

Material. NSMT-Pol. 110644, WA05-GH380D (1); NSMT-Pol. 110645, WA06-GH480D(1), NSMT-Pol. 110646, WA06-H250D (3).
Distribution. Southern California; Japan.
Chaetozone sp.
Material. NSMT-Pol. 110647, WA05-EF250D (6); NSMT-Pol. 110648, WA06-DE280D(15).
Genus Cirratulus Lamarck, 1801
Cirratulus cirratus (Müller, 1776)
Cirratulus cirratus: Fauvel, 1927: 94, fig. 33a g; Imajima and Hartman, 1964: 298; Hartmann-Schröder, 1971: 258 259, fig.
125; Imajima, 1997a: 197 198; Imajima, 2006: 381.

Material. NSMT-Pol. 110576, WA05-DE250D (2), NSMT-Pol. 110577, WA05-G280 (1),
NSMT-Pol. 110578, WA05-GH380 (4), NSMT-Pol. 110579, WA05-H310 (2), NSMT-Pol. 110580,
WA05-H350 (2), NSMT-Pol. 110581, WA05-H380 (6), NSMT-Pol. 110582, WA05-H510 (1);
NSMT-Pol. 110583, WA06-A150D (1), NSMT-Pol. 110584, WA06-B350 (1), NSMT-Pol. 110585,
WA06-DE450 (1), NSMT-Pol. 110586, WA06-G380 (1), NSMT-Pol. 110587, WA06-H310 (1);
NSMT-Pol. 110588, WA07-B310 (1), NSMT-Pol. 110589, WA07-B410 (5), NSMT-Pol. 110590,
WA07-B410D (1), NSMT-Pol. 110591, WA07-C310 (10), NSMT-Pol. 110592, WA07-C350D (1),
NSMT-Pol. 110593, WA07-C410 (1), NSMT-Pol. 110594, WA07-C450 (2). NSMT-Pol. 110598,

122

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KT07-29-M-3-1 (1).
Distribution. Western and southern Europe, central and southern California, Japan.
Genus Monticellina Laubier, 1961
Monticellina tesselata (Hartman, 1960)
Tharyx tesselata Hartman, 1960: 126 127, pl. 11, figs. 1 4; Reish, 1968: 86.
Monticellina tesselata: Blake, 1996c: 328 329, fig. 8.27; Imajima, 1997a: 198; Imajima, 2001a: 80 81.

Material. NSMT-Pol. 110595, WA05-DE380D (18); NSMT-Pol. 110597, WA06-E510D (2),
NSMT-Pol. 110596, WA06-GH480D (5).
Distribution. Central and southern California; Japan.
Genus Timarete Kinberg, 1866
Timarete sp.
Material. NSMT-Pol. 110649, WA06-F1500D II (1).
Order Cossurida
Family Cossuridae Day, 1963
Genus Cossura Webster and Benedict, 1887
Cossura spp.
Material. NSMT-Pol. 110506, WA05-EF250D (3), NSMT-Pol. 110507, WA05-FG250D (1),
NSMT-Pol. 110508, WA05-GH510D (3); NSMT-Pol. 110509, WA06-E510D (2), NSMT-Pol.
110510, WA06-G1200D (2), NSMT-Pol. 110511, WA06-GH480D (21); NSMT-Pol. 110512,
WA07-D210D (1). NSMT-Pol. 110857, KT07-29-H-1 (2), NSMT-Pol. 110856, KT07-29-M-1(1).
All specimens are missing characteristic posterior ends.
Order Flabelligerida
Family Flabelligeridae Saint-Joseph, 1894
Genus Brada Stimpson, 1854
Brada granulata Malmgren, 1867
Brada granulata Malmgren, 1867b: 194, pl. 12, fig. 71; Uschakov, 1965: 287, fig. 115A C.

Material. NSMT-Pol. 110667, WA07-C310 (1).
The species is newly added to the Japanese polychaetous fauna.
Distribution. Northern part of Sea of Japan; Sea of Okhotsk; Bering Sea; Japan.
Brada ochotensis Annenkova, 1922
Brada ochotensis Annenkova, 1922: 39; Uschakov, 1965: 287, fig. 115J.

Material. NSMT-Pol. 110668, WA07-B410D (1).
The species is newly added to the Japanese polychaetous fauna.
Distribution. Sakhalin; Bering Sea; Chuckchee Sea; Japan.

Deep-sea Benthic Polychaetes off Northern Honshu

123

Brada sachalina Annenkova, 1922
Brada sachalina Annenkova, 1922: 39; Uschakov, 1955: 310, fig. 115K L.
Brada mamillata: Imajima, 1964: 246 247, figs. 33 38 (not McIntosh, 1885).

Material. NSMT-Pol. 110664, WA05-GH510D (1); NSMT-Pol. 109820, WA06-B310D (1);
NSMT-Pol. 110665, WA07-B410D (21).
Distribution. Sakhalin; Japan.
Brada villosa (Rathke, 1843)
Siphonostoma villosum Rathke, 1843: 215, pl. 11, figs. 11 12.
Brada villosa: Haase, 1915: 203; Fauvel, 1927: 121; Okuda, 1937c: 53 54, pl. 2, fig. D; Uschakov, 1955: 311, fig. 115D H;
Imajima and Hartman, 1964: 302; Imajima, 1997a: 199; Blake, 2000a: 5 7, fig. 1.1.

Material. NSMT-Pol. 110669, WA06-E1200D (1), NSMT-Pol. 110670, WA06-F1500D (4),
NSMT-Pol. 110671, WA06-F1500D II (5), NSMT-Pol. 109821, WA06-H1500D (3); NSMT-Pol.
110672, WA07-C350D (1), NSMT-Pol. 110673, WA07-D1500D (4).
Distribution. Norway; Mediterranean Sea; Atlantic Ocean; Japan.
Genus Pherusa Oken, 1807
Pherusa plumosa (O. F. Müller, 1776)
Amphitrite plumosa O. F. Müller, 1776: 216.
Stylaroides plumosa: Okuda, 1937c: 52, pl. 2, fig. C.
Pherusa plumosa: Støp-Bowitz, 1948: 13; Imajima and Hartman, 1964: 303 304; Imajima, 1988: 127; Imajima, 1994: 118;
Imajima, 1997a: 198 199.

Material. NSMT-Pol. 109822, WA05-FG510D (4), NSMT-Pol. 109823, WA05-GH380D (1);
NSMT-Pol. 109824, WA06-E1200D (1), NSMT-Pol. 109825, WA06-F650D (1), NSMT-Pol.
109826, WA06-GH480D (3), NSMT-Pol. 109827, WA06-H1500D (1); NSMT-Pol. 110674,
WA07-D310 (1).
Distribution. Atlantic and Pacific oceans; Japan.
Family Acrocirridae Banse, 1969?
Genus Acrocirrus Grube, 1872
Acrocirrus heterochaetus Annenkova, 1934
Acrocirrus heterochaetus Annenkova, 1934b: 326 327, fig. 7; Hartman, 1948a: 38 39, fig. 11a c; Uschakov, 1965: 282, fig.
112A D.

Material. NSMT-Pol. 110513, WA06-A350 (1); NSMT-Pol. 110514, WA07-B410 (2), NSMT-Pol. 110515, WA07-C350D (1).
The species is newly added to the Japanese polychaetous fauna.
Distribution. Sea of Japan; Sea of Okhotsk; Alaska Peninsula; Japan.
Family Fauveliopsidae
Genus Fauveliopsis McIntosh, 1922
Fauveliopsis glabra (Hartman, 1960)(Fig. 40A H)

124

Minoru Imajima

Brada glabra Hartman, 1960: 129 130, pl. 14, figs. 1 2.
Fauveliopsis glabra: Hartman, 1969: 283 284, figs. 1 2; Hartman and Fauchald, 1971: 116 117; Levenstein, 1972: 173, tab.
2, fig. 1; Blake and Petersen, 2000: 35 38, figs. 3.1 3.3.

Material. NSMT-Pol. 110882 (5), 110888 (2), 110889 (1), WA06-H480, NSMT-Pol. 110758,
WA06-H1500D (5); NSMT-Pol. 110883, WA07-C1500D (3), NSMT-Pol. 110759, WA07-D900(1). NSMT-Pol. 110760, SO07-C7-B (99), NSMT-Pol. 110761, SO07-K2 (6), NSMT-Pol. 110762,

Fig. 40. Fauveliopsis glabra (Hartman). —A, entire animal in lateral view, × 15; B, anterior body, dorsal view, ×
30; C, left parapodium of setiger 1, posterior view, × 136; D, right parapodium of setiger 2, same view, × 136;
E, left parapodium of setiger 3, same view, × 136; F, left parapodium of setiger 4, same view, × 136; G, leftparapodium of setiger 10, same view, × 136; H, left parapodium of setiger 30, same view, × 136.

Deep-sea Benthic Polychaetes off Northern Honshu

125SO07-O1 (1). NSMT-Pol. 110886, KT07-29-H-1 (1), NSMT-Pol. 110763, KT07-29-M-1 (1), NSMT-Pol. 110764, KT07-29-M-3-2 (3), NSMT-Pol. 110887, KT07-29- M-3-3 (2).
Description. Body more or less club-shaped, rounded truncate anteriorly, somewhat thicker
and more cylindrical over long middle region, narrow in curved far posterior region, with segmental furrows poorly defined (Fig. 40A); 7 8 mm long, about 1 mm wide at middle region and with
34 35 setigers. Body surface with transverse ridges interrupted at more or less regular intervals by
numerous minute, narrow papillae. Color in spirit light tan.
Prostomium usually invaginated, causing some contraction of setiger 1. Anterior setal region
comprising setigers 1 3, with modified setae (Fig. 40B); notopodia of setiger 1 each with 2 weakly
falcate, slender, subequal spines; neuropodia of setiger 1 usually with 2 falcate spines; upper spine
thick, broad, and strongly hooked; lower one slender, about one-third as wide as upper one (Fig.
40C). Noto- and neuropodia of setigers 2 3 each with 2 subequal falcate spines similar to those ofnotopodia of setiger 1 (Fig. 40D E). Middle setal region beginning on setiger 4; all parapodia withsingle acicular spine and capillary seta in each ramus, with spines initially being more slender thanthose of setigers 1 3 (Fig. 40F), then becoming longer, thicker and more curved than those of setiger 4 (Fig. 40G). Posterior setal region characterized by segments with multiple neurosetae, 3 acicular spines and 2 capillaries (Fig. 40H).
Interramal papilla stalked, close to notosetae on setigers 1 4, and shifting to more medial
position between rami (Fig. 40G), then becoming closer to notosetae (Fig. 40H). Genital papillae
visible on posterior part of setiger 10, immediately anterodorsal to anteriorly oriented setae of rightparapodium 11 (Fig. 40B). Pygidium circular flattened dorsal lobe bending ventrally, forming
cover over anus.
The species is newly added to the Japanese polychaetous fauna.
Distribution. California; Japan.
Genus Laubieriopsis Petersen, 2000
Laubieriopsis brevis japonica subsp. nov.
(Fig. 41A J)
Type material. Holotype—NSMT-Pol. H 513, Stn. SO07-K3, off Kinkazan, Ishinomaki,
38˚33.7´N, 143˚41.2´E 38˚33.2´N, 143˚41.5´E, 4105 4181 m, Aug. 7, 2007. Paratypes—NSMTPol. P 514, Stn. SO07-K3 (32), same locality as holotype; NSMT-Pol. P 515, Stn. WA05-G1500D,
off Kashima Sea, 1498 m, Nov. 10, 2005 (1); NSMT-Pol. P 516, Stn. SO07-K1, off Kinkazan,
Ishinomaki, 2043 2081 m, Aug. 6, 2007 (42); NSMT-Pol. P 517, Stn. SO07-O1, off Onahama,2024 2020 m, Aug. 8, 2007 (1); NSMT-Pol. P 518, Stn. SO07-O3, off Onahama, 4128 4094 m,
Aug. 8, 2007 (1); NSMT-Pol. P 519, Stn. KT07-29-H-2, off Hachinohe, 2055 2032 m, Nov. 8,
2007 (37); NSMT-Pol. P 520, Stn. KT07-29-M-3-2, off Miyako, 1737 1709 m, Nov. 6, 2007 (6);
NSMT-Pol. P 521, Stn. KT07-29-M-3-3, off Miyako, 1733 1695 m, Nov. 9, 2007 (8).
Description. Holotype complete, linear, of similar width throughout, 5 mm long, about 0.33
mm wide at median region; length of paratypes 4 10 mm long; all specimens including holotype
and paratypes consisting of 16 segments. Body surface relatively smooth but covered with conicalmicropapillae throughout of surface under higher magnification. Anterior body with segmental
annulations and furrows poorly defined, subsequent segments with no evidence of segmental furrows (Fig. 41A).
Prostomium with rounded lobe apparent on anterior margin, most specimens retracted prostomium, with pair of nuchal organs. Peristomium distinct, about half as long as setiger 1 (Fig.
41B).

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Minoru Imajima

Fig. 41. Laubieriopsis brevis japonica subsp. nov. —A, entire animal in dorsal view, × 34; B, anterior end, dorsal
view, × 60; C, left parapodium from setiger 1, anterior view, × 287; D, cross section of second setiger, posterior view, × 137; E, right parapodium from setiger 2, anterior view, × 287; F, right parpodium from setiger 9,
same view, × 287; G, left parapodium from setiger 15, same view, × 287; H, setigers 6 8 showing genital papilla, lateral view, × 60; I, posterior end, dorsal view, × 60; J, pygidium, ventral view, × 84.

Deep-sea Benthic Polychaetes off Northern Honshu

127

First four setigers wider than long, with modified aciculars in laterally directed fascicles: eachnoto- and neuropodium with 2 thick weakly falcate, slender, subequal spines and 2 3 narrower
spines (Fig. 41C E). Parapodia of middle and posterior setal regions with gently curving, single
acicular spine and one capillary seta in each ramus (Fig. 41F G). Interramal papilla short-stalked
or sessile, situated midway between rami in anterior parapodia (Fig. 41C E), thereafter becoming
closer to notosetae (Fig. 41F G).
Genital papillae bearing on both sides at posterior edge of setiger 6, just anterior to notochaetae of setiger 7 (Fig. 41H).
Last segment well developed, with 4 large, long acicular spines slightly flexed distally, anddirected nearly straight back (Fig. 41I). Boundary between distal part of last segment and pygidium indistinct. Anus terminal, usually withdrawn within rounded pygidium, with 2 small papillae
at middorsal margin (Fig. 41J).
Remarks. The subspecies can be distinguished from the stem, Laubieriopsis brevis (Hartman,1965) from Antarctic Sea, in having the rounded pygidium instead of having a broad posterior slit
between the bases of the acicular spines.
Etymology. The subspecies is named because it is the first species of the genus Laubieriopsis
from Japanese waters.
Distribution. Japan (1498 4181 m depth).
Order Sternaspida
Family Sternaspidae Carus, 1863
Genus Sternaspis Otto, 1821
Sternaspis scutata (Ranzani, 1807)
Thalassema scutatum Ranzani, 1817: 1461.
Sternaspis scutata: Moore, 1903: 487; Okuda, 1936: 151 152, textfig. 5; Imajima and Hartman, 1964: 310 311; Imajima,
1997a: 199; Petersen, 2000: 328 329, fig. 11. 3. B C.

Material. NSMT-Pol. 109842, WA05-DE250D (42), NSMT-Pol. 109843, WA05-DE380D(264), NSMT-Pol. 109844, WA05-E1000D (1), NSMT-Pol. 109845, WA05-EF250 (1), NSMTPol. 109846, WA05-EF250D (12), NSMT-Pol. 109847, WA05-FG250D (66), NSMT-Pol. 109848,
WA05-FG480 (2), NSMT-Pol. 109849, WA05-FG510D (3), NSMT-Pol. 109850, WA05-G1500D(3), NSMT-Pol. 109851, WA05-GH380 (3), NSMT-Pol. 109852, WA05-GH380D (3), NSMT-Pol.
109853, WA05-GH510D (1); NSMT-Pol. 110736, WA06-A410 (1), NSMT-Pol. 110737, WA06C550 (1), NSMT-Pol. 109854, WA06-D210D (7), NSMT-Pol. 109855, WA06-D450D (15), NSMT-Pol. 109856, WA06-DE280D (85), NSMT-Pol. 109857, WA06-E510 (1), NSMT-Pol. 109858,
WA06-E510D (22), NSMT-Pol. 110738, WA06-E1200D (2), NSMT-Pol. 109859, WA06-EF425D(1), NSMT-Pol. 109860, WA06-F1500D II (6), NSMT-Pol. 109861, WA06-FG350D (2), NSMTPol. 110739, WA06-G510 (2), NSMT-Pol. 109862, WA06-G900D (3), NSMT-Pol. 109863, WA06G1200D (1), NSMT-Pol. 109864, WA06-GH380 (1), NSMT-Pol. 109865, WA06-GH450 (1),
NSMT-Pol. 109866, WA06-GH480D (5), NSMT-Pol. 109867, WA06-H250D (68), NSMT-Pol.
110450, WA06-H480 (6), NSMT-Pol. 109868, WA06-H1500D (4); NSMT-Pol. 110558, WA07A450 (4), NSMT-Pol. 110731, WA07-A650 (5), NSMT-Pol. 110732, WA07-B410D (116), NSMTPol. 110733, WA07-C1500D (2), NSMT-Pol. 110734, WA07-D210D (135), NSMT-Pol. 110735,
WA07-D900 (2).
Distribution. Arctic, Atlantic, Pacific and Indian oceans; Japan.

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Order Capitellida
Family Capitellidae Grube, 1862
Genus Capitella Blainville, 1828
Capitella capitata (Fabricius, 1780)
Lumbricus capitatus Fabricius, 1780: 279.
Capitella capitata: Fauvel, 1927: 154, fig. 55a h; Hartman, 1947b: 404, pl. 43, figs. 1 2; Imajima and Hartman, 1964:
311 312.

Material. NSMT-Pol. 110657, WA05-FG280 (1), NSMT-Pol. 110658, WA05-G425 (1); NSMT-Pol. 110659, WA07-C510 (1).
Distribution. North Atlantic; Southern California; Mediterranean Sea; Bering Sea; Japan.
Genus Leiochrides Augener, 1914
Leiochrides sp.
Material. NSMT-Pol. 110661, WA05-G1500D (2).
Genus Notomastus Sars, 1850
Notomastus hemipodus Hartman, 1945
Notomastus (Clistomastus) hemipodus Hartman, 1945: 38.
Notomastus hemipodus: Day, 1973: 100; Blake, 2000b: 81 83, fig. 4.13; Imajima, 2006: 383.

Material. NSMT-Pol. 110921, WA05-G1500D (2).
Distribution. North Carolina; Gulf of Mexico; California; Japan.
Notomastus latericeus Sars, 1851
Notomastus latericeus Sars, 1851: 199; Fauvel, 1927: 143, fig. 49a h; Uschakov, 1955: 325, fig. 121A B; Imajima and Hartman, 1964: 313; Imajima, 1997a: 199 200.

Material. NSMT-Pol. 109896, WA05-DE380D (1), NSMT-Pol. 109895, WA05-E1000D (8),NSMT-Pol. 110542, WA05-FG250D (1), NSMT-Pol. 109897, WA05-FG510D (1), NSMT-Pol.
109898, WA05-G1500D (1), NSMT-Pol. 109899, WA05-H900D (7); NSMT-Pol. 109900, WA06E510D (1), NSMT-Pol. 109901, WA06-E1200D (3), NSMT-Pol. 109904, WA06-F650D (1), NSMT-Pol. 109902, WA06-G900D (1), NSMT-Pol. 109903, WA06-GH480D (10); NSMT-Pol.
110543, WA07-A1500D (1), NSMT-Pol. 110545, WA07-B410D (1), NSMT-Pol. 110544, WA07B1500D (6), NSMT-Pol. 110546, WA07-C1500D (2), NSMT-Pol. 110547, WA07-D900 (1). NSMT-Pol. 110862, KT07-29-H-2 (1).
Distribution. Western and southern Europe; Sea of Okhotsk; Japan.
Genus Pseudoleiocapitella Harmelin, 1964
Pseudoleiocapitella sp.
Material. NSMT-Pol. 110662, WA06-B750D (1), NSMT-Pol. 110663, WA06-F650D (1),
NSMT-Pol. 110660, WA07-C1500D (1).
The genus is reported as a part of the Japanese polychaetous fauna for the first time.

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Family Maldanidae Malmgren, 1867
Subfamily Clymenurinae Imajima and Shiraki, 1982
Genus Clymenura Verrill, 1900
Clymenura (Cephalata) aciculata Imajima and Shiraki, 1982
Clymenura (Cephalata) aciculata Imajima and Shiraki, 1982a: 21 22, fig. 6a n.

Material. NSMT-Pol. 109927, WA05-DE250D (1), NSMT-Pol. 109928, WA05-DE380D (8),NSMT-Pol. 109929, WA05-EF450D (1).
Distribution. Japan.
Clymenura (Cephalata) columbiana (Berkeley, 1929)
Leiochone columbiana Berkeley, 1929: 315 316, pl. 1, figs. 1 9.
Clymenura (Cephalata) columbiana: Imajima and Shiraki, 1982a: 23 24, fig. 7a 1; Imajima, 1997a: 201.

Material. NSMT-Pol. 109930, WA05-DE380D (1), NSMT-Pol. 109931, WA05-EF250D (1),
NSMT-Pol. 109932, WA05-GH510D (2).
Distribution. Pacific of western Canada; Japan.
Clymenura (Cephalata) lankesteri (McIntosh, 1885)
Praxilla lankesteri McIntosh, 1885: 403 404, pl. 25A, fig. 3.
Leiochone borealis Arwidsson, 1907: 156 163, pl. 3, figs. 108 115, pl. 4, figs. 116 117, pl. 9, figs. 281 283, pl. 11, figs.
352 353.
Clymenura borealis: Hartman, 1959: 456.
Clymenura (Cephalata) lankesteri: Imajima and Shiraki, 1982a: 16 19, figs. 3a n, 4a d; Imajima, 1997a: 200 201.

Material. NSMT-Pol. 109933, WA05-GH510D (8); NSMT-Pol. 109934, WA06-EF425D (3),
NSMT-Pol. 109935, WA06-F650D (2), NSMT-Pol. 109936, WA06-GH480D (14); NSMT-Pol.
110166, WA07-B1500D (1).
Distribution. Sea of Okhotsk; west coast of Norway; Japan.
Clymenura (Cephalata) longicaudata Imajima and Shiraki, 1982
Clymenura (Clymenura) longicaudata Imajima and Shiraki, 1982a: 19 21, fig. 5a p.

Material. NSMT-Pol. 110167, WA07-B1500D (1).
Distribution. Japan.
Clymenura (Cephalata) sp.
Material. NSMT-Pol. 110818, SO07-O2 (11).
Clymenura (Clymenura) japonica Imajima and Shiraki, 1982
Clymenura (Clymenura) japonica Imajima and Shiraki, 1982a: 12 15, fig. 2a m; Imajima, 1997a: 200.

Material. NSMT-Pol. 109937, WA05-DE250D (4), NSMT-Pol. 109938, WA05-DE380D (1),
NSMT-Pol. 109939, WA05-E1000D (1), NSMT-Pol. 109940, WA05-EF250D (2), NSMT-Pol.

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109941, WA05-FG250D (7), NSMT-Pol. 109942, WA05-GH510D (1); NSMT-Pol. 109943,
WA06-G900D (4).
Distribution. Japan.
Subfamily Euclymeninae Arwidsson, 1907
Genus Clymenella Verrill, 1873
Clymenella koellikeri (McIntosh, 1885)
Praxilla koellikeri McIntosh, 1885: 402 403, pl. 46, fig. 6, pl. 25A, fig. 2, pl. 37A, figs. 3, 8.
Clymenella koellikeri: Imajima and Shiraki, 1982b: 52 54, figs. 23a h, 24a b; Imajima, 1997a: 203.

Material. NSMT-Pol. 109912, WA05-GH510D (1).
Distribution. Fiji Island; Japan.
Genus Euclymene Verrill, 1900
Euclymene uncinata Imajima and Shiraki, 1982
Euclymene uncinata Imajima and Shiraki, 1982b: 70 71, fig. 33a l.

Material. NSMT-Pol. 109944, WA05-E1000D (1); NSMT-Pol. 109945, WA06-GH480D (1).
Distribution. Japan.
Genus Isocirrus Arwidsson, 1907
Isocirrus planiceps (Sars, 1872)
Isocirrus planiceps: Arwidsson, 1907: 137 143, pl. 3, figs. 98 107, pl. 8, figs. 276 280, pl. 11, figs. 348, 351; Imajima andShiraki, 1982b: 73 74, fig. 35a-j.

Material. NSMT-Pol. 110341, WA06-A150D (1); NSMT-Pol. 110168, WA07-A650 (1).
Distribution. Norway; Japan.
Genus Maldanella McIntosh, 1885
Maldanella harai (Izuka, 1902)
Clymene harai Izuka, 1902: 111 113, pl. 3, figs. 9 12.
Maldanella harai: Fauvel, 1914: 260 261, pl. 23, fig. 1; Fauvel, 1927: 186, fig. 64i n; Fauvel, 1953: 383 384, fig. 199i n;
Uschakov, 1955: 342, fig. 126E G; Imajima and Hartman, 1964: 319 320; Imajima and Shiraki, 1982b: 55 56, fig.
25a h.
Axiothea campanulata Moore, 1903: 485 487, pl. 27, figs. 97 99.
Maldanella robusta Moore, 1906: 236 239, pl. 11, figs. 31 32.

Material. NSMT-Pol. 109962, WA05-DE380D (12), NSMT-Pol. 109963, WA05- EF250D(2), NSMT-Pol. 109964, WA05-H900D (1); NSMT-Pol. 109965, WA06-D450D (3), NSMT-Pol.
109966, WA06-EF425D (4), NSMT-Pol. 109967, WA06-FG350D (1), NSMT-Pol. 109968, WA06GH480D (4).
Distribution. Atlantic and Indian oceans; Sea of Okhotsk; Japan.
Maldanella sp.
Material. NSMT-Pol. 110802, SO07-O2 (2).

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Genus Microclymene Arwidsson, 1907
Microclymene caudata Imajima and Shiraki, 1982
Microclymene caudata Imajima and Shiraki, 1982b: 65 67, fig. 30a o; Imajima, 1997a: 204.

Material. NSMT-Pol. 109973, WA05-DE380D (2), NSMT-Pol. 109974, WA05-EF250D (1);
NSMT-Pol. 110342, WA06-EF425D (2), NSMT-Pol. 110343, WA06-H250D (4).
Distribution. Japan.
Genus Praxillella Verrill, 1881
Praxillella gracilis (Sars, 1861)
Praxillella gracilis: Arwidsson, 1907: 183 191, pl. 4, figs. 153 155, pl. 5, figs. 156 158, pl. 9, figs. 302 307, pl. 12, fig. 367;
Fauvel, 1927: 178 179, fig. 62m p; Mesnil and Fauvel, 1939: 4 5, fig. 1; Berkeley and Berkeley, 1952: 50, figs.
101 102; Hartman, 1969: 477 478; Imajima and Shiraki, 1982b: 61 63, fig. 28a k; Imajima, 1997a: 203 204.

Material. NSMT-Pol. 109984, WA05-DE380D (4), NSMT-Pol. 109985, WA05-EF250D (8),NSMT-Pol. 109986, WA05-G280 (2), NSMT-Pol. 109987, WA05-GH510D (2), NSMT-Pol.
109988, WA05-H900D (1); NSMT-Pol. 109989, WA06-B750D (2), NSMT-Pol. 109990, WA06EF425D (1), NSMT-Pol. 109991, WA06-G900D (1), NSMT-Pol. 109992, WA06-G1200D (3),
NSMT-Pol. 109993, WA06-GH480D (1), NSMT-Pol. 109994, WA06-H250D (2), NSMT-Pol.
109995, WA06-H480 (1); NSMT-Pol. 110176, WA07-B1500D (4), NSMT-Pol. 110177, WA07D210D (1).
Distribution. Southern California north to western Canada; north Atlantic and western Europe; Mediterranean Sea; Japan.
Praxillella pacifica Berkeley, 1929
Praxillella affinis var. pacifica Berkeley, 1929: 313 314; Berkeley and Berkeley, 1952: 49 50, figs. 97 100; Hartman, 1969:475 476.
Praxillella pacifica: Imajima and Shiraki, 1982b: 58 60, fig. 27a l.

Material. NSMT-Pol. 109996, WA05-EF250D (5), NSMT-Pol. 109997, WA05-GH380D (1);
NSMT-Pol. 109998, WA06-EF425D (1), NSMT-Pol. 109999, WA06-G900D (1), NSMT-Pol.
110000, WA06-G1200D (1), NSMT-Pol. 110001, WA06-GH480D (17), NSMT-Pol. 110002,
WA06-H250D (2). NSMT-Pol. 110633, KT07-29-H-2 (1).
Distribution. Southern California north to western Canada; Japan.
Praxillella praetermissa (Malmgren, 1865)
Praxilla praetermissa Malmgren, 1865: 191.
Praxillella praetermissa: Arwidsson, 1907: 192 204, pl. 4, figs. 136a 143, pl. 9, figs. 294 296, pl. 12, figs. 361 363; Day,
1967b: 642 644, fig. 30.7.i l; Imajima and Shiraki, 1982b: 63 65, fig. 29a n; Imajima, 1997a: 204.

Material. NSMT-Pol. 110003, WA05-DE380D (5), NSMT-Pol. 110004, WA05-GH510D(10); NSMT-Pol. 110005, WA06-F750 (1), NSMT-Pol. 110006, WA06-GH480D (46), NSMT-Pol.
110007, WA06-H1500D (1); NSMT-Pol. 110178, WA07-C1500D (1), NSMT-Pol. 110179, WA07D210D (1). NSMT-Pol. 110634, KT07-29-M-3-1 (1).
Distribution. Arctic; North Atlantic from Norway to Spain; Mediterranean Sea; Japan.

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Subfamily Lumbriclymeninae Arwidsson, 1907
Genus Clymenopsis Verrill, 1900
Clymenopsis cingulata (Ehlers, 1887)
Clymene cingulata Ehlers, 1887: 185 188, pl. 47, figs. 2 5.
Clymenopsis cingulata: Hartman, 1960: 144 145; Imajima and Shiraki, 1982a: 30 32, fig. 12a k; Imajima, 1997a: 201 202.

Material. NSMT-Pol. 109913, WA05-DE380D (1), NSMT-Pol. 109914, WA05-E1000D (1),
NSMT-Pol. 109915, WA05-EF250D (4), NSMT-Pol. 109916, WA05-G1500D (1); NSMT-Pol.
109917, WA06-E510D (2), NSMT-Pol. 109918, WA06-E1200D (1), NSMT-Pol. 109919, WA06EF425D (3), NSMT-Pol. 109920, WA06-F1500D (1), NSMT-Pol. 109921, WA06-F1500D II (3),
NSMT-Pol. 109922, WA06-G900D (3), NSMT-Pol. 109923, WA06-G1200D (6), NSMT-Pol.
109924, WA06-GH480D (17), NSMT-Pol. 109925, WA06-H250D (10), NSMT-Pol. 109926,
WA06-H1500D (5).
Distribution. Southern California; Greenland; Japan.
Genus Lumbriclymene Sars, 1872
Lumbriclymene japonica (McIntosh, 1885)
Nicomache japonica McIntosh, 1885: 399 400, pl. 46, fig. 5, pl. 24A, fig. 20.
Lumbriclymene japonica: Imajima and Shiraki, 1982a: 26 28, fig. 9a r, fig. 10a d; Imajima, 1997a: 201.

Material. NSMT-Pol. 109946, WA05-EF250D (1), NSMT-Pol. 109947, WA05-GH510D (3);
NSMT-Pol. 109948, WA06-G900D (1). NSMT-Pol. 110819, SO07-O2 (3).
Distribution. Japan.
Genus Notoproctus Arwidsson, 1907
Notoproctus pacificus (Moore, 1906)
Lumbriclymene pacifica Moore, 1906: 246 248, pl. 12, figs. 40 42.
Notoproctus pacificus: Berkeley and Berkeley, 1952: 56 67, figs. 117 118; Hartman, 1948a: 8; Hartman, 1969: 469; Imajima,
1964: 249 251, figs. 42 50; Imajima and Shiraki, 1982a: 24 26, fig. 8a m; Imajima, 2001a: 83.

Material. NSMT-Pol. 109979, WA05-DE380D (1), NSMT-Pol. 109980, WA05-EF250D (1),
NSMT-Pol. 109981, WA05-GH380D (7), NSMT-Pol. 109982, WA05-GH510D (10); NSMT-Pol.
109983, WA06-EF425D (1); NSMT-Pol. 110174, WA07-B410D (4), NSMT-Pol. 110175, WA07C1500D (1).
Distribution. Southern California; western Canada and Alaska; Japan.
Genus Praxillura Verrill, 1880
Praxillura tanseiana Imajima and Shiraki, 1982
Praxillura tanseiana Imajima and Shiraki, 1982a: 29 30, fig. 11a n.

Material. NSMT-Pol. 110008, WA05-EF250D (1), NSMT-Pol. 110009, WA05-GH380D (3);
NSMT-Pol. 110010, WA06-G900D (2), NSMT-Pol. 110011, WA06-H250D (1).
Distribution. Japan.

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Subfamily Maldaninae Arwidsson, 1907
Genus Asychis Kinberg, 1867
Asychis auritus Uschakov, 1950
(Fig. 42A I)
Asychis auritus Uschakov, 1950: 214; Uschakov, 1965: 321 322, fig. 127 H I; Light, 1991: 140.

Material. NSMT-Pol. 110335, WA05-E1000D (3), NSMT-Pol. 110336, WA05-GH510D (2);
NSMT-Pol. 110337, WA06-B750D (23), NSMT-Pol. 110338, WA06-C900D (1), NSMT-Pol.
110339, WA06-F650D (2).
Description. Largest complete specimen 200 mm long, 6 mm wide for 19 setigerous segments, an apodous preanal segment and pygidium. First seven setigers distinctively biannulate.
First setiger with low collar ventrally (Fig. 42A).
Cephalic plate elliptical; prostomium broad, rounded with smooth margin. Cephalic rim divided into three lobes by deep lateral notches; lateral rim auricular, wide, more elevated withsmooth margin. Posterior rim crenulated by small serrations. Cephalic keel broad and short. Nuchalorgans deeply curved; inner shanks nearly parallel to each other, less than half length of cephalicplate; outer shanks turn outward posteriorly, about half as long as inner shanks (Fig. 42 B).
First setigerous segment uniramous, with only notopodial fascicle consisting of short andlong limbate capillaries arranged in two rows (Fig. 42C D). Median notopodial fascicle with long,
distally spinous setae (Fig. 42E). Neuropodial uncini present from second setiger with six uncini
in linear series; median neuropodia with about 30 uncini. Uncini typical rostrate, with teeth intransverse arc above main fang and gular bristles (Fig. 42F G).
Anal plate forming deep, funnel-like pocket; posterior wall of dorsal lobe expanded, with twoacute angles near lateral incisions produced into cirri; and median elongated cirrus; ventral lobe
with undulating margin (Fig. 42H I). Anal pore present dorsally on plate (Fig. 42H).
The species is newly added to the Japanese polychaetous fauna.
Distribution. West coast of south Sakhalin; Kamchatka; Sea of Okhotsk; Japan.
Genus Chirimia Light, 1991
Chirimia biceps (Sars, 1861)
Asychis biceps: Arwidsson, 1907: 263 271, pl. 6, figs. 200 207, pl. 10, figs. 339 344; Imajima and Shiraki, 1982b: 77 80, fig.
37a t.
Asychis lacera Moore, 1923: 235 237.
Asychis lobata Fauchald, 1972b: 256 258, pl. 52, figs. a f.
Chirimia biceps biceps: Light, 1991: 139; Imajima, 1997a: 205.

Material. NSMT-Pol. 109906, WA05-GH380D (1), NSMT-Pol. 109907, WA05-GH510D(2), NSMT-Pol. 109908, WA05-H900D (2); NSMT-Pol. 109909, WA06-F650D (1), NSMT-Pol.
109910, WA06-G900D (3), NSMT-Pol. 109911, WA06-GH480D (7).
Distribution. Iceland; Greenland; California; Scotland; Atlantic coast of Europe; western
Mexico; Japan.
Genus Maldane Grube, 1860
Maldane cristata Treadwell, 1923
Maldane cristata Treadwell, 1923: 9 10, figs. 5 8; Hartman, 1956: 295 296; Imajima and Shiraki, 1982b: 84 86, fig. 40a n;
Imajima, 1997a: 205 206; Imajima, 2006: 386.

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Fig. 42. Asychis auritus Uschakov. —A, anterior end, lateral view, × 7; B, cephalic plate, frontal view, × 7; C; partof notosetal fascicle, × 50; D, short limbate capillary notoseta, × 238; E, spinous capillary notoseta, × 250,
with detail of part, × 642; F, rostrate uncinus, × 118; G, distal end of same, × 426; H, posterior end, lateralview, × 6; I, anal plate, posterior view, × 6.

Maldane sarsi: McIntosh, 1885: 392 393; Okuda, 1939: 239 240; Imajima and Hartman, 1964: 317 318.

Material. NSMT-Pol. 109949, WA05-DE380D (2), NSMT-Pol. 109950, WA05-EF250D (4),
NSMT-Pol. 109951, WA05-GH510D (5), NSMT-Pol. 109952, WA05-H900D (7); NSMT-Pol.
109953, WA06-E510D (14), NSMT-Pol. 109954, WA06-F650D (6), NSMT-Pol. 109955, WA06-

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G900D (75), NSMT-Pol. 109956, WA06-G1200D (1), NSMT-Pol. 109957, WA06-GH480D (22),
NSMT-Pol. 109958, WA06-H250D (58), NSMT-Pol. 109959, WA06-H1500D (1); NSMT-Pol.
110169, WA07-D210D (10), NSMT-Pol. 110170, WA07-D900 (6). NSMT-Pol. 110633, KT0729-H-2 (1).
Distribution. Southern California to western Mexico; Japan.
Maldane pigmentata (Imajima and Shiraki, 1982)
Asychis pigmentata Imajima and Shiraki, 1982b: 82 83, fig. 39a k.
Maldane pigmentata: Imajima, 1996: 288, fig. 233; Imajima, 1997a: 205; Imajima, 2001a: 85 86; Imajima, 2006: 387.

Material. NSMT-Pol. 109960, WA05-EF250D (1); NSMT-Pol. 109961, WA06-H250D (2).
Distribution. Japan.
Maldane sp.
Material. NSMT-Pol. 110820, SO07-C7-B (3).
Genus Metasychis Light, 1991
Metasychis disparidentata (Moore, 1904)
Maldane disparidentata Moore, 1904: 494 496, pl. 38, figs. 28 31.
Maldane (Asychis) disparidentata: Takahashi, 1938: 209 211, textfig. 12.
Asychis disparidentata: Berkeley and Berkeley, 1952: 46 47, figs. 89 90; Imajima and Hartman, 1964: 316 317; Hartman,
1969: 423; Imajima and Shiraki, 1982b: 80 82, fig. 38a k.
Metasychis disparidentata: Light, 1991: 141.

Material. NSMT-Pol. 109969, WA06-A900 (2), NSMT-Pol. 109970, WA06-F650D (2).
Distribution. Western Canada south of southern California; Japan.
Metasychis gotoi (Izuka, 1902)
Maldane gotoi Izuka, 1902: 109 111, pl. 3, figs. 1 8.
Maldane coronata Moore, 1903: 483 485, pl. 27, figs. 94 96.
Asychis gotoi: Fauvel, 1932b: 205; Fauvel, 1953: 387, fig. 200a b; Okuda, 1938a: 100; Okuda, 1939: 239; Mesnil and Fauvel, 1939: 16 17, fig. 11; Uschakov, 1955: 345, fig. 127A D; Imajima and Hartman, 1964: 317; Imajima, 1972: 13;
Imajima and Shiraki, 1982b: 75 77, fig. 36a l.
Asychis shaccotanus Uchida, 1968: 603 604.
Metasychis gotoi: Light, 1991: 139; Imajima, 1997a: 204 205; Imajima, 2001a: 86.

Material. NSMT-Pol. 109971, WA06-E550 (1), NSMT-Pol. 109972, WA06-H250D (2).
Distribution. Indo-Pacific areas; Adriatic Sea; California; Japan.
Subfamily Nicomachinae Arwidsson, 1907
Genus Nicomache Malmgren, 1865
Nicomache (Nicomache) lumbricalis (Fabricius, 1780)
Nicomache lumbricalis: Arwidsson, 1907: 86 93, pl. 8, figs. 244 245; Fauvel, 1927: 190 191, fig. 66a i; Wesenberg-Lund,
1948: 23 27, figs. 10, 11d f; Wesenberg-Lund, 1950: 41; Hartman, 1948a: 42; Hartman, 1969: 465; Berkeley andBerkeley, 1952: 54 55, figs. 111 112; Pettibone, 1954: 305 306, fig. 34i j; Uschakov, 1955: 336, fig. 124A D; Imajima,
1964: 248 249, figs. 39 41; Day, 1967b: 621, fig. 30.1.i o; Hartmann-Schröder, 1971: 428 430, fig. 149a f; Imajima

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and Shiraki, 1982a: 35 37, fig. 14a n; Imajima, 1997a: 202.
Nicomache (Nicomache) lumbricalis: Imajima, 2001a: 86.
Nicomache benthaliana McIntosh, 1885: 400, pl. 46, fig. 8, pl. 24A, fig. 21.

Material. NSMT-Pol. 109975, WA05-E1000D (1), NSMT-Pol. 109976, WA05-GH510D (3);
NSMT-Pol. 109977, WA06-D210D (1), NSMT-Pol. 109978, WA06-D1500D (1); NSMT-Pol.
110171, WA07-A510 (1), NSMT-Pol. 110172, WA07-B1500D (1), NSMT-Pol. 110173, WA07D1500D (1).
Distribution. Greenland; North Sea; Kara Sea, Bering Sea; Pacific coast of North America;
Japan.
Nicomache (Nicomache) sp.
Material. NSMT-Pol. 110821, SO06-M1-B (1), NSMT-Pol. 110822, SO06-M3-B2 (1), NSMT-Pol. 110823, SO06-M4-B (1); NSMT-Pol. 110824, SO07-O2 (1).
Subfamily Rhodininae Arwidsson, 1907
Genus Rhodine Malmgren, 1865
Rhodine loveni Malmgren, 1865
Rhodine loveni: Arwidsson, 1907: 64 74, pl. 2, figs. 39 52, pl. 7, figs. 235 236, pl. 11, figs. 346 347; Hartman, 1966: 72, pl.
23, figs. 9 11; Imajima and Shiraki, 1982a: 32 35, fig. 13a m; Imajima, 1997a: 202.

Material. NSMT-Pol. 110012, WA05-DE380D (8), NSMT-Pol. 110013, WA05-EF250D (4),
NSMT-Pol. 110014, WA05-FG510D (1), NSMT-Pol. 110015, WA05-GH380D (1), NSMT-Pol.
110016, WA05-GH510D (3); NSMT-Pol. 110017, WA06-E510D (1), NSMT-Pol. 110018, WA06EF425D (4), NSMT-Pol. 110019, WA06-FG350D (1), NSMT-Pol. 110020, WA06-GH480D (16);
NSMT-Pol. 110180, WA07-A650 (1), NSMT-Pol. 110181, WA07-A1500D (20), NSMT-Pol.
110182, WA07-B1500D (1), NSMT-Pol. 110183, WA07-D210D (16), NSMT-Pol. 110184, WA07D1500D (1). NSMT-Pol. 110825, SO07-C7-B (1).
Distribution. Arctic boreal; Japan.
Order Opheliida
Family Opheliidae Malmgren, 1867
Genus Ophelina Oersted, 1843
Ophelina acuminata Oersted, 1843
Ophelina acuminata: Hobson and Banse, 1981: 62, fig. 13f; Hartmann-Schröder, 1996: 427, fig. 209; Blake, 2000c: 158 159,
fig. 7.5.
Ammotrypane aulogaster: Berkeley and Berkeley, 1952: 92, figs. 186 187; Okuda, 1936: 149 150, textfigs. 2 3; Imajima andHartman, 1964: 305 306; Imajima, 1997a: 206.

Material. NSMT-Pol. 109885, WA05-DE250D (8), NSMT-Pol. 110773, WA05-DE380D (2),
NSMT-Pol. 109886, WA05-FG250D (43), NSMT-Pol. 109887, WA05-FG510D (11), NSMT-Pol.
109888, WA05-G1500D (2), NSMT-Pol. 110774, WA05-GH510D (6); NSMT-Pol. 109889,
WA06-D210D (1), NSMT-Pol. 109890, WA06-E510D (3), NSMT-Pol. 109891, WA06-E1200D(6), NSMT-Pol. 109892, WA06-F1500D II (1), NSMT-Pol. 109893, WA06-FG350D (2), NSMTPol. 109894, WA06-GH480D (9). NSMT-Pol. 110861, KT07-29-H-1 (7).
Distribution. Atlantic, Arctic, Pacific and Indian oceans; Japan.

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137

Ophelina arctica (McIntosh, 1879) comb. nov.
(Fig. 43A D)
Ammotrypanella arctica McIntosh, 1879: 505, pl. 65, fig. 12; Fauvel, 1914: 246 247, pl. 22, figs. 14 19.

Material. NSMT-Pol. 110775, WA05-FG250D (20); NSMT-Pol. 110776, WA06-DE280D(15), NSMT-Pol. 110777, WA06-GH480D (3). NSMT-Pol. 110859, KT07-29-H-1 (1), NSMTPol. 110860, KT07-29-H-2 (1), NSMT-Pol. 110858, KT07-29-M-1 (56).
Description. Body elongate, weakly annulated with deep midventral groove and 2 lateralgrooves throughout; largest specimen 12 mm long, 0.8 mm wide for 35 setigers; anal tube 0.6 mm
long. Branchiae from setiger 2, continuing to posterior end, except 4 abranchiate prepygidial segments.
Prostomium narrow, sharply pointed, with terminal palpode; prostomial and lateral eyes absent. Lateral nuchal organs conspicuous, forming a deep fissure (Fig. 43A). Parapodia biramous,
reduced; noto- and neuropodia with small fascicles of slender capillaries (Fig. 43B). Posterior end
terminating in distinct cylindrical anal tube, with terminal opening, and bearing 24 26 short subequal digitate papillae and one long, unpaired ventral cirrus at end (Fig. 43C D). Small specimen

Fig. 43. Ophelina arctica (McIntosh). —A, anterior end, lateral view, × 26; B, median parapodium, anterior view,
× 72; C, posterior end, lateral view, × 53; D, same, ventral view, × 53.

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Minoru Imajima

with anal tube bearing 8 short papillae at end.
The species is newly added to the Japanese polychaetous fauna.
Remarks. Ophelina arctica is similar to O. breviata (Ehlers, 1913) in having long cylindricalanal tube. However, the anal tube of O. arctica has more distinct conical terminal papillae rather
than a lobulate edge. Moreover, the both species are distinguished in the structure of the abranchiate prepygidial segments. The anal tubes of the examined specimens are provided with 24 26
digitate papillae rather than short papillae in the McIntosh’s specimen. The differentiation of thischaracter appears to be an environmental effect.
Distribution. Atlantic Ocean; Japan.
Ophelina breviata (Ehlers, 1913).
(Fig. 44A F)
Ammotrypane breviata Ehlers, 1913: 523 524, pl. 39, figs. 1 7; Pettibone, 1954: 295 296, fig. 33l n; Berkeley, 1966:
847 848.
Ophelina breviata: Hobson and Banse, 1981: 62, fig. 13g.

Material. NSMT-Pol. 110809, WA06-DE280D (1); NSMT-Pol. 110810, WA06-E510D (7);WA06-GH480D (6).
Description. Largest specimen 12.4 mm long, anal tube 0.6 mm long for total length of 13mm, about 0.5 mm wide across middle of body, with 28 setigers. Body subcylindrical, further backcompressed from sides, more or less rigid, with midventral groove present throughout; segmental
furrows and annulations poorly developed. Branchiae from setiger 2 (Fig. 44A), continuing toposterior end except 4 abranchial prepygidial segments (Fig. 44B D).
Prostomium conical, tapering to narrow tip bearing terminal palpode; prostomial and lateraleyes absent. Lateral nuchal organs conspicuous, forming a deep fissure faintly pigmented at bottom. Proboscis soft, with lobes, not fringed (Fig. 44A).
Parapodia biramous, reduced; noto- and neuropodia with small fascicles of slender capillaries(Fig. 44E F); notosetae longer than neurosetae. Posterior end terminating in distinct cylindricalanal tube, with smooth or wrinkles along its hinder margin except at its dorsal peak.
The species is newly added to the Japanese polychaetous fauna.
Distribution. Kaiser Wilhelm Land; East Greenland; Arctic Alaska; British Columbia; Japan.
Genus Travisia Johnston, 1840
Travisia brevis Moore, 1923
Travisia brevis Moore, 1923: 220 221; Berkeley and Berkeley, 1952: 90 91, fig. 183; Imajima, 1964: 247; Hobson and Banse,
1981: 62, fig. 13h; Blake, 2000c: 161 162, fig. 7.7.

Material. NSMT-Pol. 110516, WA05-DE380D (11), NSMT-Pol. 110517, WA05-E750 (2),
NSMT-Pol. 110518, WA05-EF250D (5), NSMT-Pol. 110519, WA05-EF450D (5), NSMT-Pol.
110520, WA05-F900 (7), NSMT-Pol. 110521, WA05-FG250D (8), NSMT-Pol. 110522, WA05FG510D (1), NSMT-Pol. 110523, WA05-G750 (1), NSMT-Pol. 110524, WA05-G900 (2), NSMTPol. 110525, WA05-G1500D (1), NSMT-Pol. 110526, WA05-H650 (2), NSMT-Pol. 110527,
WA05-H750 (1), NSMT-Pol. 110528, WA05-H900D (2); NSMT-Pol. 110529, WA06-B750D (5),NSMT-Pol. 110530, WA06-D450D (4), NSMT-Pol. 110666, WA06-DE280D (2), NSMT-Pol.
110531, WA06-E510D (1), NSMT-Pol. 110532, WA06-EF425D (22), NSMT-Pol. 110533, WA06F650D (3), NSMT-Pol. 110534, WA06-G900D (7), NSMT-Pol. 110535, WA06-GH480D (3);
NSMT-Pol. 110536, WA07-B1500D (2), NSMT-Pol. 110537, WA07-C350D (1), NSMT-Pol.

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139

Fig. 44. Ophelina breviata (Ehlers). —A, anterior end, dorsal view, × 32; B, part of body, showing setigers 11 14,
lateral view, × 32; C, posterior end, same view, × 32; D, same, ventral view, × 32; E, parapodium from setiger
16, anterior view, × 65; F, notoseta, × 266.

110538, WA07-C750 (1), NSMT-Pol. 110539, WA07-D750 (3), NSMT-Pol. 110540, WA07-D900(1).
Distribution. Arctic-boreal; circumpolar; Japan.
Travisia pupa Moore, 1906
(Fig. 45A E)
Travisia pupa Moore, 1906: 228, pl. 11, fig. 23; Berkeley and Berkeley, 1952: 89 90, figs. 181 182; Uschakov, 1965: 301, fig.
120F G; Hobson, 1976: 139; Hobson and Banse, 1981: 62, fig. 13j; Blake, 2000c: 163, fig. 7.8.

Material. NSMT-Pol. 110651, WA05-GH510D (2); NSMT-Pol. 110652, WA06-D750 (1),
NSMT-Pol. 110653, WA06-G650 (2), NSMT-Pol. 110654, WA06-H550 (1), NSMT-Pol. 110655,
WA06-H650 (3), NSMT-Pol. 110656, WA06-H750 (1).
Description. Largest specimen 35 mm long, 9 mm wide for 29 setigers and added to 1 achaetous anterior segment and 4 achaetous preanal segments. Body distinctly fusiform, covered withnumerous vesicles or pustules in several sizes from anterior to posterior end; these vesicles

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Minoru Imajima

becoming large, warty on posterior part of each annulus. Anterior setigers triannulate to about 16(Fig. 45A, C D), thereafter uniannulate (Fig. 45E).
Prostomium small, conical; eyes absent (Fig. 45A). Peristomium uniannulate, achaetous; first
setigers forming mouth (Fig. 45B). Parapodial lappets inconspicuous. Nephridiopores present on
setigers 3 14 (Fig. 45D). Setae all capillaries, with fine hairs. Branchiae simple, cirriform, fromsetiger 2 to 22 (Fig. 45A E). Pygidium with ring of 17 small anal papillae, preceded by 3 6 preanal
constricted setigers.
The species is newly added to the Japanese polychaetous fauna.
Distribution. Alaska to western Mexico; Sea of Okhotsk; Japan.

Fig. 45. Travisia pupa Moore. —A, anterior end, dorsal view, × 11; B, same, ventral view, × 11; C, right parapodium from setiger 3, dorsal view, × 25; D, setigers 10 and 11, lateral view, × 17; E, setigers 18 and 19, sameview, × 17.

Deep-sea Benthic Polychaetes off Northern Honshu

141

Family Scalibregmatidae Malmgren, 1867
Genus Asclerocheilus Ashworth, 1901
Asclerocheilus beringianus Uschakov, 1955
(Figs. 46A D, 47A F)
Asclerocheilus beringianus Uschakov, 1955: 315, fig. 116I L; Uschakov, 1965: 292, fig. 116I L; Levenstein, 1966: 48; Hobson and Banse, 1981: 60, fig. 12a; Blake, 2000d: 137 138, fig. 6.2.

Material. NSMT-Pol. 109834, WA06-E1200D (1).
Description. Body missing posterior end 19 mm long, 3 mm wide with 22 setigerous segments, somewhat thicker anteriorly, but not swollen or distended, narrowing posteriorly. Body
surface covered with annulated rings, up to 4 per segment, each with square, blocked-shaped elevated pads forming transverse rows on dorsum and ventrum (Fig. 46A B).
Prostomium reduced, triangular, bearing 2 round frontal horns; prostomium may be withdrawn deeply into peristomium, only fully visible in frontal view (Fig. 46C). Multiple nuchal organs visible on either side of prostomium, large and bulbous; eyes absent. Peristomium narrowachaetous ring, forming lateral lips around mouth (Fig. 46C).

Fig. 46. Asclerocheilus beringianus Uschakov. —A, anterior end, dorsal view, × 14; B, same, ventral view, × 14;
C, prostomium and setiger 1, frontal view, × 27; D, anterior end, lateral view, × 14.

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Minoru Imajima

Fig. 47. Asclerocheilus beringianus Uschakov. —A, right parapodium from setiger 1, anterior view, × 36; B, same fromsetiger 15, same view, × 36; C D, hirsute spines from setiger 1 (C) and setiger 2 (D), × 367; E F, lyrate setae, ×
500.

Setigers 1 and 2 not subdivided by annuli (Fig. 46A D); setiger 3 triannulate; setiger 4 andfollowing segments quadriannulate; annuli divided dorsally and ventrally into lumpy blocked pads
providing complex areolated appearance to body surface. Pads of ventral midline of setigers 3 and4, forming distinct longitudinal row, large, rectangular, and biannulate from setiger 5 (Fig. 46B).
Branchiae absent.
Parapodia swollen, with setal tori throughout; parapodial cirri and lamellae absent (Fig.
47A B). Interramal sense organs small, inconspicuous, occurring from setiger 1 (Fig. 47D). All
setigers with noto- and neuropodial fascicles of slender capillaries, setigers 1 and 2 with additional 2 rows of curved, hirsute notopodial spines (Fig. 47C D); these sharply pointed, 17 18
spines in setiger 1, 7 10 spines in setiger 2 (Fig. 47A). Setiger 3 and following with capillaries andlyrate setae (Fig. 47E F), few at first, then up to 10 per fascicle, with unequal tynes bearing distinct
bristles on inner margins. Pygidium unknown.
The species is newly added to the Japanese polychaetous fauna.
Distribution. Bering Sea; northern California; Japan.
Genus Pseudoscalibregma Ashworth, 1901
Pseudoscalibregma orientalis sp. nov.
(Figs. 48A D, 49A G)
Type material. Holotype—NSMT-Pol. H 522, Stn. WA05-GH510D, off Kashima Sea,

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143

Fig. 48. Pseudoscalibregma orientalis sp. nov. —A, entire animal in dorsal view, × 5; B, anterior end, dorsal view,
× 15; C, same, lateral view, × 20; D, same, ventral view, × 20.

36˚41.06´N, 141˚22.04´E, 512 508 m, Nov. 11, 2005. Paratypes—NSMT-Pol. P 523, Stn. WA05GH510D, same locality as holotype (8); NSMT-Pol. P 524, Stn. WA05-E1000D, off Sanriku,
1005 1004 m, Oct. 26, 2005 (1); NSMT-Pol. P 525, Stn. WA05-GH380D, off Kashima Sea,
378 373 m, Nov. 12, 2005 (7); NSMT-Pol. P 526, Stn. WA06-EF425D, off Sanriku, 420 424 m,
Nov. 21, 2006 (2); NSMT-Pol. P 527, Stn. WA06-F650D, off Sanriku, 647 641 m, Oct. 31, 2006(1); NSMT-Pol. P 528, Stn. WA06-GH480D, off Kashima Sea, 483 478 m, Nov. 18, 2006 (18).
Description. Body more or less arenicoliform, anterior half expanded to variable degree, posterior region tapered. Holotype consisting of anterior region of 21 mm long, 11 mm wide with 13
setigers, and posterior region of 25 mm long, 5 mm wide with 24 setigers (Fig. 48A). Largest

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paratype 43 mm long for 37 setigers.
Prostomium T-shaped, with lateral processes projecting anterolaterally, nuchal organs visible
on either side of prostomium, eyes lacking (Fig. 48B). Peristomium achaetous, not concealing
posterior margin of prostomium, narrow dorsally and laterally, but broad ventrally. Mouth ventral,
represented by short transverse slit at junction between peristomium and setiger 1. Proboscis papillated sac, with 24 25 longitudinal rows of small conical papillae (Fig. 48C D).
All setigers with biannulate. Body surface of anterior region tessellate, each segmental annulation with single series of raised rectangular or square pads. Epidermal pads above notopodiaof setigers 1 3 inconspicuous. Venter with distinct mid-ventral row of large epidermal pads; pads
bulbous, rectangular, becoming smaller and biannulate from about setiger 6 7. Branchiae absent.
Parapodia on first 13 setigers small and conical with low presetal lobes (Fig. 49A C), without
dorsal and ventral cirri; from setiger 14 noto- and neuropodia developed, with dorsal and ventral
cirri (Fig. 49A), thereafter increasing in size, continuing to posterior end. Noto- and neuropodia inposterior region elongated, each subequal in size, with distally rounded pre- and postsetal lobes(Fig. 49D E). Dorsal and ventral cirri digitate, conspicuous, dorsal cirri longer than ventral ones.
In posteriormost region preanal segments lacking parapodia and setae. Interramal sense organs
from setiger 1 as single small ciliated papillae. Pygidium with smooth margin around anal aperture.
All setigers with slender capillaries in both rami, with those of anterior fascicles more numerous. Setiger 1 with additional row of very short, slender, inconspicuous, pointed and bifurcate
spines (Fig. 49F). Setiger 2 and following with furcate setae in same anterior position as shortspines of setiger 1; each furcate seta with unequal tynes having numerous spinose on inner margin(Fig. 49G).
Remarks. Following Ashworth (1901) and Kudenov and Blake (1978), the scalibregmid genus Pseudoscalibregma included P. parvum (Hansen, 1878) from Norway, P. pallens Levenstein,
1962 from Central Pacific and P. chilensis (Hartmann-Schröder, 1965) from Chile.
Pseudoscalibregma orientalis is distinguishable from above three species in having distinctly
more robust noto- and neuropodia in the posterior region.
The genus is reported as a part of the Japanese fauna for the first time.
Etymology. The species is named because it is the first species of the genus, Pseudoscalibregma from the Asian district.
Distribution. Japan (373 1005 m depth).
Genus Scalibregma Rathke, 1843
Scalibregma inflatum Rathke, 1843
Scalibregma inflatum: Okuda, 1936: 148 149, textfig. 1; Okuda, 1938a: 99; Imajima and Hartman, 1964: 305; Imajima,
1997a: 206; Imajima, 2001a: 87.

Material. NSMT-Pol. 109835, WA05-DE250D (7), NSMT-Pol. 109836, WA05-EF250D (1),
NSMT-Pol. 109837, WA05-FG250D (8); NSMT-Pol. 109838, WA06-B750D (1), NSMT-Pol.
109839, WA06-DE280D (7), NSMT-Pol. 109840, WA06-E510D (1), NSMT-Pol. 109841, WA06GH480D (1); NSMT-Pol. 110868, WA07-B410D (1). NSMT-Pol. 110869, SO07-K2 (1).
Distribution. Norway; Atlantic and Pacific oceans; Alaska; Japan.

Deep-sea Benthic Polychaetes off Northern Honshu

Fig. 49. Pseudoscalibregma orientalis sp. nov. —A, right parapodia on setigers 12 16, dorsal view, × 10; B, leftparapodium from setiger 1, anterior view, × 28; C, same from setiger 8, same view, × 28; D, same from setiger
15, same view, × 27; E, same from setiger 25, posterior view, × 28; F, bifurcate spines from setiger 1, × 578;
G, furcate seta, × 578.

145

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Minoru Imajima

Order Oweniida
Family Oweniidae Rioja, 1917
Genus Galathowenia Kirkegaad, 1959
Galathowenia oculata (Zaks, 1922)
Myriochele oculata Zaks, 1922: 171 174, figs. 1 3; Nilsen and Holthe, 1985: 23 25, figs. 7 8.
Galathowenia oculata: Kirkegaard, 1983: 604 605; Blake, 1984: 114; Imajima and Morita, 1987: 94 97, figs. 6a j, 8c d;
Imajima, 1994: 119 120; Imajima, 1997a: 207; Blake, 2000e: 103 104, fig. 5.1.

Material. NSMT-Pol. 110038, WA05-DE250D (97), NSMT-Pol. 110064, WA05-DE380D(33), NSMT-Pol. 110065, WA05-FG250D (2); NSMT-Pol. 110548, WA06-D210D (1), NSMTPol. 110549, WA06-DE280D (2).
Distribution. Kara Sea; Norwegian Sea; Bering Sea; British Colombia; Japan.
Galathowenia scotiae (Hartman, 1978)
Myriochele scotiae Hartman, 1978: 188 190, fig. 32a d.
Galathowenia scotiae: Parapar, 2001: 404 412, figs. 1 2, tab. 1.
Galathowenia wilsoni Blake, 1984: 114 116, fig. 2a d; Imajima and Morita, 1987: 98, figs. 7a k, 8e f; Imajima, 1997a: 207;
Imajima, 2001a: 88.

Material. NSMT-Pol. 110550, WA05-DE250D (58), NSMT-Pol. 110551, WA06- DE380D(2), NSMT-Pol. 110043, WA05-E1000D (975), NSMT-Pol. 110552, WA05-FG250D (4); NSMTPol. 110740, WA06-A1200D (1), NSMT-Pol. 110741, WA06-E1200D (150), NSMT-Pol. 110742,
WA06-F1500D II (3), NSMT-Pol. 110743, WA06-G900D (15).
Distribution. Ross Sea; Amundsen Sea; Japan.
Genus Myriochele Malmgren, 1867
Myriochele heeri Malmgren, 1867
Myriochele heeri Malmgren, 1867a: 101 102, tab. 7, fig. 37; Blake and Dean, 1973: 37, fig. 2; Nilsen and Holthe, 1985:
21 22, figs. 3 4, 11c e; Imajima and Morita, 1987: 90 91, figs. 3a k, 4e h; Imajima, 1997a: 208; Imajima, 2001a: 88.

Material. NSMT-Pol. 110744, WA05-DE250D (31), NSMT-Pol. 110745, WA05-DE380D(39), NSMT-Pol. 110746, WA05-EF250D (2), NSMT-Pol. 110747, WA05-FG250D (18); NSMTPol. 110748, WA06-D210D (54), NSMT-Pol. 110749, WA06-D450D (1), NSMT-Pol. 110750,
WA06-E510D (4), NSMT-Pol. 110751, WA06-EF425D (6), NSMT-Pol. 110752, WA06-G900D(14).
Distribution. Norwegian Sea; South Atlantic Ocean; Polar Sea; Sea of Okhotsk; Japan.
Myriochele olgae Blake, 2000
Myriochele olgae Blake, 2000e: 111 113, fig. 5.5.

Material. NSMT-Pol. 110753, WA05-DE250D (31), NSMT-Pol. 110754, WA05-DE380D(4), NSMT-Pol. 110755, WA05-E1000D (1), NSMT-Pol. 110756, WA05-FG250D (2).
The species is newly added to the Japanese polychaetous fauna.
Distribution. California; Japan.

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147

Myriochele sp.
Material. NSMT-Pol. 110872, KT07-29-H-2 (2), NSMT-Pol. 110870, KT07-29-M-3-2 (7),
NSMT-Pol. 110871, KT07-29-M-3-3 (1).
Genus Owenia dell Chiaje, 1842
Owenia spp.
Material. NSMT-Pol. 110765, WA05-DE250D (100), NSMT-Pol. 110766, WA05-DE380D(4); NSMT-Pol. 110767, WA06-A150D (1); NSMT-Pol. 110768, WA07-A250D (4). NSMT-Pol.
110873, KT07-29-H-1 (3).
Order Terebellida
Family Pectinariidae Quatrefages, 1865
Genus Amphictene Savigny, 1818
Amphictene japonica (Nilsson, 1928)
Pectinaria (Amphictene) japonica Nilsson, 1928: 52 54, textfig. 16.
Pectinaria japonica: Okuda, 1934: 321 324, textfigs. 1 3; Imajima and Hartman, 1964: 327.

Material. NSMT-Pol. 110044, WA05-DE250D (266), NSMT-Pol. 110045, WA05-DE380D(32), NSMT-Pol. 110046, WA05-E1000D (36), NSMT-Pol. 110047, WA05-EF250D (35), NSMTPol. 110048, WA05-FG250D (4), NSMT-Pol. 110049, WA05-G280 (1); NSMT-Pol. 110050,
WA06-A1200D (5), NSMT-Pol. 110051, WA06-D450D (2), NSMT-Pol. 110052, WA06-DE280D(1), NSMT-Pol. 110053, WA06-E510D (1), NSMT-Pol. 110054, WA06- E1200D (77), NSMT-Pol.
110055, WA06-EF425D (9), NSMT-Pol. 110056, WA06-F1500D II (22), NSMT-Pol. 110057,
WA06-FG350D (1), NSMT-Pol. 110058, WA06-G900D (5), NSMT-Pol. 110059, WA06-G1200D(10), NSMT-Pol. 110060, WA06-GH380 (1), NSMT-Pol. 110063, WA06-GH480D (5), NSMTPol. 110061, WA06-H250D (9), NSMT-Pol. 110062, WA06- H1500D (111); NSMT-Pol. 110702,
WA07-A1500D (4), NSMT-Pol. 110703, WA07-B1200 (4), NSMT-Pol. 110704, WA07-C1500D(12), NSMT-Pol. 110705, WA07-D900 (33). NSMT-Pol. 110874, SO07-C7-B (5). NSMT-Pol.
110863, KT07-29-M-3-3 (2).
Distribution. Japan.
Genus Cistenides Malmgren, 1866
Cistenides hyperborea Malmgren, 1866
Cistenides hyperborea: Moore, 1903: 479.
Pectinaria (Cistenides) hyperborea: Okuda, 1937c: 56 57, pl. 2, fig. F, textfig. 5.

Material. NSMT-Pol. 110706, WA07-A250D (8).
Distribution. Atlantic Ocean; Arctic boreal regions; Japan.
Family Sabellariidae Johnston, 1865
Genus Lygdamis Kinberg, 1867
Lygdamis giardi (McIntosh, 1885)
Sabellaria (Pallasia) giardi McIntosh, 1885: 421, pl. 47, fig. 7; pl. 26A, figs. 13 15.

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Minoru Imajima

Eupallasia giardi: Augener, 1927: 236, textfig. 14.
Lygdamis giardi: Okuda, 1938b: 237 241, figs. 1 3; Imajima, 2001a: 89.

Material. NSMT-Pol. 110553, WA06-GH425 (1).
Distribution. Western Australia; Japan.
Family Ampharetidae Malmgren, 1867
Genus Amage Malmgren, 1866
Amage auricula Malmgren, 1866
Amage auricula Malmgren, 1866: 371; Marenzeller, 1884: 198 199, pl. 2, fig. 6; Hessle, 1917: 120 121; Imajima and Hartman, 1964: 330; Imajima, 1997a: 209 210.

Material. NSMT-Pol. 110920, WA05-DE380D (1).
Distribution. Boreal Atlantic and Arctic oceans; Japan.
Amage delus (Chamberlin, 1919)(Fig. 50A N)
Sabellides delus Chamberlin, 1919: 455 456, pl. 77, fig. 13.
Amage delus: Fauchald, 1972b: 278 280, pl. 56, figs. a c.

Material. NSMT-Pol. 110191, WA06-A650 (3), NSMT-Pol. 110192, WA06-E510D (2), NSMT-Pol. 110193, WA06-E550 (1); NSMT-Pol. 110461, WA07-A550 (16), NSMT-Pol. 110462,
WA07-A650 (11), NSMT-Pol. 110463, WA07-A750 (2), NSMT-Pol. 110464, WA07-C900 (20).
NSMT-Pol. 110834, SO07-C7-B (13), NSMT-Pol. 110885, SO07-O3 (1).
Description. Largest specimen 30 mm long, 6 mm wide at thorax, consisting of 14 thoracic
setigers, 11 uncinigerous and 12 abdominal segments. Body short, tumid, with wide and squarethorax; abdomen quickly tapering to small pygidium.
Prostomium with rounded mid-superior lobe bearing lateral extensions, posteriorly borderedby transverse nuchal organs; U-shaped inferior lobe surrounding superior lobe (Fig. 50 A B). Segments 1 and 2 fused dorsally and laterally, separate ventrally, both forming large lower lip (Fig.
50C); buccal tentacles smooth. Paleae absent. Four pairs of smooth, large branchiae on 3 visible
segments, arranged nearly segmentally with 2 pairs on segment 3 (setiger 1) and 1 pair each on
segments 4 and 5 (setigers 2, 3); posterior pair connected to base of notopodia by thickened ridges(Fig. 50D E).
Segments 3 and 4 fused laterally, bearing first notopodia; reduced, displaced ventrally byouter branchia, bearing few fine, short setae (Fig. 50E F). Subsequent notopodia gradually lengthening, conical throughout with short, triangular postsetal lobe in addition to conical ventral cirri(Fig. 50G). Neuropodia with uncini first present on setiger 4, paddle-shaped without dorsal cirri(Fig. 50D, G). Abdominal notopodial rudiments large, bluntly digitiform and curved inward (Fig.
50H); neuropodia progressingly elongate toward pygidium, last pair projecting beyond anus, withsmall dorsal cirri (Fig. 50N). Notosetae fairly stout, limbate (Fig. 50I); thoracic and abdominal
uncini with 4 5 teeth in single row above inconspicuous rostral point and rounded basal prow (Fig.
50J M).
Pygidium with 2 thick, clavate dorsolateral anal cirri and several low, rounded ventral lobes(Fig. 50I).
The species is newly added to the Japanese polychaetous fauna.
Distribution. Southern California; Japan.

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Fig. 50. Amage delus (Chamberlin). —A, anterior end, dorsal view, × 7; B, prostomium, same view, × 15; C; anterior end, ventral view, × 7; D, anterior end, lateral view, × 7; E, segments 3 to 6, same view, × 15; F, leftparapodia of segments 3 to 7, dorsal view, × 15; G, last thoracic parapodium, anterior view, × 15; H, abdominal parapodium, same view, × 15; I, notoseta, × 120; J K, thoracic uncini, × 705; L M, abdominal uncini, ×705; N, posterior end, ventral view, × 22.

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Minoru Imajima

Genus Ampharete Malmgren, 1866
Ampharete longipaleolata Uschakov, 1950
(Fig. 51A N)
Ampharete longipaleolata Uschakov, 1950: 218 219, pl. 2, fig. 33; Uschakov, 1955: 367 368, fig. 136E G.

Fig. 51. Ampharete longipaleolata Uschakov. —A, last thoracic and anterior abdominal segments, dorsal view, ×
6; B, anterior end, same view, × 9; C, same from others, same view, × 9; D E, paleae, × 140; F, thoracic parapodium, anterior view, × 25; G, notoseta, × 127; H J, thoracic uncini, lateral and frontal views, × 748; K, abdominal parapodium, anterior view, × 25; L M, abdominal uncini, lateral and frontal views, × 957; N, posterior end, ventral view, × 17.

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Material. NSMT-Pol. 110194, WA05-DE380D (2), NSMT-Pol. 110195, WA05-FG410 (1),
NSMT-Pol. 110196, WA05-FG425 (1), NSMT-Pol. 110197, WA05-FG450 (1); NSMT-Pol.
110198, WA06-E450 (4), NSMT-Pol. 110199, WA06-E480 (3), NSMT-Pol. 110200, WA06-E510(5), NSMT-Pol. 110201, WA06-E510D (5), NSMT-Pol. 110202, WA06-E550 (1), NSMT-Pol.
110203, WA06-F410 (1), NSMT-Pol. 110204, WA06-F480 (1); NSMT-Pol. 110468, WA07A1500D (1).
Description. Largest specimen 43 mm long, 4.5 mm wide at thorax, consisting of 14 thoracic
setigers, 12 uncinigerous and 13 abdominal segments. Body slender, widest anteriorly, graduallytapering toward pygidium, with distinct constriction between third and fourth abdominal segment(Fig. 51A).
Prostomium roughly pentagonal, with large mid-superior lobe surrounded by inferior lobe,
with nuchal organs at postectal corners of mid-superior lobe (Fig. 51B). Segments 1 and 2 separate
dorsally and ventrally, with segment 1 forming smooth lower lip; segment 2 very short but distinct.
Buccal tentacles slender, smooth.
Segment 3 with anterior 3 pairs of branchiae and paleae; branchiae subulate, usually slightly
separated by narrow dorsomedian gap and bases of inner branchiae fused for short length (Fig.
51C). Paleae wide, very long, about 4 times as long as prostomium, abruptly tapering to acute tips,
in crowded semicircular fascicles 22 24 per fascicle (Fig. 51B, D E). Segments 4 and 5 fused laterally, bearing bases of fourth pair of branchiae fused to dorsal body wall, branchia emerging between inner and middle anterior pairs and 2 minute nephridial papillae just inside branchial bases,
carrying small first pair of notopodia (Fig. 51B).
Following notopodia cylindrical (Fig. 51F), with limbate notosetae, tapering to acute tip (Fig.
51G). Thoracic uncinigers first present on setiger 3, with short neuropodial cirri. Uncini withdouble row of 5 7 teeth above rostral point and basal prow (Fig. 51H J). Anterior abdominal segments with tiny, papilliform notopodial rudiments (Fig. 51K), posteriorly gradually decreasing andreplaced by low glandular ridges above neuropodial lobes. Neuropodia with short, conical dorsal
cirri; uncini smaller than thoracic uncini, with 2 3 rows of teeth (Fig. 51L M).
Pygidium short, narrow, with 2 slender lateral cirri and circlet of low, rounded papillae between cirri (Fig. 51N).
The species is newly added to the Japanese polychaetous fauna.
Distribution. Sea of Okhotsk; Japan.
Genus Amphicteis Grube, 1850
Amphicteis mederi Annenkova, 1929
(Figs. 52A D, 53A I)
Amphicteis mederi Annenkova, 1929: 499 500, pl. 38, figs. 45 46, pl. 39, fig. 58; Uschakov, 1955: 376, fig. 139K M.

Material. NSMT-Pol. 110205, WA05-DE380D (9), NSMT-Pol. 110206, WA05-FG510D (2),
NSMT-Pol. 110207, WA05-GH510D (6), NSMT-Pol. 110208, WA05-H900D (2); NSMT-Pol.
110209, WA06-A150D (2), NSMT-Pol. 110210, WA06-A250D (1), NSMT-Pol. 110211, WA06DE280D (1), NSMT-Pol. 110212, WA06-G900D (1); NSMT-Pol. 110469, WA07-A250D (4),
NSMT-Pol. 110470, WA07-D900 (1).
Description. Largest specimen 42 mm long, 5 mm wide at thorax, consisting of 17 thoracic
setigers, 14 uncinigerous and 15 abdominal segments. Body long, slender, gently tapering toward
pygidium.
Prostomium longer than wide, with mid-superior lobe divided into 2 straight, thick ridges;
posterior prostomial lobe surrounding mid-superior lobe, bearing long, conspicuous glandular

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Fig. 52. Amphicteis mederi Annenkova. —A, anterior end, dorsal view, × 9; B, prostomium, same view, × 14; C,anterior end, lateral view, × 9; D, anterior segments, dorsolateral view, × 26.

ridges on anterior margin; central part of prostomium bearing 2 lateral groups of small eyespots,
usually concealed by glandular ridges (Fig. 52A B). Single papilla occurring between glandular
ridges (Fig. 52B). Segments 1 and 2 fused, produced into crenulated lower lip ventrally. Branchiae
long, with broad bases leaving fairly large scars when breaking off (Fig. 52A, D), tapered to filiform tips (Fig. 52C); in 2 groups of 2, separated by narrow median gap occupied by notched branchial pad (Fig. 52A).
Paleae well developed, about 9 10 per fascicle (Fig. 52A), coppery, tapering to slightly
hooked with rounded tip (Fig. 53A). First 2 notopodia on segments 4, 5 small and conical, withtriangular postsetal lobes and fine setae (Fig. 52D); subsequent notopodia larger, with distinct foliaceous ventral cirri and conical presetal lobes (Fig. 53B). Fully developed notosetae stiff, limbate. Thoracic neuropodia from segment 7 (setiger 4) well-developed, paddle-shaped; all neuropodia in same width. Thoracic uncini with single row of 5 teeth above inconspicuous rostral point andnarrow basal prow (Fig. 53C D). Abdominal notopodia rudiments present throughout, small but
distinct, involute-digitiform to rounded (Fig. 53E). Abdominal neuropodia narrower than thoracic
ones, gradually lengthening toward pygidium, with short dorsal cirri (Fig. 53E). Abdominal uncini

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153

Fig. 53. Amphicteis mederi Annenkova. —A, palea, × 70; B, thoracic parapodium, anterior view, × 25; C D, thoracic uncini, lateral and frontal views, × 717; E, abdominal parapodium, anterior view, × 32; F H, abdominal
uncini, lateral and frontal views, × 717; I, posterior end, ventral view, × 17.

smaller than thoracic ones, with single row of 5 teeth above small rostral point and narrow basal
prow (Fig. 53F H).
Pygidium small, with 2 long, filiform lateral anal cirri and 6 dorsal and 5 ventral rounded
lobes (Fig. 53I).
The species is newly added to the Japanese polychaetous fauna.
Distribution. Seas of Japan and Okhotsk; Kuril Islands; Japan.
Amphicteis scaphobranchiata Moore, 1906
(Figs. 54A H, 55A H)
Amphicteis scaphobranchiata Moore, 1906: 255 257, pl. 12, figs. 54 61; Berkeley and Berkeley, 1952: 68 69, figs. 139 141;
Uschakov, 1955: 376; Imajima, 1961: 95 96, textfig. 11a c; Hartman, 1969: 549 550, figs. 1 6; Fauchald, 1972b:
286 287, pl. 58, fig. c; Uebelacker, 1984b: 51-18 20, figs. 51-13, 14a e; Hilbig, 2000: 188 190, fig. 8.7.

Material. NSMT-Pol. 110213, WA06-E510D (5), NSMT-Pol. 110214, WA06-EF425D (7),
NSMT-Pol. 110215, WA06-F750 (2), NSMT-Pol. 110216, WA06-H480 (2), NSMT-Pol. 110217,
WA06-H1500D (3); NSMT-Pol. 110471, WA07-A650 (1).
Description. Largest specimen 47 mm long, about 5 mm wide at thorax, consisting of 17
thoracic setigers, 14 uncinigerous and 15 abdominal segments. Body with wide thorax tapering
toward pygidium.

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Fig. 54. Amphicteis scaphobranchiata Moore. —A, anterior end, branchiae omitted, dorsal view, × 11; B, anterior
end, same view, × 16; C, anteromedial branchia, × 18; D, anteromedial and anterolateral branchiae, × 16; E,
palea, × 67; F, thoracic parapodium, lateral view, × 32; G, abdominal parapodium, same view, × 23; H, notoseta,
× 108.

Prostomium wider than long, with mid-superior lobe divided into 2 straigh, thick ridges
joined by thin, lamellar middle part. Posterior prostomial lobe bearing long, conspicuous glandular
ridges on anterior margin; central part of prostomium bearing 2 lateral groups of small eyespots
and pairs nuchal organs median to eyes, usually concealed by glandular ridges (Fig. 54A).
Segments 1 and 2 fused, produced ventrally into smooth lower lip. Branchiae large, as longas 9 setigers, anteromedial pair broadly foliaceous with scalloped anterior margin bearing a

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155

Fig. 55. Amphicteis scaphobranchiata Moore. —A B, thoracic uncini, lateral view, × 935; C E, abdominal anterior uncini, lateral and frontal views, × 935; F G, abdominal posterior uncini, lateral and frontal views, × 935;
H, posterior end, ventral view, × 25.

filiform hooked tip, others cylindrical and tapered to filiform tips (Fig. 54B D); in 2 rows of 2,
separated by broad median gap occupied by large branchial membrane or pad about half as wide
as branchial bases (Fig. 54A).
Paleae well developed, tapering abruptly to hairlike tips (Fig. 54E), about 15 23 per fascicle.
First 2 notopodia small and conical, with triangular postsetal lobes and few fine setae (Fig. 54A);
subsequent notopodia larger, with foliaceous ventral cirri in addition to presetal lobes (Fig. 54F);
thoracic uncinigers well-developed, paddle-shaped with conical dorsal cirri. Abdominal notopodial rudiments present throughout involute-digitiform to rounded (Fig. 54G); neuropodia progressingly elongate toward pygidium, last pair projecting beyond anus, with small dorsal cirri. Notosetae fairly stout, limbate (Fig. 54H); thoracic (Fig. 55A B) and anterior abdominal uncini (Fig.
55C E) with 5 6 teeth in single row above inconspicuous rostral point and rounded basal prow;
middle and posterior abdominal uncini with several rows of teeth above large basal prow (Fig.
55F G).
Pygidium with 2 slender, clavate dorsolateral anal cirri and several low, rounded lateral andventral lobes (Fig. 55H).
The species is newly added to the Japanese polychaetous fauna.
Distribution. British Colombia to California; Gulf of Mexico; Sea of Okhotsk; Japan.
Genus Amphisamytha Hessle, 1917
Amphisamytha japonica Hessle, 1917
(Fig. 56A L)
Amphisamytha japonica Hessle, 1917: 114 115, pl. 1, figs. 7 8; Imajima and Hartman, 1964: 332.

Material. NSMT-Pol. 110218, WA06-H250D (1).
Description. Specimen 16 mm long, 3 mm wide at thorax, consisting of 17 thoracic setigers,
14 uncinigerous and 13 abdominal segments. Body slender, widest anteriorly, gradually tapering
toward pygidium.

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Fig. 56. Amphisamytha japonica Hessle. —A, anterior end, dorsal view, × 12; B, anterior end, lateral view, × 26;
C, thoracic parapodium, anterior view, × 26; D, notoseta, × 170; E, abdominal neuropodia, lateral view, × 26;
F I, thoracic uncini, lateral and frontal views, × 990; J K, abdominal uncini, lateral view, × 990; L, posterior
end, ventral view, × 26.

Prostomium distinctly trilobed, with large mid-superior lobe surrounded by inferior lobe,
with eyes at postectal corners of mid-superior lobe (Fig. 56A). Segments 1 and 2 separate ventrally, with segment 1 forming crenulated lower lip; segment 2 short but distinct (Fig. 56B). Segment 3 with first 2 pairs of branchiae; segment 4 with posterior 2 pairs of branchiae and minute first
notopodia bearing few very fine setae. Posterior pairs of branchiae associated with segments 5 and6, bases fused to dorsal body wall; branchiae separated by narrow dorsomedian gap. Branchiae
slender, cylindrical about 1.5 times as long as body width. Paleae absent.
Fully developed biramous parapodia from segment 7 (setiger 4) through segment 20.

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157

Thoracic notopodia slightly inflated to cylindrical, simple throughout with presetal lobes; limbate
notosetae tapering to acute tip. Thoracic uncinigers small rounded pinnules with crowded uncini(Fig. 56C D). Abdominal neuropodia longer than thoracic, lengthening toward posterior end, lastpair reaching edge of pygidium; parapodial cirri and notopodial rudiments absent (Fig. 56E). Thoracic uncini with 2 3 rows of 6 teeth above basal prow (Fig. 56F I); abdominal uncini smaller thanthoracic, with 3 rows of 5 teeth above basal prow (Fig. 56J K).
Pygidium with 2 short ventrolateral anal cirri (Fig. 56L).
Remarks. The branchiae of the type specimen from Sagami Bay, in 300 m are arranged as 3
anterior and 1 posterior pair, whereas the examined specimen has 2 anterior and 2 posterior pairs.
Hilbig (2000) said; the branchiae on 2 or 3 visible segments in this genus, arranged 3+1, 2+2, or
2+1+1 are depending on the degree of relaxation at the time of death, therefore that branchial configurations are not a good character in this genus.
Distribution. Japan.
Genus Anobothrus Levinsen, 1883
Anobothrus gracilis (Malmgren, 1866)(Figs. 57A G, 58A H)
Ampharete gracilis Malmgren, 1866: 365, pl. 26, fig. 75.
Anobothrus gracilis: Levinsen, 1883: 162; Hilbig, 2000: 192 194, fig. 8.9.

Material. NSMT-Pol. 110219, WA05-EF250D (1), NSMT-Pol. 110220, WA05-FG510D (2),
NSMT-Pol. 110221, WA05-G280 (3), NSMT-Pol. 110222, WA05-GH510D (14); NSMT-Pol.
110223, WA06-H250D (16). NSMT-Pol. 110472, WA07-A1500D (1), NSMT-Pol. 110473, WA07B1500D (1), NSMT-Pol. 110474, WA07-C1500D (1), NSMT-Pol. 110475, WA07-D210D (1).
NSMT-Pol. 110224, SO06-M3-B1 (1), NSMT-Pol. 110803, SO07-K3 (3).
Description. Largest specimen 26 mm long, about 3 mm wide at thorax, consisting of 16
thoracic setigers including paleae, 12 uncinigerous and 14 abdominal segments. Body slender,
linear, tapering posteriorly to rounded pygidium, with more or less distinct constriction between
second and third abdominal segment (Fig. 57A).
Prostomium roughly pentagonal, with large mid-superior lobe surrounded by inferior lobe,
with nuchal organs at postectal corners of mid-superior lobe, eyespots not recognized (Fig. 57B).
Peristomium and segment 2 fused laterally and ventrally, produced into smooth lower lip. Fourpairs of smooth branchiae cylindrical, slender, on 2 visible segments, arranged in a transverse rowof 3 pairs and single pair behind on either side, left and right group not separated by dorsomediangap and bases of inner branchiae fused for short length (Fig. 57B D).
Paleae slender, tapering gradually to hairlike tips (Fig. 57E), in crowded semicircular fascicles of about 20 (Fig. 57B). First notopodia on segment 4, smallest, directly behind base of outermost branchia, reduced to thin fascicle of short, very fine setae (Fig. 57B D). Following notopodia
increasing in size, cylindrical, without lobes or cirri (Fig. 58A), largest in mid thoracic segments,
bearing larger, limbate setae (Fig. 57F). Thoracic uncinigers beginning on setiger 4 wide, paddleshaped; abdominal neuropodia (Fig. 58B) narrower and gradually lengthening toward pygidium.
Notopodia in setiger 5 (unciniger 2) connected by often weakly defined dorsal ridge; notopodia of fifth-to-last setiger (setiger 11, unciniger 8) slightly elevated dorsally, connected by thickened dorsal glandular ridge (Fig. 57C D), and bearing modified setae with finely pilose, more
abruptly tapering tips (Fig. 57G). Thoracic uncini in a single series numbering 60 in anterior unciniger, and fewer in the more posterior unciniger. Thoracic uncini avicular with doble row of 7teeth above rounded basal prow (Fig. 58C E), abdominal uncini with 3 4 rows of 3 7 teeth above

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Minoru Imajima

Fig. 57. Anobothrus gracilis (Malmgren). —A, last thoracic and three abdominal segments, lateral view, × 12; B,
anterior end, dorsal view, × 20; C, anterior end, same view, × 12; D, same, lateral view, × 12; E, tip of palea,
× 100; F, tip of regular notoseta, × 245; G, tip of modified notoseta, × 287.

rostral point and basal prow (Fig. 58F G).
Pygidium with terminal anus surrounded by 2 large dorsal and 3 small ventrolateral papillae(Fig. 58H). Tubes with mucous inner lining and thin outer layer of fine mud particles.
The species is newly added to the Japanese polychaetous fauna.
Distribution. Arctic and boreal Atlantic Ocean; California; Japan.

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159

Fig. 58. Anobothrus gracilis (Malmgren). —A, thoracic parapodium, × 38; B, anterior abdominal unciniger, × 38;
C E, thoracic uncini, lateral and frontal views, × 1045; F G, abdominal uncini, lateral and frontal views, ×1045; H, posterior end, ventrolateral view, × 28.

Anobothrus wakatakamaruae sp. nov.
(Figs. 59A F, 60A L)
Type material. Holotype—NSMT-Pol. H 529, Stn. WA05-EF450D, off Sanriku, 38˚02.18´N,
142˚04.82´E, 452 454 m, Nov. 18, 2005. Paratypes—NSMT-Pol. P 530, Stn. WA05-FG510D, off
Sanriku, 516 515 m, Nov. 15, 2005 (3).
Description. Holotype 15 mm long, about 1.8 mm wide at thorax, consisting of 14 thoracic
setigers, 12 uncinigerous and 12 abdominal segments. Body slender, linear, gently tapering toward
pygidium, with distinct constriction between third and fourth abdominal segment (Fig. 59A).
Prostomium roughly rectangular, distinctly trilobed with large mid-superior lobe surrounded
by inferior lobe, concealed by paleae and branchial bases; nuchal organs present but eyes not observed (Fig. 59B). Segments 1 and 2 free dorsally, fused and somewhat obscured laterally and freeventrally, produced into smooth lower lip. Buccal tentacles numerous, slender, with 4 longitudinalrows of short pinnae on lateral sides (Fig. 59C). Branchiae about as long as first 11 setigers, cylindrical, slender, arising from segment 3 in a straight row of 3 and 1 behind on either side, separated
by narrow median gap; posterior pair of branchiae associated with segment 6 (setiger 2), bases
fused to dorsal body wall (Fig. 59D E). Paleae prominent, wide and slender, tapering gradually to
long, curved hairlike tips, arising from large cylindrical lobes in dense, fan-shaped fascicles ofabout 25 26 per fascicle (Figs. 59F, 60A). Segments 4 and 5 fused, bearing small but distinct notopodium with developed notosetae; second notopodium on segment 6, also bearing developed
notosetae (Fig. 60B). Fully developed, biramous parapodia from segment 7. Notopodia 4 and 5
connected by dorsal glandular bands (Fig. 59D E); notopodia of fourth-to-last setiger (setiger 11)

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Fig. 59. Anobothrus wakatakamaruae sp. nov. —A, last thoracic and four abdominal segments, lateral view, × 10;
B, anterior end, dorsal view, × 27; C, tip of buccal tentacle, × 256; D, anterior end, lateral view, × 14; E, anterior end, dorsal view, × 16; F, paleae in fascicles, frontal view, × 27.

slightly elevated dorsally, connected by prominent, thickened ciliated dorsal ridge and bearing
modified setae with finely pilose (Fig. 60C D).
Thoracic notopodia simple, cylindrical throughout, without lobes or cirri; thoracic uncinigers
first present in setiger 3 wide, paddle-shaped (Fig. 60E). Abdominal neuropodia narrower andgradually lengthening toward pygidium; dorsal cirri absent (Fig. 60F). Thoracic and anterior abdominal uncini with double row of 6 7 teeth above rostral point and rounded basal prow (Fig. 60G,H); middle and posterior abdominal uncini with 3 rows of 6 7teeth above rostral point and rounded basal prow (Fig. 60I K).
Pygidium with terminal anus surrounded by 2 slender lateral cirri and 5 very small papillae in
dorsal arc and 7 papillae in ventral arcs (Fig. 60L).
Remarks. Anobothrus wakatakamaruae resembles A. paleatus Hilbig, 2000 from California

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Fig. 60. Anobothrus wakatakamaruae sp. nov. —A, palea, × 112, with detail of distal region, × 162; B, notoseta, ×206; C, posterior thoracic segments, showing ciliated dorsal ridge, lateral view, × 29; D, modified seta, × 320,with detail of distal region, × 642; E, thoracic parapodium, anterior view, × 34; F, abdominal unciniger, sameview, × 34; G H, thoracic uncini, in lateral and frontal views, × 935; I K, abdominal uncini, lateral and frontal
views, × 935; L, posterior end, viewed from the rear, × 50.

(in 550 to 610 m) in the notopodia of the fourth-to-last setiger (setiger 11, unciniger 9) are connected by thickened dorsal ridge and the position of the abdominal constriction. However, A.
wakatakamaruae can be distinguished from A. paleatus in that the notopodia 4 and 5 are connected by the dorsal glandular ridges rather than only notopodium 5, and both are also differentiated each other in the morphology of the paleae and modified setae.
Etymology. The species is named after the R/V Wakataka-maru, FRA, which collected these
specimens.
Distribution. Japan (452 516 m depth).
Genus Auchenoplax Ehlers, 1887
Auchenoplax crinita Ehlers, 1887
Auchenoplax crinita Ehlers, 1887: 209 214, pl. 44, figs. 10 16; Fauvel, 1936b: 95 96; Hartman, 1965: 216 217, pl. 47, figs.
a d; Imajima, 1997a: 210 211, fig. 13a h.

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Material. NSMT-Pol. 109420, WA05-DE380D (7), NSMT-Pol. 109421, WA05-EF250D (2),
NSMT-Pol. 109422, WA05-EF450D (13), NSMT-Pol. 109423, WA05-FG510D (4); NSMT-Pol.
110225, WA06-DE280D (1), NSMT-Pol. 110226, WA06-E510D (11), NSMT-Pol. 110227, WA06EF425D (17), NSMT-Pol. 110228, WA06-FG350D (2), NSMT-Pol. 110229, WA06-FG480 (1),
NSMT-Pol. 110230, WA06-G900D (3), NSMT-Pol. 110231, WA06-GH480D (15), NSMT-Pol.
110232, WA06-H250D (13).
Distribution. Atlantic of New England; northeastern South America; southern Florida; Japan.
Genus Lysippe Malmgren, 1866
Lysippe labiata Malmgren, 1866
(Fig. 61A M)
Lysippe labiata Malmgren, 1866: 367, pl. 26, fig. 78; Hessle, 1917: 109 110; Berkeley and Berkeley, 1956: 241; Hilbig, 2000:
206 207, fig. 8.16.

Material. NSMT-Pol. 110234, WA05-DE380D (1), NSMT-Pol. 110235, WA05-EF250D (2);
NSMT-Pol. 110236, WA06-E510D (1), NSMT-Pol. 110237, WA06-EF425D (1), NSMT-Pol.
110239, WA06-GH480D (1), NSMT-Pol. 110238, WA06-H250D (4).
Description. Largest specimen 16 mm long, 2 mm wide at thorax, consisting of 16 thoracic
setigers, 13 uncinigerous 12 abdominal segments. Body slender, widest in thorax, gradually tapering toward pygidium.
Prostomium trilobed, with wide mid-superior lobe surrounded by inferior lobes with patchesof brown pigment; nuchal organs at postectal corners of mid-superior lobe, without eyes and glandular ridges (Fig. 61A B). Buccal tentacles few, pinnate. Segments 1 and 2 fused, ventrally produced into large, scoop-shaped, crenulated lower lip usually protruding beyond prostomium andvisible in dorsal view (Fig. 61B). Segment 3 with small, thin paleae arising from large cylindricallobe (Fig. 61C) and anterior 2 pairs of branchiae; posterior 2 pairs of branchiae associated withsegments 4 and 5, bases fused to dorsal body wall; left and right group separated by wide dorsomedian gap (Fig. 61A B). Branchiae slender, cylindrical, as long as 8 setigers (Fig. 61D).
First notopodia on segment 4 reduced in size but bearing developed notosetae; second notopodia on segment 5, also somewhat reduced in size but bearing well-developed notosetae. Notopodia from segment 7 fully developed, cylindrical, without lobes and cirri (Fig. 61E), with stiff,
long setae (Fig. 61F). First thoracic uncinigers on setiger 4 small pinnules, lengthening in posterior abdominal segments; dorsal cirri absent (Fig. 61G). Thoracic uncini with 3 rows of 4 5 teeth
above rostral point and basal prow (Fig. 61H J); abdominal uncini crested, with about 4 rows ofteeth above rostral point and basal prow (Fig. 61K L).
Pygidium with 2 slender lateral anal cirri, about as wide as last segment, and low, rounded
lobes (Fig. 61M).
The species is newly added to the Japanese polychaetous fauna.
Distribution. Arctic; Greenland; Iceland; Denmark; Alaska to California; Sea of Japan, Japan.
Genus Melinna Malmgren, 1867
Melinna cristata (Sars, 1851)
Melinna cristata: Wollebaek, 1912: 65 67, pl. 14, figs. 1 9; Hessle, 1917: 92 93; Uschakov, 1955: 363, fig. 134G; Hartman,
1965: 218 219; Fauchald, 1972b: 418; Yang and Sun, 1988: 280, fig. 134a g; Mackie and Pleijel, 1995: 104 111, figs.
1 3, tab. 1; Imajima, 1997a: 209.

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Fig. 61. Lysippe labiata Malmgren. —A, anterior end, dorsal view, × 29; B, anterior end from others, same view,
× 29; C; palea, × 280; D, anterior end, lateral view, × 29; E, thoracic parapodium, anterior view, × 49; F, notoseta, × 255; G, abdominal unciniger, anterior view, × 49; H J, thoracic uncini, lateral and frontal views, ×1045; K L, abdominal uncini, lateral and frontal views, × 1045; M, posterior end, ventral view, × 42.

Material. NSMT-Pol. 110240, WA05-E1000D (1); NSMT-Pol. 110241, WA06-A1200D (18),
NSMT-Pol. 110242, WA06-E510D (1), NSMT-Pol. 110243,WA06-G900D (1), NSMT-Pol.
110244, WA06-GH480D (1); NSMT-Pol. 110465, WA07-A1500D (3).
Distribution. Arctic and boreal seas; Japan.

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Melinna elisabethae McIntosh, 1914
Melinna elisabethae McIntosh, 1914: 106; McIntosh, 1922: 86 88, pl. 119, fig. 1, pl. 125, figs. 2 2b; Uschakov, 1955: 363;
Banse and Hobson, 1968: 45; Fauchald, 1972b: 418; Banse, 1979: 1549; Imajima, 1994:121 122, fig. 3a k.

Material. NSMT-Pol. 110245, WA06-DE280D (3); NSMT-Pol. 110466, WA07-A250D (1),
NSMT-Pol. 110467, WA07-D210D (2).
Distribution. Arctic Sea; Sea of Okhotsk; Puget Sound, Washington; Japan.
Genus Melinnopsis McIntosh, 1885
Melinnopsis atlantica McIntosh, 1885
(Figs. 62A I, 63A G)
Melinnopsis atlantica McIntosh, 1885: 441 442, pl. 27A, fig. 18.

Material. NSMT-Pol. 110246, WA06-H1500D (7). NSMT-Pol. 110884, SO07-O3 (5).
Description. Largest complete specimen 28 mm long, 2.5 mm wide at thorax, consisting of
18 thoracic segments, 15 setigerous, 13 uncinigerous and 39 abdominal segments. Body slender,
almost linear, widest at level of postbranchial membrane, gradually tapering toward pygidium.
Prostomium wider than long, with scalloped anterior margin; posterior edge developed intoraised transverse glandular ridge, and paired nuchal organs just anterior to ridge (Fig. 62A). Eyes
lacking. Segments 1 and 2 fused to prostomium dorsally, forming smooth, padlike lower lip ventrally. Buccal tentacles smooth, slender to stout.
Segments 3 to 6 fused dorsally and laterally, forming lateral wings with 4 rows of needlelike
neurosetae (Fig. 62B C); segments 3 and 4 fused ventrally, forming ventral fold with smooth edge.
Segment 3 bearing branchiae, arranged 2+2, fused basally, long and subulate. Nuchal hooks absent. Segment 4 with neurosetae only (Fig. 62D); segment 5 with first notosetae, very fine andshort (Fig. 62F), emerging more or less directly from body wall, no visible notopodia. Postbranchial membrane with smooth anterior margin (Fig. 62B). Segment 6 also with small fascicle of finenotosetae and short transverse linear series of needlelike neurosetae (Fig. 62E).
Regular biramous parapodia from segment 7, with developed notopodia and neuropodial pinnules bearing uncini. Notopodia slightly inflated to cylindrical, with presetal lobes and stiff, longsetae; thoracic uncinigers small, paddle-shaped, with short dorsal cirri and numerous crowded
uncini (Fig. 62G). Abdominal notopodial rudiments small conical but distinct; neuropodial pinnules becoming narrower with short linear series of uncini (Fig. 62H). Notosetae in two rows,
anterior short limbate setae and posterior long ones (Fig. 62I). Thoracic uncini with single row of
4 5 teeth over basal prow (Fig. 63A C), abdominal uncini with 2 3 large teeth surmounted by crestof about 7 smaller ones, large rostral point, and rounded basal prow (Fig. 63D F).
Pygidium cylindrical, with extending dorsal papilla and slightly crenulated posterior margin(Fig. 63G).
The species is newly added to the Japanese polychaetous fauna.
Remarks. The holotype collected from off Chesapeake Bay, North America, 1700 fathoms issomewhat damaged, but it has some characteristics peculiar to the species.
Distribution. Off Chesapeake Bay, North America; Japan.

Deep-sea Benthic Polychaetes off Northern Honshu

Fig. 62. Melinnopsis atlantica McIntosh. —A, prostomium, dorsal view, × 14; B, anterior end, same view, × 14; C,
same, lateral view, × 14; D, needlelike neurosetae from segment 4, × 560; E, same from segment 6, × 560; F,
first notosetae, × 300; G, thoracic parapodia, anterior view, × 42; H, abdominal parapodium, same view, × 56;
I, notoseta, × 150.

165

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Fig. 63. Melinnopsis atlantica McIntosh. —A C, thoracic uncini, lateral and frontal views, × 1100; D F, abdominal
uncini, lateral and frontal views, × 1100; G, pygidium, lateral view, × 35.

Genus Paramage Caullery, 1944
Paramage scutata (Moore, 1923)(Fig. 64A M)
Amage scutata Moore, 1923: 210 212, pl. 17, figs. 19 24.
Paramage scutata: Williams, 1987: 254 255, fig. 2B; Hilbig, 2000: 214 215, fig. 8.20.

Material. NSMT-Pol. 110476, WA06-A650 (1), NSMT-Pol. 110477, WA06-B750D (1), NSMT-Pol. 110478, WA06-E510D (2), NSMT-Pol. 110479, WA06-FG350D (1); NSMT-Pol. 110480,
WA07-D900 (1).
Description. Largest complete specimen 36 mm long, 10 mm wide including parapodia at
thorax, consisting of 12 thoracic setigers and 2 asetigerous, 11 uncinigerous and 12 abdominal
segments. Body widest in thorax, tumid, quickly tapering in abdomen toward small pygidium.
Prostomium wider than long, distinctly trilobed, with nuchal organs at postectal corners of
mid-superior lobe, without glandular ridges (Fig. 64A). Paleae absent. Segments 1 and 2 separate
dorsally, fused laterally and ventrally, produced into lower lip and lateral auricular folds. Segment
3 also produced into lateral auricular folds, bearing first 2 pairs of branchiae and 1 pair of vestigial
notopodia. Segments 4 and 5 with 1 pair of branchiae each and vestigial notopodia; vestigial notopodia of segment 5 setigerous, notosetae very fine, barely emerging from parapodium (Fig. 64B).
Branchiae smooth, cylindrical and tapered (Fig. 64C). Segment 6 with fully developed notopodia;
following 10 setigers with regular, biramous parapodia.
Notopodia cylindrical, simple throughout with triangular presetal lobes in addition to foliaceous ventral cirri; uncinigers well-developed, paddle-shaped (Fig. 64D). Abdominal notopodial
rudiments conspicuous, distally inflated, coiled inwards; neuropodia shorter, with short dorsal
cirri (Fig. 64E). Notosetae in dense fascicles, long, slender, narrowly limbate (Fig. 64F). Thoracic
uncini bearing single row of 5 teeth and rounded basal prow (Fig. 64G); inferior thoracic unciniwith three rows of 4 large teeth (Fig. 64H). Abdominal uncini with single or three rows of 4 teeth
over basal prow (Fig. 64I L).

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Fig. 64. Paramage scutata (Moore). —A, anterior end, dorsal view, × 14; B, same, lateral view, × 14; C; branchia,
× 16; D, last thoracic parapodium, anterior view, × 10; E, second abdominal parapodium, same view, × 16; F,
notoseta, × 60; G, superior thoracic uncinus, lateral view, × 880; H, inferior thoracic uncinus, same view, ×880; I L, abdominal uncini, lateral and frontal views, × 880; M, posterior end, ventral view, × 10.

Pygidium with 2 digitiform ventrolateral anal cirri and several small, rounded lobes (Fig.
64M).
The species is newly added to the Japanese polychaetous fauna.
Distribution. California; Japan.

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Genus Samythella Verrill, 1873
Samythella bathycola Uschakov, 1950
(Fig. 65A K)
Samythella bathycola Uschakov, 1950: 220 221, pl. 2, 10, fig. 34; Uschakov, 1955: 374, fig. 139A F.

Material. NSMT-Pol. 110247, WA05-FG450 (1).
Description. Specimen 35 mm long, about 3 mm wide at thorax, consisting of 15 thoracic
setigers, 12 uncinigerous and 28 abdominal segments. Body slender, widest in thorax, graduallytapering toward pygidium.
Prostomium wider than long, trilobed, with mid-superior lobe wider than lateral lobes, without eyes, nuchal and glandular ridges; anterior end of prostomium slightly upturned. Paleae absent.
Buccal tentacles conspicuous, densely covered with cilia on lateral and ventral sides (Fig. 65A B).
Segments 1 and 2 fused, concealed by branchial bases dorsally, forming large, somewhat inflated,finely crenulated lower lip ventrally. Segment 3 with first notopodia bearing fine, short setae andwith 3 pairs of branchiae, filiform, in nearly straight row, 1 pair slightly behind on either side;
branchial bases of left and right group fused each others (Fig. 65C). Segments 5 to 7 with gradually increasing notopodia, and parapodia in segment 8 (setiger 5, unciniger 2) extremely smaller
than others, with short setae.
Fully developed biramous parapodia from setiger 6; thoracic notopodia simple, cylindricalthroughout, neuropodia fairly long pinnules with very crowded and numerous uncini (Fig. 65D).
Abdominal neuropodia much narrower but longer than thoracic, with fewer uncini and digitate
dorsal cirri, shorter than pinnule; notopodial rudiments conical (Fig. 65E). Notosetae narrowly
limbate; thoracic and abdominal uncini similar, with single row of 5 7 teeth above small rostral
point and narrow basal prow (Fig. 65F J).
Pygidium with many cylindrical papillae (Fig. 65K).
The species is newly added to the Japanese polychaetous fauna.
Distribution. Sea of Okhotsk; Japan.
Samythella neglecta Wollebaek, 1912
(Fig. 66A L)
Samythella neglecta Wollebaek, 1912: 62 64, pl. 12, figs. 1 9.

Material. NSMT-Pol. 110282, WA05-FG410 (2), NSMT-Pol. 110283, WA05-FG510D (1).
Description. Largest specimen 32 mm long, 3 mm wide at thorax, consisting of 15 thoracic
setigers, 12 uncinigerous and 32 abdominal segments. Body linear, tapering slightly toward pygidium.
Prostomium approximately as wide as long; lateral sides tapering anteriorly, with truncatedanterior margin and two poorly marked longitudinal ridges; yellowly pigmented, crescent-shaped
nuchal organs along posterior margin of inferior lobe (Fig. 66A). Segments 1 and 2 fused, dorsallyconcealed by branchial bases, ventrally produced into slightly inflated, distinctly crenulated lower
lip. Paleae absent. Buccal tentacles conspicuous, slightly shorter than branchiae. Segment 3 with 3
pairs of branchiae in nearly straight row, 1 pair slightly behind on either side and bearing first small
notopodia with reduced setal fascicle; each branchia connected basally by membrane about aswide as branchial bases (Fig. 66B C).
First fully developed biramous parapodia in segment 6; notopodia simple, cylindrical throughout, uncinigers long, narrow pinnules (Fig. 66D). Abdominal neuropodia somewhat longer, with

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169

Fig. 65. Samythella bathycola Uschakov. —A, anterior end, lateral view, × 18; B, distal part of buccal tentacle, ×
153; C; anterior end, dorsal view, × 12; D, thoracic parapodium, anterior view, × 36; E, abdominal parapodium, same view, × 53; F G, thoracic uncini, lateral view, × 842; H J, abdominal uncini, lateral and frontal
views, × 842; K, pygidium, ventrolateral view, × 53.

tiny notopodial rudiments (Fig. 66E). Notosetae narrowly limbate (Fig. 66F); thoracic and abdominal uncini similar, with single row of 5 6 teeth above small rostral point and narrow basal
prow (Fig. 66G J).
Pygidium very small, with terminal anus surrounded by 10 conical papillae (Fig. 66K L).
The species is newly added to the Japanese polychaetous fauna.

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Fig. 66. Samythella neglecta Wollebaek. —A, prostomium, dorsal view, × 15; B, anterior end, same view, × 10; C;
same, lateral view, × 10; D, thoracic parapodium, anterior view, × 34; E, abdominal neuropodium, same view,
× 34; F, notoseta, × 160; G H, thoracic uncini, × 817; I J, abdominal uncini, × 817; K L, posterior end, lateral(K) and ventral (L) views, × 34.

Distribution. Norwegian Sea (depth: 225 1977 m); Japan.

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171

Family Trichobranchidae Malmgren, 1866
Genus Terebellides Sars, 1835
Terebellides horikoshii Imajima and Williams, 1985
Terebellides horikoshii Imajima and Williams, 1985: 15 16, fig. 4d f.

Material. NSMT-Pol. 110284, WA06-A1200D (1), NSMT-Pol. 110285, WA06-B750D (2),
NSMT-Pol. 110286, WA06-E1200D (1), NSMT-Pol. 110287, WA06-EF425D (1), NSMT-Pol.
110288, WA06-GH480D (7), NSMT-Pol. 110289, WA06-H1500D (3); NSMT-Pol. 110707,
WA07-A250D (1), NSMT-Pol. 110708, WA07-A450 (3), NSMT-Pol. 110709, WA07-A650 (4),
NSMT-Pol. 110710, WA07-C1500D (1).
Distribution. Off Kamchatka; Japan.
Terebellides intoshi Caullery, 1915
Terebellides intoshi Caullery, 1915: 111, figs. 1 2; Caullery, 1944: 186, fig. 149; Hessle, 1917: 142; Imajima and Williams,
1985: 9 10, fig. 2a c; Imajima, 1997a: 212.

Material. NSMT-Pol. 110291, WA05-E1000D (1), NSMT-Pol. 110292, WA05-GH510D (1).
Distribution. East Indies; Japan.
Terebellides japonica Moore, 1903
Terebellides stroemi var. japonica Moore, 1903: 478; Hessle, 1917: 142.
Terebellides japonica: Imajima and Williams, 1985: 10 11, fig. 2d f; Imajima, 1997a: 211.

Material. NSMT-Pol. 110293, WA05-GH510D (1).
Distribution. Japan.
Terebellides kobei Hessle, 1917
Terebellides kobei Hessle, 1917: 140, pl. 1, fig. 9, textfig. 32; Imajima and Hartman, 1964: 352; Imajima and Williams, 1985:
11 13, fig. 3a c; Imajima, 1997a: 211 212.

Material. NSMT-Pol. 110294, WA06-GH480D (11), NSMT-Pol. 110711, WA06-H480 (1).
Distribution. Japan.
Terebellides lineata Imajima and Williams, 1985
Terebellides lineata Imajima and Williams, 1985: 14 15, fig. 4a c; Imajima, 1997a: 212; Imajima, 2006: 391.

Material. NSMT-Pol. 110295, WA05-FG250D (7); NSMT-Pol. 110296, WA06-F650D (2),
NSMT-Pol. 110297, WA06-G900D (9), NSMT-Pol. 110298, WA06-G1200D (1), NSMT-Pol.
110299, WA06-H1500D (2); NSMT-Pol. 110712, WA07-A250D (1), NSMT-Pol. 110713, WA07D210D (2). NSMT-Pol. 110876, KT07-29-M-1 (1), NSMT-Pol. 110877, KT07-29-M-3-1 (1).
Distribution. Japan.

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Genus Trichobranchus Malmgren, 1865
Trichobranchus bibranchiatus Moore, 1903
Trichobranchus bibranchiatus Moore, 1903: 477 478, pl. 27, figs. 83 85; Hessle, 1917: 134; Imajima and Williams, 1985:
16 17, fig. 5a h.

Material. NSMT-Pol. 110300, WA06-GH480D (3).
Distribution. Japan.
Trichobranchus glacialis Malmgren, 1866
Trichobranchus glacialis Malmgren, 1866: 395, pl. 24, fig. 65; Fauvel, 1927: 288 289, fig. 100a h; Uschakov, 1955: 382, fig.
142E G.

Material. NSMT-Pol. 110928, WA06-E510D (1).
The species is newly added to the Japanese polychaetous fauna.
Distribution. Sea of Japan; Sea of Okhotsk; Bering Sea; Japan.
Family Terebellidae Malmgren, 1867
Genus Amphitrite Müller, 1771
Amphitrite cirrata Müller, 1771
Amphitrite cirrata: Moore, 1903: 473; Fauvel, 1927: 251 252, fig. 86i o; Okuda, 1937c: 58; Okuda, 1938a: 102; Imajima andHartman, 1964: 335; Day, 1967b: 746 747, fig. 36.9.m q; Imajima, 1997a: 212; Imajima, 2001a: 92.

Material. NSMT-Pol. 110255, WA05-FG480 (1), NSMT-Pol. 110256, WA05-G450 (1), NSMT-Pol. 110257, WA05-G750 (2); NSMT-Pol. 110258, WA06-EF425D (1), NSMT-Pol. 110259,
WA06-F510 (5), NSMT-Pol. 110260, WA06-F750 (4), NSMT-Pol. 110261, WA06-G900D (1),
NSMT-Pol. 110262, WA06-G1200D (5), NSMT-Pol. 110263, WA06-GH480D (8), NSMT-Pol.
110264, WA06-H1500D (1).
Distribution. North Atlantic; Bering Sea; Pacific of western Canada to central California;
Japan.
Amphitrite edwardsii (Quatrefages, 1865)
Amphitrite edwardsii: Fauvel, 1927: 245 246, fig. 84a i; Okuda, 1939: 240, textfig. 12; Imajima and Hartman, 1964: 335 336;
Imajima, 1997a: 212 213.

Material. NSMT-Pol. 110265, WA05-GH380 (1), NSMT-Pol. 110266, WA05-GH380D (1).
Distribution. Western Europe; Falkland Islands; Japan.
Amphitrite ramosissima Marenzeller, 1884
Amphitrite ramosissima Marenzeller, 1884: 200 201, pl. 1, fig. 2.
Amphitrite bifurcata Moore, 1903: 471 472, pl. 26, fig. 78.
Neoamphitrite ramosissima Hessle, 1917: 181.
? Amphitrite bifurcata Okuda, 1936: 155.

Material. NSMT-Pol. 110875, KT07-29-M-3-3 (2).
Distribution. Japan.

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Genus Artacama Malmgren, 1866
Artacama proboscidea Malmgren, 1866
Artacama proboscidea Malmgren, 1866: 394 395, pl. 23, fig. 60; Hessle, 1917: 194 195, pl. 2, fig. 13; Uschakov, 1955: 400,
fig. 152I L.

Material. NSMT-Pol. 110267, WA05-DE380D (1); NSMT-Pol. 110268, WA06-B750D (1),
NSMT-Pol. 110269, WA06-F650D (1), NSMT-Pol. 110270, WA06-FG350D (1), NSMT-Pol.
110271, WA06-GH480D (7).
The species is newly added to the Japanese polychaetous fauna.
Distribution. Seas of Japan and Okhotsk; Japan.
Genus Eupolymnia Verrill, 1900
Eupolymnia congruens (Marenzeller, 1884)
Polymnia congruens Marenzeller, 1884: 207 208, pl. 2, fig. 3.
Polymnia nesidensis japonica Moore, 1903: 475.
Eupolymnia congruens: Imajima and Hartman, 1964: 337 338.

Material. NSMT-Pol. 109828, WA06-A1200D (1), NSMT-Pol. 109829, WA06-D550 (1),
NSMT-Pol. 109830, WA06-EF425D (10), NSMT-Pol. 109831, WA06-F380 (1), NSMT-Pol.
109832, WA06-F425 (1), NSMT-Pol. 109833, WA06-F550 (1), NSMT-Pol. 110922, WA06-F1500(2), NSMT-Pol. 110923, WA06-FG425 (3), NSMT-Pol. 110924, WA06-FG480 (8), NSMT-Pol.
110925, WA06-G550 (1), NSMT-Pol. 110926, WA06-G900D (1), NSMT-Pol. 110927, WA06GH480D (1).
Distribution. Alaska; California; Japan.
Genus Nicolea Malmgren, 1866
Nicolea zostericola (Oersted, 1844)
Nicolea zostericola: Malmgren, 1865: 381, pl. 26, fig. 76; Fauvel, 1927: 261 262, fig. 90g n; Uschakov, 1950: 223; Uschakov, 1965: 366, fig. 146B D.

Material. NSMT-Pol. 110929, WA05-DE380D (42), NSMT-Pol. 110930, WA05-EF450D(3).
The species is newly added to the Japanese polychaetous fauna.
Distribution. Seas of Japan and Okhotsk; Bering Sea.
Genus Pista Malmgren, 1866
Pista cristata (Müller, 1776)
Pista cristata: Malmgren, 1866: 382, pl. 22, fig. 59; Okuda, 1938a: 102; Uschakov, 1965: 364, fig. 144A C.

Material. NSMT-Pol. 110273, WA06-EF425D (4), NSMT-Pol. 110274, WA06-GH480D(45).
Distribution. Atlantic; Seas of Japan and Okhotsk; Japan.
Pista macrolobata Hessle, 1917
Pista macrolobata Hessle, 1917: 157, pl. 2, fig. 4.

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Material. NSMT-Pol. 110276, WA05-DE380D (1), NSMT-Pol. 110277, WA05-GH380D (1).
Distribution. Japan (Ogasawara Islands).
Genus Scionella Moore, 1903
Scionella vinogradovi (Uschakov, 1955)
Pista vinogradovi Uschakov, 1955: 386, fig. 143D F.
Scionella vinogradovi: Hartman, 1965: 74; Banse and Hobson, 1968: 46; Saphronova, 1991: 241 243, figs. 1 3.

Material. NSMT-Pol. 110278, WA05-DE380D (1), NSMT-Pol. 110279, WA05-FG510D (1);
NSMT-Pol. 110280, WA06-D450D (1), NSMT-Pol. 110281, WA06-GH480D (5).
The species is newly added to the Japanese polychaetous fauna.
Distribution. Sea of Okhotsk; Bering Sea; Japan.
Genus Streblosoma Sars, 1872
Streblosoma bairdi (Malmgren, 1866)
Grymaea bairdi Malmgren, 1866: 388, pl. 19, fig. 69.
Streblosoma Bairdi: Hessle, 1917: 211; Uschakov, 1965: 378, fig. 152A E.

Material. NSMT-Pol. 110784, WA05-E1000D (1), NSMT-Pol. 110785, WA05-G1500D (2);
NSMT-Pol. 110786, WA06-E1200D (1), NSMT-Pol. 110787, WA06-H1500D (1).
The species is newly added to the Japanese polychaetous fauna.
Distribution. Atlantic; Seas of Japan and Okhotsk; Japan.
Order Sabellida
Family Sabellidae Malmgren, 1867
Genus Chone Kröyer, 1856
Chone gracilis Moore, 1906
Chone gracilis Moore, 1906: 257 259, pl. 12, figs. 62 66; Banse, 1972: 470 472, fig. 4.

Material. NSMT-Pol. 110677, WA05-FG510D (3), NSMT-Pol. 110678, WA05-G280 (15),NSMT-Pol. 110679, WA05-GH510D (6); NSMT-Pol. 110684, WA06-E510D (1), NSMT-Pol.
110685, WA06-F650D (6).
The species is newly added to the Japanese polychaetous fauna.
Distribution. Alaska; Japan.
Chone magna (Moore, 1923)
Euchone magna Moore, 1923: 245 246, pl. 18, figs. 45 46.
Chone magna: Hartman, 1969: 669; Banse, 1972: 472 473, fig. 5.

Material. NSMT-Pol. 110675, WA05-EF250D (15); NSMT-Pol. 110686, WA06-E750 (7),
NSMT-Pol. 110687, WA06-F900 (7), NSMT-Pol. 110688, WA06-F1500 (12).
The species is newly added to the Japanese polychaetous fauna.
Distribution. California; Japan.

Deep-sea Benthic Polychaetes off Northern Honshu

175

Chone sp.
Material. NSMT-Pol. 110878, KT07-29-M-2 (1), NSMT-Pol. 110879, KT07-29-M-3-1 (1),
NSMT-Pol. 110880, KT07-29-M-3-2 (14), NSMT-Pol. 110881, KT07-29-M-3-3 (3).
Genus Demonax Kinberg, 1867
Demonax aulaconota (Marenzeller, 1884)
Sabella aulaconota Marenzeller, 1884: 210, pl. 2, fig. 8; Johansson, 1927: 124 125; Fauvel, 1936a: 84.
Demonax aulaconota: Imajima and Hartman, 1964: 356.

Material. NSMT-Pol. 110689, WA05-E1000D (8), NSMT-Pol. 110690, WA05-F510 (1),
NSMT-Pol. 110691, WA05-FG425 (1), NSMT-Pol. 110692, WA05-G450 (1), NSMT-Pol. 110693,
WA05-G750 (28), NSMT-Pol. 110694, WA05-G1500 (1), NSMT-Pol. 110695, WA05-GH380D(1); NSMT-Pol. 110696, WA06-F510 (1), NSMT-Pol. 110697, WA06-F1500D II (3), NSMT-Pol.
110698, WA06-G480 (1), NSMT-Pol. 110699, WA06-G750 (13), NSMT-Pol. 110700, WA06G900D (92), NSMT-Pol. 110701, WA06-H1500D (74).
Distribution. Japan.
Genus Euchone Malmgren, 1866
Euchone analis (Kröyer, 1856)
Sabella analis Kröyer, 1856: 17.
Euchone analis: Malmgren, 1866: 406; 1867: 223; Berkeley and Berkeley, 1952: 121 122, figs. 250 251; Pettibone, 1954:
339; Uschakov, 1965: 395; Hartmann- Schröder, 1971: 521; Banse, 1972: 482 483, fig. 9a-c.

Material. NSMT-Pol. 110680, WA05-EF250D (1), NSMT-Pol. 110681, WA05-F1200 (2),
NSMT-Pol. 110682, WA05-FG410 (4), NSMT-Pol. 110683, WA05-FG450 (2).
The species is newly added to the Japanese polychaetous fauna.
Distribution. North Atlantic; Alaska; Sea of Okhotsk; Japan.
Family Serpulidae Savigny, 1818
Genus Crucigera Benedict, 1887
Crucigera zygophora (Johnson, 1901)
Crucigera zygophora Johnson, 1901: 433 434, pl. 19, figs. 205 208; Bush, 1904: 233, pl. 29, fig. 5, pl. 31, fig. 2, pl. 33, fig.
3, pl. 39, figs. 8 13, 15, 17, 20; Imajima, 1988: 128; Imajima, 1996: 305, fig. 243.
Serpula (Crucigera) zygophora: Uschakov, 1965: 400, fig. 160F.

Material. NSMT-Pol. 110676, WA07-A250 (1).
Distribution. Sea of Okhotsk; Bering Sea; Alaska; California; Japan.
Summary of Result
A total of 243 species including 32 indeterminable species, belonging to 43 families, wereobtained by this survey. The following breakdown illustrates the degree of their endemism, andother facts of their distributional character:14 species are newly described.
37 species are newly added to the Japanese polychaetous fauna.

176

Minoru Imajima

Table 2. Species of polychaetes, with the degree of their endemism and other facts of their distributional characters.

FamilyAphroditidae

Polynoidae

Acoetidae
Pholoidae
Sigalionidae

Phyllodocidae

Glyceridae

Goniadidae

Sphaerodoridae

Species

Aphrodita aculeata
A. goolmarris
A. negligens
A. nipponensis
A. sibogae
A. talpa
Laetmonice japonica
L. producta
Laetmonice sp.
Bylgides macrolepida
Eunoe barbata
E. depressa
E. shirikishinai
Gattyana ciliata
Harmothoe aspera
H. extenuata
H. grandispina
H. imbricata
H. imparLepidasthenia interrupta
L. izukai
L. magnacornuta
Lepidonotus glaber
Parahalosydna krassini
Weberia abyssicola
Acoetes jogasimae
Pholoe sp.
Labioleanira sp.
Leanira sp.
Neoleanira areolata
Sigalion orientalis
Eulalia bilineata
Eulalia sp.
Mysta sp.
Paranaitis polynoides
Phyllodoce groenlandica
P. lineata tosaensis
P. madeirensis
Glycera branchiopoda
G. capitata
G. neorobusta
G. okaiG. semibranchiopoda
G. tesselata
Glycera sp.
Glycinde sp.
Goniada brunnea
G. foliacea
G. maculata
G. vorax
Clavodorum sp.
Ephesiella brevicapitis

New species

New toJapan

Also innorthwestern Pacificareas

Endemic to
Japan

×
×
×
×
×

×
×
×
×

×

×

×
×
×
×
×
×
×
×
×

×

×

×

×
×

×
×
×
×

×
×

×
×
×
×

×

×
×
×
×

Collected
depth (m)898 1499
377 904
146 357
305 904
213 1214277 509299 3960
146 5215219 5268
294 556
209 551276 758
153 156
253 497509 510508 900
276 383
409 415383 1737
380 512373 454410 427294 318420 424
1709 1737
248 248249 285
816 8204094 4128
210 1005
146 214747 750478 483
478 483
354 355
213 1505
213 750
210 1470
346 750
213 355
1512 1513
284 1402
253 904
153 905
816 8202948 2991
248 1505
248 1498
213 512515 516454 3032
284 4181

177

Deep-sea Benthic Polychaetes off Northern Honshu

Table 2. (Continued)

FamilyHesionidae
Pilargidae

Syllidae

Nereididae

Nephtyidae

Paralacydoniidae
Euphrosinidae
Onuphidae

Eunicidae
Lumbrineridae

Species

Sphaerodoropsis biserialis
S. minuta
Sphaerodorum gracilis
Podarkeopsis glabra
Ancistrosyllis groenlandica
Cabira sp.
Sigambra bassi
S. bidentata
Sigambra spp.
Autolytus sp.
Eusyllis blomstrandi
Exogone breviantennata
Exogone sp.
Sphaerosyllis erinaceus
Sphaerosyllis sp.
Syllis spongiphila
Typosyllis alternata
Typosyllis spp.
Ceratocephala borealis
Nereis abyssa
Rullierinereis profunda
Aglaophamus japonicus
A. malmgreni
A. sinensis
Aglaophamus spp.
Nephtys caeca
N. longosetosa
N. oligobranchia
N. paradoxa
N. punctata
Paralacydonia paradoxa
Euphrosine digitalis
E. pseudonotialis
Anchinothria macrobranchiata
Epidiopatra rugosa
Nothria grossa
Onuphis geophiliformis
O. holobranchiata
O. imajimai
O. opalina
O. taraba
Paradiopatra crassa
P. gracilis
P. striata
Eunice mucronata
Augeneria bidens
Eranno abyssicola
E. bifurcata
E. tosaensis
Lumbrineris inflata
L. japonica
L. latreilli

New species

New toJapan

Also innorthwestern Pacificareas

Endemic to
Japan

×
×
×

×

×

×
×
×
×
×
×

×
×
×
×
×
×
×
×
×
×
×
×
×
×
×
×
×
×
×
×
×

Collected
depth (m)284 1737454 497
213 925
253 255
508 1514
2032 2055
253 2551450 1498
816 4181375
213 483375 4742018454 4972018
376 427284 925
249 416
248 1498
151 14992018
253 255
248 3032
146 147
213 1733
213 213213 893454 1603
248 1728
213 2055
253 2055
420 454420 4282948 2991
375 1515
154 1498
248 1470
213 1470
478 1505
213 1515
248 1471
213 213213 1511
146 1499
146 1214
1498 4951
213 4128213 1202
249 483
416 820454 2055
213 1728

178

Minoru Imajima

Table 2. (Continued)

FamilyOenonidae

Orbiniidae

Paraonidae

Poecilochaetidae
Spionidae

Chaetopteridae
Cirratulidae

Cossuridae
Flabelligeridae

Acrocirridae
Fauveliopsidae
Sternaspidae
Capitellidae

SpeciesNinoe palmata
Paraninoe simpla
Scoletoma longifolia
S. nipponica
Drilonereis falcata japonica
D. filum
Notocirrus japonicus
Califia calida
Leitoscoloplos pugettensis
Naineris japonica
Phylo felix
P. fimbriatus
P. nudus
Scoloplos (Leodamas) robustus
Scoloplos (Scoloplos) similis
Aricidea (Acmira) simplex
A. (Aedicira) belgicae
A. (Allia) antennata
Aricidea (Allia) sp.
Cirrophorus branchiatus
Levinsenia gracilis
Paradoneis lyra
Poecilochaetus granulatus
Laonice cirrata
Prionospio (Prionospio) bocki
P. (P.) depauperata
P. (P.) dubia
Prionospio sp.
Spiophanes japonicum
S. kroyeri
Spiochaetopterus typicus
Aphelochaeta sp.
Chaetozone setosa
C. spinosa
Chaetozone sp.
Cirratulus cirratus
Monticellina tesselata
Timarete sp.
Cossura spp.
Brada granulata
B. ochotensis
B. sachalina
B. villosa
Pherusa plumosa
Acrocirrus heterochaetus
Acrocirrus sp.
Fauveliopsis glabra
Laubieriopsis brevis japonica
Sternaspis scutata
Capitella capitata
Leiochrides sp.
Notomastus hemipodus

New species

New toJapan

Also innorthwestern Pacificareas

Endemic to
Japan

×
×
×
×
×

×
×
×
×
×
×
×

×
×
×

×
×
×

×
×
×

×

×

×
×
×

Collected
depth (m)253 4181641 2055
454 4128416 1499
253 1499
253 259
146 1201
508 512284 285354 424354 483
373 512416 1202
373 378
253 503375 483
284 1470
478 512454 4971695 1733
478 1214375 512478 483
253 1402
375 503249 285
213 483
284 497248 248
248 1470
213 1514
248 1498
248 497
248 483
253 285
146 1728
375 503
1466 1471
213 1201
294 318416 416305 512354 1511
303 1470
354 4201709 1737
454 3032
1498 4181
213 1499
276 521
1498
1498

179

Deep-sea Benthic Polychaetes off Northern Honshu

Table 2. (Continued)

FamilyMaldanidae

Opheliidae

Scalibregmatidae

Oweniidae

Pectinariidae
Sabellariidae
Ampharetidae

SpeciesN. latericeus
Pseudoleiocapitella sp.
Clymenura (Cephalata) aciculata
C. (C.) columbiana
C. (C.) lankesteri
C. (C.) longicaudata
Clymenura (Cephalata) sp.
Clymenura (Clymenura) japonica
Clymenella koellikeri
Euclymene uncinata
Isocirrus planiceps
Maldanella harai
Maldanella sp.
Microclymene caudata
Praxillella gracilis
P. pacifica
P. praetermissa
Clymenopsis cingulata
Lumbriclymene japonica
Notoproctus pacificus
Praxillura tanseiana
Asychis auritus
Chirimia biceps
Maldane cristata
M. pigmentata
Maldane sp.
Metasychis disparidentata
M. gotoiNicomache (Nicomache) lumbricalis
Nicomache (Nicomache) sp.
Rhodine loveni
Ophelina acuminata
O. arctica
O. breviata
Travisia brevis
T. pupa
Asclerocheilus beringianus
Pseudoscalibregma orientalis
Scalibregma inflatum
Galathowenia oculata
G. scotiae
Myriochele heeri
M. olgae
Myriochele sp.
Owenia spp.
Amphictene japonica
Cistenides hyperborea
Lygdamis giardi
Amage auricula
A. delus
Ampharete longipaleolata

New species

New toJapan

Also innorthwestern Pacificareas

Endemic to
Japan

×
×
×
×
×
×
×
×
×

×
×
×
×

×

×

×

×
×
×
×
×

×
×
×
×

×
×
×
×
×

×
×
×
×
×

×

Collected
depth (m)253 2055
641 750
249 454
253 512420 1514
1511 1514
1511 2991
249 1005
508 512478 1005
146 662
253 904
2948 2991
248 424
213 1514
248 2055
213 1728
248 1515
253 2991
253 1499
248 904
508 1005
373 925
213 2055
248 259816 820647 883
248 553
213 1521
2018 4951
213 1514
213 1498
253 2055
284 503253 1514
508 749
1213 1214
373 1005
249 3032
213 375
249 1471
213 925
249 1005
2032 2055
146 497
248 1733
258 258
422 425375
661 4128375 1402

180

Minoru Imajima

Table 2. (Continued)

FamilyTrichobranchidae

Terebellidae

Sabellidae

Serpulidae

Species

Amphicteis mederi
A. scaphobranchiata
Amphisamytha japonica
Anobothrus gracilis
A. wakatakamaruae
Auchenoplax crinita
Lysippe labiata
Melinna cristata
M. elisabethae
Melinnopsis atlantica
Paramage scutata
Samythella bathycola
S. neglecta
Terebellides horikoshii
T. intoshi
T. japonica
T. kobei
T. lineata
Trichobranchus bibranchiatus
T. glacialis
Amphitrite cirrata
A. edwardsii
A. ramosissima
Artacama proboscidea
Eupolymnia congruens
Nicolea zostericola
Pista cristata
P. macrolobata
Scionella vinogradovi
Streblosoma bairdi
Chone gracilis
C. magna
Chone sp.
Demonax aulaconota
Euchone analis
Crucigera zygophora

New species

New toJapan

Also innorthwestern Pacific
areas

×
×

×
×

×

×

×

×
×
×

Endemic to
Japan

×
×

×
×
×
×

×
×
×
×
×
×

×

×
×

×

×
×
×
×

×

×

×
×
×
×
×

×
×
×

×

×
×

×

Collected
depth (m)146 925420 1470
248 248
213 4181452 516
248 925
248 503478 1402
213 2851450 4128
346 905
446 450410 516258 1499
508 1005
508 512478 483
213 1728
478 483
498 503420 1470
373 3811695 1733
346 750379 1515
375 454420 483
373 378375 516
1004 1498
277 647
253 1515
1528 1737
373 1498
253 1196
258 273

40 species are known only from Japan, or endemic.
86 species are known also from the northwestern Pacific Ocean, including Sea of Okhotsk
and Bering Sea.
These data are analyzed in the Table 2.
Of these samples, the species-rich families were Maldanidae (27 spp.) Ampharetidae (16spp.), Polynoidae (16 spp.), Onuphidae (11 spp.) and Lumbrineridae (11 spp.). Eleven families
were represented by single species.
The most species were collected from the bathyal zone (400 2000 m). However, the following species are extended to the abyssal zone (2000 6000 m):
Laetmonice japonica
Laubieriopsis brevis japonica
Laetmonice sp.
Notomastus latericeus

Deep-sea Benthic Polychaetes off Northern Honshu

Leanira sp.
Glycinde sp.
Clavodorum sp.
Ephesiella brevicapitis
Cabira sp.
Aglaophamus malmgreni
Nephtys punctata
Paralacydonia paradoxa
Anchinothria macrobranchia
Augeneria bidens
Eranno abyssicola
Fauveliopsis glabra

181

Clymenura (Cephalata) sp.
Maldanella sp.
Praxillella pacifica
Maldane cristata
Nicomache (Nicomache) sp.
Ophelina arctica
Scalibregma inflatum
Myriochele sp.
Amage delus
Anobothrus gracilis
Melinnopsis atlantica

Acknowledgements
This study would not have been possible without the cooperation of the staff members of the
Department of Zoology of the National Museum of Nature and Science, Tokyo. Thanks are due tothe captain and crew of the R/Vs Wakataka-maru, Soyo-maru and Tansei-maru. I also thank to Dr.
Hannelore Paxton, Macquarie University, Australia, for giving me the personal communicationand the criticism of the manuscript on Paradiopatra gracilis. Thanks are extended to Dr. Angelika
Brandt, Zoologisches Institut und Zoologisches Museum, Hamburg, Germany for the loan of thetype material of Scalibregma chilensis, Ms. Emma Sherlock, The Natural History Museum, London for the loan of the type material of Melinnopsis atlantica and Dr. Jon Anders Kongsrud, University of Bergen-Museum of Zoology, Norway for the loan of the type material of Pseudoscalibregma parvum.
References
Annenkova, N. P., 1922. Aperçu de la famille des Chloraemidae (Annelida Polychaeta) de la collection du Musée Zoologiquede l’Académie des Sciences de Russie. (A Survey of the Family Chloraemidae (Annelida, Polychaeta) from the collection of the Zoological Museum of the Russian Academy of Sciences). Doklady Akademii Nauk SSSR: 38 40.
Annenkova, N. P., 1929. Beiträge zur Kenntnis der Polychaeten-Fauna der USSR. I. Fam. Pectinariidae Quatrefages (Amphictenidae Malmgren) und Ampharetidae Malmgren. Annuaire du Musée Zoologique de l’Académie des Sciences del’URSS, 30: 477 503, pls. 37 39.
Annenkova, N. P., 1934a. Paraoniden der Meeren des Fernen Ostens der USSR. Doklady Akademii Nauk SSSR, 3: 656 661.
(In Russian with German summary.)
Annenkova, N. P., 1934b. Kurze übersicht der Polychaeten der Litoralzone der Bering-Insel (Kommandor-Inseln), nebst
Beschreibung neuer Arten. Zoologischer Anzeiger, 106: 322 331.
Annenkova, N. P., 1937. The Polychaete fauna of the northern part of the Sea of Japan. Explorations des Mers de l’URSS,(23): 139 216, pls.1 5. (In Russian with brief English translation.)
Annenkova, N. P., 1952. Novye vidy mnogoschchetinkovykh chervei (Polychaeta) (new polychaete species). Trudy Zoologicheskogo Instituta, Akademii Nauk SSSR, 12: 148 154.
Arwidsson, I., 1907. Studien über die skandinavischen und arktischen Maldaniden nebst Zusammenstellung der übringenbisher bekannten Arten dieser Familie. Zoologische Jahrbücher. Abteilung für Systematik, Geographie und Biologie,
Der Tiere. Supplement 9: 1 308, pls. 1 12.
Ashworth, J. H., 1901. The anatomy of Scalibregma inflatum Rathke. Quarterly Journal of Microscopical Science, 45:
237 309.
Augener, H., 1927. Papers from Dr. T. H. Mortensen’s Pacific Expedition 1914 16. No. 38. Polychaeten von Südost- undSüd-Australien. Videnskabelige Meddelelser fra Dansk naturhistorisk Førening i Kjøbenhavn, 83: 71 275.
Banse, K., 1972. Redescription of some species of Chone Kröyer and Euchone Malmgren, and three new species (Sabellidae,
Polychaeta). Fishery Bulletin, 70: 459 495.
Banse, K., 1979. Ampharetidae (Polychaeta) from British Columbia and Washington. Canadian Journal of Zoology, 57:

182

Minoru Imajima

1543 1552.
Banse, K. and K. D. Hobson, 1968. Benthic polychaetes from Puget Sound, Washington, with remarks on four other species.
Proceedings of the United States National Museum, 125: 1 53.
Banse, K. and K. D. Hobson, 1974. Benthic errantiate polychaetes of British Columbia and Washington. Bulletin of the Fisheries Research Board of Canada, 185: 1 111.
Barnich, R. and D. Fiege, 2000. Review of the North East Atlantic and Mediterranean species of Aphrodita Linnaeus,
1758 and Aphroditella Roule, 1898 (Polychaeta: Aphroditidae). Ophelia, 753: 131 140.
Bergström, E., 1914. Zur Systematik der Polychaetenfamilie der Phyllodociden. Zoologiska Bidrag från Uppsala, 3:
37 224.
Berkeley, C., 1966. Records of some species of Polychaeta new to British Columbia and of extensions in distribution of someothers. Canadian Journal of Zoology, 44: 839 849.
Berkeley, E., 1929. Polychaetous annelids from the Nanaimo district. 4. Chaetopteridae and Maldanidae. Contributions to
Canadian Biology, New Series, 4: 305 316, 1 pl.
Berkeley, E. and C. Berkeley, 1942. North Pacific Polychaeta, chiefly from the west coast of Vancouver Island, Alaska andBering Sea. Canadian Journal of Research, 20: 183 208.
Berkeley, E. and C. Berkeley, 1944. Polychaeta from the western Canadian Arctic region. Canadian Journal of Research, 22:
1 5.
Berkeley, E. and C. Berkeley, 1948. Annelida. Polychaeta Errantia. Canadian Pacific Fauna. Fisheries Research Board of
Canada, (9b-1): 1 100.
Berkeley, E. and C. Berkeley, 1952. Annelida. Polychaeta Sedentaria. Canadian Pacific Fauna. Fisheries Research Board of
Canada, (9b-2): 1 139.
Berkeley, E. and C. Berkeley, 1956. On a collection of polychaetous annelids from northern Banks Island, from the south
Beaufort Sea, and from northwest Alaska; together with some new records from the east of Canada. Journal of the Fisheries Research Board of Canada, 13: 233 246.
Blake, J. A., 1984. Polychaeta Oweniidae from Antarctic seas collected by the United States Antarctic Research Program.
Proceedings of the First International Polychaete Conference, Sydney, Australia, July 1983: 112 117.
Blake, J. A., 1994a. 4. Family Phyllodocidae Savigny, 1818. In: Blake, J. A. and B. Hilbig (eds.), Taxonomic Atlas of thebenthic fauna of the Santa Maria Basin and Western Santa Barbara Channel, Volume 4. The Annelida Part 1. Oligochaeta and Polychaeta: Phyllodocida (Phyllodocidae to Paralacydoniidae), pp. 115 186. Santa Barbara Museum of
Natural History, Santa Barbara, California.
Blake, J. A., 1994b. 4. Family Pilargidae Saint Joseph, 1899. In: Blake, J. A. and B. Hilbig (eds.), Taxonomic Atlas of thebenthic fauna of the Santa Maria Basin and Western Santa Barbara Channel, Volume 4. The Annelida Part 1. Oligochaeta and Polychaeta: Phyllodocida (Phyllodocidae to Paralacydoniidae), pp. 271 294. Santa Barbara Museum of
Natural History, Santa Barbara, California.
Blake, J. A., 1994c. 14. Family Paralacydoniidae Pettibone, 1963. In: Blake, J. A. and B. Hilbig (eds.), Taxonomic Atlas of
the benthic fauna of the Santa Maria Basin and Western Santa Barbara Channel, Volume 4. The Annelida Part 1. Oligochaeta and Polychaeta: Phyllodocida (Phyllodocidae to Paralacydoniidae), pp. 363 367. Santa Barbara Museum of
Natural History, Santa Barbara, California.
Blake, J. A., 1996a. 1. Family Orbiniidae Hartman, 1942. In: Blake, J. A., B. Hilbig and P. H. Scott (eds.), Taxonomic Atlasof the benthic fauna of the Santa Maria Basin and Western Santa Barbara Channel, Volume 6. The Annelida Part 3.
Polychaeta: Orbiniidae to Cossuridae, pp. 1 26. Santa Barbara Museum of Natural History, Santa Barbara, California.
Blake, J. A., 1996b. 2. Family Paraonidae Cerruti, 1909. In: Blake, J. A., B. Hilbig and P. H. Scott (eds.), Taxonomic Atlas of
the benthic fauna of the Santa Maria Basin and Western Santa Barbara Channel, Volume 6. The Annelida Part 3. Polychaeta: Orbiniidae to Cossuridae, pp. 27 70. Santa Barbara Museum of Natural History, Santa Barbara, California.
Blake, J. A., 1996c. 2. Family Cirratulidae Ryckholdt, 1851. In: Blake, J. A., B. Hilbig and P. H. Scott (eds.), Taxonomic Atlasof the benthic fauna of the Santa Maria Basin and Western Santa Barbara Channel, Volume 6. The Annelida Part 3.
Polychaeta: Orbiniidae to Cossuridae, pp. 263 384. Santa Barbara Museum of Natural History, Santa Barbara, California.
Blake, J. A., 2000a. 1. Family Flabelligeridae Saint Joseph, 1894. In: Blake, J. A., B. Hilbig and P. V. Scott (eds.), Taxonomic Atlas of the benthic fauna of the Santa Maria Basin and Western Santa Barbara Channel, Volume 7. The AnnelidaPart 4. Polychaeta: Flabelligeridae to Sternaspidae, pp. 1 23. Santa Barbara Museum of Natural History, Santa Barbara, California.
Blake, J. A., 2000b. 4. Family Capitellidae Grube, 1862. In: Blake, J. A., B. Hilbig and P. V. Scott (eds.), Taxonomic Atlas of
the benthic fauna of the Santa Maria Basin and Western Santa Barbara Channel, Volume 7. The Annelida Part 4. Polychaeta: Flabelligeridae to Sternaspidae, pp. 47 96. Santa Barbara Museum of Natural History, Santa Barbara, California.
Blake, J. A., 2000c. 7. Family Opheliidae Malmgren, 1867. In: Blake, J. A., B. Hilbig and P. V. Scott (eds.), Taxonomic Atlasof the benthic fauna of the Santa Maria Basin and Western Santa Barbara Channel, Volume 7. The Annelida Part 4.

Deep-sea Benthic Polychaetes off Northern Honshu

183

Polychaeta: Flabelligeridae to Sternaspidae, pp. 145 168. Santa Barbara Museum of Natural History, Santa Barbara,
California.
Blake, J. A., 2000d. 6. Family Scalibregmatidae Malmgren, 1867. In: Blake, J. A., B. Hilbig and P. V. Scott (eds.), Taxonomic Atlas of the benthic fauna of the Santa Maria Basin and Western Santa Barbara Channel, Volume 7. The AnnelidaPart 4. Polychaeta: Flabelligeridae to Sternaspidae, pp. 129 144. Santa Barbara Museum of Natural History, Santa
Barbara, California.
Blake, J. A., 2000e. 5. Family Oweniidae Rioja, 1917. In: Blake, J. A., B. Hilbig and P. V. Scott (eds.), Taxonomic Atlas of thebenthic fauna of the Santa Maria Basin and Western Santa Barbara Channel, Volume 7. The Annelida Part 4. Polychaeta: Flabelligeridae to Sternaspidae, pp. 97 128. Santa Barbara Museum of Natural History, Santa Barbara, California.
Blake, J. A. and D. Dean, 1973. Polychaetous annelids collected by the R/V Hero from Baffin Island, Davis Strait and West
Greenland in 1968. Bulletin of the Southern California Academy of Sciences, 72: 31 39.
Blake, J. A. and M. E. Petersen, 2000. 3. Family Fauveliopsidae Hartman, 1971. In: Blake, J. A., B. Hilbig and P. V. Scott(eds.), Taxonomic Atlas of the benthic fauna of the Santa Maria Basin and Western Santa Barbara Channel, Volume 7.
The Annelida Part 4. Polychaeta: Flabelligeridae to Sternaspidae, pp. 31 45. Santa Barbara Museum of Natural History, Santa Barbara, California.
Böggemann, M., 1998. Polychaeten aus der Deutschen Bucht. Courier Forschungsinstitut Senckenberg, 202: 1 315. [erroneously dated 1997]
Böggemann, M., 2002. Revision of the Glyceridae Grube, 1850 (Annelida: Polychaeta). Abhandlungen der Senckenbergischen Naturforschenden Gesellschaft, 555: 1 249.
Böggemann, M., 2005. Revision of the Goniadidae (Annelida, Polychaeta). Abhandlungen des Naturwissenschaftlichen Vereins in Hamburg, (29): 1 354.
Böggemann, M. and D. Fiege, 2001. Description of seven new species of the genus Glycera Savigny, 1818 (Annelida: Polychaeta: Glyceridae). Ophelia, 54: 29 49.
Britaev, T. A. and M. A. Saphronova, 1981. New species of the family Pilargidae (Polychaeta) from the Sea of Japan andrevision of the genus Cabira. Zoologishetskij Zhurnal, Akademii Nauk SSSR, 60: 1314 1324. (In Russian.)Bush, K. J., 1904.Tubicolous Annelids of the Tribes Sabellides and Serpulides from the Pacific Ocean. Harriman Alaska
Expedition, 12: 169 355, pls. 21 24.
Cantone, G., 1982. Primo rinvenimento in Mediterraneo di Rullierinereis Pettibone, 1971 (Policheti Nereidi) con descrizione
di una nuova specie. Animalia, 9: 103 107.
Caullery, M., 1915. Sur les Terebellides Malmgren du Siboga et les Terébélliens voisins. Bulletin de la Société Zoologique deFrance, 40: 111 116.
Caullery, M., 1944. Polychètes sédentaires de l’Expédition du Siboga: Ariciidae, Spionidae, Chaetopteridae, Chloraemidae,
Opheliidae, Oweniidae, Sabellariidae, Sternaspidae, Amphictenidae, Ampharetidae, Terebellidae. Siboga-Expedition,
Leiden, 24 (2 bis): 1 204.
Chamberlin, R.V., 1919. The Annelida Polychaeta. Memoirs of the Museum of Comparative Zoology, 48: 1 514, pls. 1 80.
Chambers, S. J. and A. I. Muir, 1997. Polychaetes: British Chrysopetaloidea, Pisionoidea and Aphroditoidea. 202 pp. The
Linnean Society of London and The Estuarine and Coastal Sciences Association by Field Studies Council, Oegstgeest.
Claparède, E.,1863. Beobachtungen über Anatomie und Entwicklungsgeschichte wirbelloser Thiere an der Küste von Normandie angestellt. 120 pp., 18 pls. W. Engelmann, Leipzig.
Claparède, E., 1868. Les Annélides Chétopodes du Golfe de Naples. Mémoires de la Société de Physique et d’Histoire Naturelle de Genève, 19: 313 584, 16 pls.
Cuvier, G. L. C. F. D., 1830. Le Règne Animal. Distribué d’après son organisation pour servir de base à l’Histoire Naturelledes Animaux et d’introduction à l’anatomie comparée. Nouvelle édition, revue et augmentée, Volume 3. Déterville andCrochard, Paris.
Day, J. H., 1961. The polychaete fauna of South Africa. Part. 6: Sedentary species dredged off Cape coasts with a few newrecords from the shore. Journal of the Linnean Society of London, 44: 463 560.
Day, J. H., 1963a. The Polychaete fauna of South Africa. Part 7: Species from depths between 1,000 and 3,300 metres westof Cape Town. Annals of the South African Museum, 46: 353 371.
Day, J. H., 1963b. The Polychaete fauna of South Africa. Part 8: New species and records from grab samples and dredgings.
Bulletin of the British Museum (Natural History), Zoology, 10: 383 445.
Day, J. H., 1967a. A monograph on the Polychaeta of Southern Africa. Part 1, Errantia. 458 pp. + xxix. Trustees of the British
Museum (Natural History), London.
Day, J. H., 1967b. A monograph on the Polychaeta of Southern Africa. Part 2, Sedentaria. pp. 459 878+xvii. Trustees of the
British Museum (Natural History), London.
Day, J. H., 1973. New Polychaeta from Beaufort, with a key to all species recorded from North Carolina. NOAA Technical
Report, NMFS Circular, 375: 1 140.
Ditlevsen, H., 1917. Annels. I. Danish Ingolf-Expedition, 4: 1 71.

184

Minoru Imajima

Ehlers, E., 1868. Die Borstenwürmer nach systematischen und anatomischen Untersuchungen dergestellt. pp. 269 748, pls.
12 24. Wilhelm Engelmann, Leipzig.
Ehlers, E., 1887. Report on the annelids of the dredging expedition of the U.S. Coast Survey Steamer “Blake”. Memoirs of
the Museum of Comparative Zoology, 15: 1 335, 60 pls.
Ehlers, E., 1900. Magellanische Anneliden gesammelt während der schwedischen Expedition nach den Magellansländern.
Nachrichten von der Königlichen Gesellschaft der Wissenschaften zu Göttingen, 2: 206 223.
Ehlers, E., 1908. Die Bodensässigen Anneliden aus den Sammlungen der deutschen Tiefsee- Expedition. In: C. Chun (ed.),
Wissenschaftliche Ergebruisse der deutschen Tiefsee- Expedition auf dem Dampfer “Valdivia” 1898–1899. Volume 16,
167 pp., 23 pls. Gustav Fischer, Jena.
Ehlers, E., 1913. Die Polychaeten-Sammlung der deutschen Südpolar-Expedition 1901 1903. Deutsche Südpolar-Expedition,
13: 397 598, pls. 26 46.
Fabricius, O., 1780. Fauna Groenlandica, systematice sistens, Animalia Groenlandica occidentalis hactenus indagata, quoad
nomen specificum, triviale, vernaculumque; synonyma auctorum plurium, descriptionem, locum, generationem, mores,
usum, capturamque singuli; pront detegendi occasio fuit, maximaque parti secundum proprias observationes. 452 pp.
(Polychaeta, pp. 279 315.) Hafniae et Lipsiae.
Fauchald, K., 1963. Nephtyidae (Polychaeta) from the Bay of Nha Trang, South Viet Nam. NAGA Report, 4: 1 33.
Fauchald, K., 1970. Polychaetous annelids of the families Eunicidae, Lumbrineridae, Iphitimidae, Arabellidae, Lisaretidae
and Dorvilleidae from western Mexico. Allan Hancock Monographs in Marine Biology, 5: 1 335, 27 pls.
Fauchald, K., 1972a. Some polychaetous annelids from deep basins in Sognefjorden, western Norway. Sarsia, 49: 89 106.
Fauchald, K., 1972b. Benthic polychaetous annelids from deep water off western Mexico and adjacent areas in the eastern
Pacific Ocean. Allan Hancock Monographs in Marine Biology , 7: 1 575.
Fauchald, K., 1974. Sphaerodoridae (Polychaeta: Errantia) from world-wide areas. Journal of Natural History, 8: 257 289.
Fauchald, K., 1977. Some nereid polychaetes from deep water. Essays on polychaetous annelids in memory of Dr. Olga Hartman. pp. 157 171. Allan Hancock Foundation, University of Southern California, Los Angeles.
Fauchald, K., 1982. Revision of Onuphis, Nothria and Paradiopatra (Polychaeta: Onuphidae) based upon type material.
Smithsonian Contributions to Zoology, (356): i vi + 1 109.
Fauvel, P., 1913. Quatrième note préliminaire sur les Polychètes provenant des campagnes de l'Hirondelle et de la PrincesseAlice, ou déposées dans le Musée Océanographique de Monaco. Bulletin de l’ Institut Océanographique de Monaco,(270): 1 80.
Fauvel, P., 1914. Annélides polychètes non-pèlagiques provenant des campagnes de l'Hirondelle et de la Princesse-Alice (1885 1910). Résultats des campagnes scientifiques accompliés sur son yacht par le Pince Albert 1, 46: 1 432, 31
pls.
Fauvel, P., 1923. Polychètes errantes. Faune de France, 5: 1 488.
Fauvel, P., 1925. Sur quelques espèces du genre Aphrodita. Bulletin sociéte zoologique de France, Paris, 50: 131 150.
Fauvel, P., 1927. Polychètes sédentaires. Addenda aux errantes, archiannélides, myzostomaires. Faune de France, 16:
1 494.
Fauvel, P., 1932a. Polychètes nouvelles de Che-Foo (China). Bulletin du Muséum National d’Histoire Naturelle, Paris, Series
2, 4: 536 538.
Fauvel, P., 1932b. Annelida Polychaeta of the Indian Museum, Calcutta. Memoirs of the Indian Museum, 12: 1-262, 9 pls.
Fauvel, P., 1936a. Annélides Polychètes du Japon. Memoirs of the College of Science, Kyoto Imperial University, Series B,
12: 41 92.
Fauvel, P., 1936b. Contribution à la Faune des Annélides Polychètes du Maroc. Mémoires de la Société des Sciences Naturelles du Maroc, 43: 1 143.
Fauvel, P., 1953. Annelida Polychaeta. The Fauna of India, including Pakistan, Ceylon, Burma and Malaya, 507 pp. The Indian Press Ltd, Allahabad.
Frame, A. B., 1992. The lumbrinerids (Annelida: Polychaeta) collected in two northwestern Atlantic surveys with descriptions of a new genus and two new species. Proceedings of the Biological Society of Washington, 105: 185 218.
Gallardo, V. A., 1968. Polychaeta from the Bay of Nha Trang, South Viet Nam. NAGA Report, Scientific Results of Marine
Investigations of the South China Sea and the Gulf of Thailand 1959–1961, 4: 35 279, pls. 1 59.
Gardiner, S. L., 1976. Errant polychaete annelids from North Carolina. Journal of the Elisha Mitchell Scientific Society, 91:
77 220.
Gravier, C., 1909. Annélides Polychètes recueillis à Payta (Pérou) par M. le Dr. Rivet. Archives de Zoologie Expérimentale
et Générale, Series 4, 10: 617 659, pls. 16 17.
Grube, A. E., 1840. Actinien, Echinodermen und Würmer des Adriatischen und Mittelmeeres nach eigenen Sammlungenbeschrieben. 92 pp. J. H. Bon, Königsberg.
Grube, A. E., 1860. Beschreibung neuer oder wenig bekannter Anneliden. Archiv für Naturgeschichte, Berlin, 26: 71 118.
Grube, A. E., 1863. Beschreibung neuer oder wenig bekannter Anneliden. Zahlreiche Gattungen. Archiv für Naturgeschichte,
Berlin, 29: 37 69.

Deep-sea Benthic Polychaetes off Northern Honshu

185

Grube, A. E., 1877. Die von der Gazelle mitgebrachten Anneliden, zu denen noch zwei von Dr. Buchholz gesammelte kommen. Monatsberichte der Königlich preussischen Akademie der Wissenschaften zu Berlin, 1877: 509 554.
Haase, P., 1915. Boreale und arktische Chloraemiden. Wissenschaftliche Meeresunter- suchungen, 17: 169 228, pls. 1 2.
Hansen, G. A., 1878. Anneliden fra den norske Nordhavsexpedition i 1876. Nyt Magazin for Naturvidenskaberne, Oslo, 24:
1 17, 10 pls.
Hartman, O., 1936. A review of the Phyllodocidae (Annelida Polychaeta) of the coast of California, with descriptions of nine
new species. University of California Publications in Zoology, 41: 117 132.
Hartman, O., 1938. Review of the annelid worms of the family Nephtyidae from the northeast Pacific, with descriptions offive new species. Proceedings of the United States National Museum, 85: 143 158.
Hartman, O., 1944a. New England Annelida. Part 2. Including the unpublished plates by Verrill with reconstructed captions.
Bulletin of the American Museum of Natural History, 82: 327 344.
Hartman, O., 1944b. Polychaetous annelids. Part 5. Eunicea. Allan Hancock Pacific Expeditions, 10: 1 238.
Hartman, O., 1945. The marine annelids of North Carolina. Bulletin of Duke University Marine Station, 2: 1 54, 10 pls., 2
charts.
Hartman, O., 1947a. Polychaetous annelids. Part 8. Pilargiidae. Allan Hancock Pacific Expeditions, 10: 483 524.
Hartman, O., 1947b. Polychaetous annelids. Part 7. Capitellidae. Allan Hancock Pacific Expeditions, 10: 391 481.
Hartman, O., 1948a. The polychaetous annelids of Alaska. Pacific Science, 2: 3 58.
Hartman, O., 1948b. The marine annelids erected by Kinberg with notes on some other types in the Swedish State Museum.
Arkiv för Zoologi, Stockholm, 42A: 1 137.
Hartman, O., 1950. Goniadidae, Glyceridae and Nephtyidae. Allan Hancock Pacific Expeditions, 15: 1 181, 19 pls.
Hartman, O., 1953. Non-pelagic Polychaeta of the Swedish Antarctic Expedition 1901 1903. Further Zoological Results, 4:
1 83.
Hartman, O., 1955. Quantitative survey of the benthos of San Pedro Basin, southern California. Part I. Preliminary results.
Allan Hancock Pacific Expeditions, 19: 1 185, 7 pls, 2 charts.
Hartman, O., 1956. Polychaetous annelids erected by Treadwell, 1891 to 1948, together with a brief chronology. Bulletin of
the American Museum of Natural History, 109: 239 310.
Hartman, O., 1957. Orbiniidae, Apistobranchidae, Paraonidae and Longosomidae. Allan Hancock Pacific Expeditions, 15:
211 393.
Hartman, O., 1959. Catalogue of the polychaetous annelids of the world. Parts I, II. Allan Hancock Foundation Publications
Occasional Paper, (23): 628 pp.
Hartman, O., 1960. Systematic account of some marine invertebrate animals from the deep basins off Southern California. In:
The benthic fauna of the deep basins off Southern California, Part II. Allan Hancock Pacific Expeditions, 22: 69 215.
Hartman, O., 1961. Polychaetous annelids from California. Allan Hancock Pacific Expeditions, 25: 1 226.
Hartman, O., 1965. Deep-water benthic polychaetous annelids off New England to Bermuda and other North Atlantic areas.
Allan Hancock Foundation Publications Occasional Paper, (28): 1 378.
Hartman, O., 1966. Polychaeta Myzostomidae and Sedentaria of Antarctica. Antarctic Research Series, 7: 1 158.
Hartman, O., 1968. Atlas of the errantiate polychaetous annelids from California, 828 pp. Allan Hancock Foundation, University of Southern California, Los Angeles.
Hartman, O., 1969. Atlas of the sedentariate polychaetous annelids from California, 812 pp. Allan Hancock Foundation,
University of Southern California, Los Angeles.
Hartman, O., 1978. Polychaeta from the Weddell Sea Quadrant Antarctica. Antarctic Research Series, 26: 125 223.
Hartman, O. and K. Fauchald, 1971. Deep-water benthic polychaetous annelids off New England to Bermuda and other North
Atlantic areas. Part II. Allan Hancock Monographs in Marine Biology, 6: 1 327, 34 pls.
Hartman, O. and D. J. Reish, 1950. The marine annelids of Oregon. Oregon State College Monographs in Zoology, 6: 1 64.
Hartmann-Schröder, G., 1958. Einige Polychaeten aus dem Küstengrundwasser der Bimini- Inseln (Bahamas). Kieler
Meeresforschungen, 14: 233 240.
Hartmann-Schröder, G., 1959. Zur Ökologie der Polychaeten des Mangrove-Estero-Gebietes von El Salvador. Beiträge zue
Neotropischen Fauna, 1: 69 183.
Hartmann-Schröder, G., 1965. Die Polychaeten des Sublitorals. Zur Kenntnis des Sublitorals der chilenischen Küste unterbesonderer Berücksichtigung der Polychaeten und Ostracoden. Mitteilungen aus dem Hamburgischen Zoologischen
Museum und Institut, 62: 59 305.
Hartmann-Schröder, G., 1971. Annelida, Borstenwürmer, Polychaeta. Die Tierwelt Deutschlands und der angrenzenden
Meeresteile nach ihren Merkmalen und nach ihrer Lebensweise, 58: 594 pp.
Hartmann-Schröder, G., 1996. Annelida, Borstenwürmer, Polychaeta. Die Tierwelt Deutschlands und der angrenzenden
Meeresteile nach ihren Merkmalen und nach ihrer Lebensweise, 2. Auflage. Volume 58. 648 pp. Gustav Fischer, Jena.
Hessle, C., 1917. Zur Kenntnis der terebellomorphen Polychaeten. Zoologiska Bidrag från Uppsala, 5: 39 258.
Hilbig, B., 1994a. 7. Family Goniadidae Kinberg, 1866. In: Blake, J. A. and B. Hilbig (eds.), Taxonomic Atlas of the benthic
fauna of the Santa Maria Basin and Western Santa Barbara Channel, Volume 4. The Annelida Part 1. Oligochaeta and186

Minoru Imajima

Polychaeta: Phyllodocida (Phyllodocidae to Paralacydoniidae), pp. 215 230. Santa Barbara Museum of Natural History, Santa Barbara, California.
Hilbig, B., 1994b. 9. Family Hesionidae Sars, 1862. In: Blake, J. A. and B. Hilbig (eds.), Taxonomic Atlas of the benthic
fauna of the Santa Maria Basin and Western Santa Barbara Channel, Volume 4. The Annelida Part 1. Oligochaeta andPolychaeta: Phyllodocida (Phyllodocidae to Paralacydoniidae), pp. 243 269. Santa Barbara Museum of Natural History, Santa Barbara, California.
Hilbig, B., 2000. 8. Family Ampharetidae Malmgren, 1867. In: Blake, J. A., B. Hilbig and P. V. Scott (eds.), Taxonomic Atlasof the benthic fauna of the Santa Maria Basin and Western Santa Barbara Channel, Volume 7. The Annelida Part 4.
Polychaeta: Flabelligeridae to Sternaspidae, pp. 169 230. Santa Barbara Museum of Natural History, Santa Barbara,
California.
Hobson, K. D., 1976. Notes on benthic sedentariate Polychaeta (Annelida) from British Columbia and Washington. Syesis, 9:
135 142.
Hobson, K. D. and K. Banse, 1981. Sedentariate and archiannelid polychaetes of British Columbia and Washington. Canadian Bulletin of Fisheries and Aquatic Sciences, (209): 1 144.
Horst, R., 1903. New species of the genus Euphrosine from the Siboga-Expedition, with a table of the species hitherto known.
Notes of the Leyden Museum, 23: 213 222.
Horst, R., 1912. Polychaeta errantia of the Siboga-Expedition. Part 1. Amphinomidae. Siboga- Expeditie, Leiden, 24a: 1 43,
10 pls.
Horst, R., 1915. On new and little-known species of Polynoinae from the Netherlands’ East Indies. Zoologische Mededeelingen, Leiden, 1: 2 20.
Horst, R., 1916. Malayan species of the genera Aphroditella, Hermione, Laetmonice and Aphrogenia. Zoologische Mededeelingen, Leiden, 2: 65 77.
Horst, R., 1917. Polychaeta Errantia of the Siboga-Expedition. Part 2. Aphroditidae and Chryso- petalidae. Siboga- Expeditie,
Leiden, 24b: 1 140.
Hutchings, P. A. and J. McRae, 1993. The Aproditidae (Polychaeta) from Australia, together with a redescription of the Aphroditidae collected during the Siboga-Expedition. Records of the Australian Museum, 45: 279 363.
Imajima, M., 1961. Polychaetous annelids collected off the west coast of Kamchatka. I. Notes on species found in the collection of 1957 58. Publications of the Seto Marine Biological Laboratory, 9: 81 102.
Imajima, M., 1963. Polychaetous annelids collected off the west coast of Kamchatka. II. Notes on species found in the collection of 1959. Publications of the Seto Marine Biological Laboratory, 11: 175 202.
Imajima, M., 1964. Benthic polychaetes collected by the second cruise of the Japanese Expedition of Deep Seas (JEDS 2).
Bulletin of the National Science Museum, 7: 235 254.
Imajima, M., 1966a. The Syllidae (polychaetous annelids) from Japan. III. Eusyllinae. Publications of the Seto Marine Biological Laboratory, 14: 85 116.
Imajima, M., 1966b. The Syllidae (polychaetous annelids) from Japan. I. Exogoninae. Publications of the Seto Marine Biological Laboratory, 13: 385 404.
Imajima, M, 1966c. The Syllidae (polychaetous annelids) from Japan. IV. Syllinae (1). Publications of the Seto Marine Biological Laboratory, 14: 219 252.
Imajima, M., 1966d. The Syllidae (polychaetous annelids) from Japan. V. Syllinae (2). Publications of the Seto Marine Biological Laboratory, 14: 253 294.
Imajima, M., 1968. Benthonic polychaetes of Miyako Bay. Memoirs of the National Science Museum, (1): 139 144. (InJapanese.)
Imajima, M., 1969. Three species of the family Sphaerodoridae (polychaetous annelids) from Japan. Bulletin of the National
Science Museum, 12: 151 156.
Imajima, M., 1972. Biomass of the benthonic polychaetes at the fishing-ground for gill net off Samani, Hokkaido. Memoirsof the National Science Museum, (5): 11 16. (In Japanese.)
Imajima, M., 1973. Paraonidae (Polychaeta) from Japan. Bulletin of the National Science Museum, 16: 253 292.
Imajima, M., 1986. Eight species of Onuphidae (Polychaeta) in and offshore of Otsuchi Bay, northeastern Japan. Bulletin of
the National Science Museum, Series A (Zoology), 12: 93 116.
Imajima, M., 1987. Pilargidae (Annelida, Polychaeta) from Japan (Part 1). Bulletin of the National Science Museum, Series
A (Zoology), 13: 151 164.
Imajima, M., 1988. Polychaetous annelids of Ishikari Bay, Hokkaido. Memoirs of the National Science Museum, (21):123 129. (In Japanese.)
Imajima, M., 1989a. Two new species of Nothria (Polychaeta, Onuphidae) from off Sanriku, northeastern Honshu, Japan.
Bulletin of the National Science Museum, Series A (Zoology), 15: 1 10.
Imajima, M., 1989b. Poecilochaetidae (Annelida, Polychaeta) from Japan. Bulletin of the National Science Museum, Series
A (Zoology), 15: 61 103.
Imajima, M., 1990. Spionidae (Annelida, Polychaeta) from Japan. IV. The genus Prionospio (Prionospio). Bulletin of theDeep-sea Benthic Polychaetes off Northern Honshu

187

National Science Museum, Series A (Zoology), 16: 105 140.
Imajima, M., 1991. Spionidae (Annelida, Polychaeta) from Japan. VII. The genus Spiophanes. Bulletin of the National Science Museum, Series A (Zoology), 17: 115 137.
Imajima, M., 1992. Polychaetous annelids around Sarufutsu, northern Hokkaido. Memoirs of the National Science Museum,(25): 125 133. (In Japanese.)
Imajima, M., 1994. Polychaetous annelids of Akkeshi Bay, Hokkaido. Memoirs of the National Science Museum, (27):
105 126.
Imajima, M., 1996. Polychaeta. Syllidae, Nereididae, Nephtyidae, Spionidae, Maldanidae, Serpulidae. 530 pp. Seibutsu Kenkyusha Co., Ltd., Tokyo. (In Japanese.)
Imajima, M., 1997a. Polychaetous annelids from Suruga Bay, Central Japan. In: Deep-sea Fauna and Pollutants in Suruga
Bay. National Science Museum Monographs, (12): 149 228.
Imajima, M., 1997b. Polychaetous annelids from Sagami Bay and Sagami Sea collected by the Emperor Showa of Japan anddeposited at the Showa Memorial Institute, National Science Museum, Tokyo. Families Polynoidae and Acoetidae.
National Science Museum Monographs, (13): 1 131.
Imajima, M., 1999. Onuphidae (Annelida, Polychaeta) from Japan, excluding the genus Onuphis. National Science Museum
Monographs, (16): 1 115.
Imajima, M., 2001a. Deep-sea benthic polychaetous annelids of Tosa Bay, southwestern Japan. In: Fujita, T., H. Saito and M.
Takeda (eds.), Deep-Sea Fauna and Pollutants in Tosa Bay. National Science Museum Monographs, (20): 31 100.
Imajima, M., 2001b. Polychaeta II. Polynoidae, Acoetidae, Eulepethidae, Sphaerodoridae, Pilargidae, Paralacydoniidae, Onuphidae, Lumbrineridae, Dorvilleidae, Paraonidae, Apistobranchidae, Trochochaetidae, Poecilochaetidae, Heterospionidae, Oweniidae, Trichobranchidae. 542 pp. Seibutsu Kenkyusha Co., Ltd., Tokyo. (In Japanese.)
Imajima, M., 2003. Polychaetous annelids from Sagami Bay and Sagami Sea collected by the Emperor Showa of Japan anddeposited at the Showa Memorial Institute, National Science Museum, Tokyo II. Orders included within the Phyllodocida, Amphinomida, Spintherida and Eunicida. National Science Museum Monographs, (23): 1 221.
Imajima, M., 2005. Deep-sea benthic polychaetous annelids from around Nansei Islands. In: Hasegawa, K., G. Shinohara and
M. Takeda (eds.), Deep-sea Fauna and Pollutants in Nansei Islands. National Science Museum Monographs, (29):37 99.
Imajima, M., 2006. Polychaetous annelids from Sagami Bay and Sagami Sea, central Japan. Memoirs of the National Science
Museum, (40): 317 408.
Imajima, M., 2007. Polychaeta III. Aphroditidae, Sigalionidae, Pholoidae, Spintheridae, Glyceridae, Eunicidae, Hesionidae,
Euphrosinidae. 499 pp. Seibutsu Kenkyusha Co., Ltd., Tokyo. (In Japanese.)
Imajima, M. and O. Hartman, 1964. The polychaetous annelids of Japan. Parts I, II. Allan Hancock Foundation Publications
Occasional Paper, (26): 1 452, 38 pls.
Imajima, M. and K. Hayashi, 1969. Seasonal changes of polychaetes living among the attaching organisms. Proceedings of
the Japanese Society of Systematic Zoology, (5): 2 15. (In Japanese and English.)Imajima, M. and M. Higuchi, 1975. Lumbrineridae of polychaetous annelids from Japan, with descriptions of six new species. Bulletin of the National Science Museum, Series A (Zoology), 1: 5 37.
Imajima, M. and Y. Morita, 1987. Oweniidae (Annelida, Polychaeta) from Japan. Bulletin of the National Science Museum,Series A (Zoology), 13: 85 102.
Imajima, M. and Y. Shiraki, 1982a. Maldanidae (Annelida: Polychaeta) from Japan. Part 1. Bulletin of the National Science
Museum, Series A (Zoology), 8: 7 46.
Imajima, M. and Y. Shiraki, 1982b. Maldanidae (Annelida: Polychaeta) from Japan. Part 2. Bulletin of the National Science
Museum, Series A (Zoology), 8: 47 88.
Imajima, M. and Y. Takeda, 1985. Nephtyidae (Polychaeta) from Japan. I. The genera Inermonephtys, Micronephtys andAglaophamus. Bulletin of the National Science Museum, Series A (Zoology), 11: 57 90.
Imajima, M. and Y. Takeda, 1987. Nephtyidae (Polychaeta) from Japan. II. The genera Dentinephtys and Nephtys. Bulletin of
the National Science Museum, Series A (Zoology), 13: 41 77.
Imajima, M. and S. J. Williams, 1985. Trichobranchidae (Polychaeta) chiefly from the Sagami and Suruga bays, collected by
R/V Tansei-Maru (Cruises KT-65〜76). Bulletin of the National Science Museum, Series A (Zoology), 11: 7 18.
Izuka, A., 1902. On two new species of the family Maldanidae from the Sagami Bay. Annotationes Zoologicae Japonenses,
4: 109 114, pl. 3.
Izuka, A., 1912. The errantiate Polychaeta of Japan. The Journal of the College of Science, Imperial University of Tokyo, 30:
1 262.
Johansson, K. E., 1927. Beiträge zur Kenntniss der Polychaeten-Familien Hermelidae, Sabellidae und Serpulidae. Zoologiska
Bidrag från Uppsala, 11: 1 184, 5 pls.
Johnson, H. P., 1901. The Polychaeta of the Puget Sound Region. Proceedings of the Boston Society of Natural History, 29:
381 437.
Johnston, G., 1839. The British Aphroditacea. Annals and Magazine of Natural History, London, Series 1, 2: 424 441.

188

Minoru Imajima

Kinberg, J. G. H., 1856. Nya slägten och arter af Annelider. Öfversigt af Kongliga Vetenskaps- Akademiens Förhandlingar,
Stockholm, 12: 381 388.
Kinberg, J. G. H., 1866. Annulata nova. Öfversigt af Kongliga Vetenskaps-Akademiens Förhandlingar, Stockholm, 22:239 258.
Kinberg, J. G. H., 1910. Annulater. Kongliga Svenska Fregatten Eugenies Resa omkring jorden under befäl af C.A.Virgin åren1851–1853, Uppsala and Stockholm, 7: 33 78, pls. 9 29.
Kirkegaard, J. B., 1983. Bathyal benthic polychaetes from the N.E. Atlantic Ocean, S.W. of the British Isles. Journal of the
Marine Biological Association of the United Kingdom, 63: 593 608.
Kirkegaard, J. B., 1995. Bathyal and abyssal polychaetes (errant species). Galathea Reports, 17: 7 56.
Kröyer, H., 1856. Bidrag til kundskab af Sabellerne. Öfversigt af Kongliga Danske Videnskabernes Selskab Förhandlingar,
1 36.
Kucheruk, N. V., 1978. Deep-water Onuphidae (Polychaeta) from the collections of the 16th cruise of the R/V Dmitry Mendeleev (to the generic classification of the family Onuphidae). Trudy Instituta Okeanologii, Akademii Nauk SSSR, 113:
88 106. (In Russian with English summary.)
Kucheruk, N. V., 1979. New species of the family Onuphidae (Polychaeta) from the Pacific. Zoologisheskij Zhurnal, Akademii Nauk SSSR, 58: 1227 1230. (In Russian with English summary.)
Kudenov, J. D., 1993. Amphinomidae and Euphrosinidae (Annelida: Polychaeta) principally from Antarctica, the Southern
Ocean, and Subantarctic regions. Biology of the Antarctic Seas XXII. Antarctic Research Series, 58: 93 150.
Kudenov, J. D., 1994. 8. Family Sphaerodoridae Malmgren, 1867. In: Blake, J. A. and B. Hilbig (eds.), Taxonomic Atlas of
the benthic fauna of the Santa Maria Basin and Western Santa Barbara Channel, Volume 4. The Annelida Part 1. Oligochaeta and Polychaeta: Phyllodocida (Phyllodocidae to Paralacydoniidae), pp. 231 242. Santa Barbara Museum of
Natural History, Santa Barbara, California.
Kudenov, J. D. and J. A. Blake, 1978. A review of the genera and species of the Scalibregmidae (Polychaeta) with descriptions of one new genus and three new species from Australia. Journal of Natural History, 12: 427 444.
Langerhans, P., 1880. Die Wurmfauna von Madeira. Part 2. Zeitschrift für wissenschaftliche Zoologie, Leipzig, 33:
267 316.
Lee, J. H. and J. G. Jae, 1983. Polychaetous annelids from the Yellow Sea. I. Family Nephtyidae. Bulletin of Korea Ocean
Research and Development Institute, 5: 19 27.
Levenstein, R. Ya., 1962. The polychaetes from three abyssal trenches in the Pacific Ocean. Zoologisheskij Zhurnal, Akademii
Nauk SSSR, 41: 1142 1148. (In Russian with English summary.)
Levenstein, R. Ya., 1966. Polychaeta of the western part of the Bering Sea. Trudy Instituta Okeanologii P. P. Shirshov, Akademii Nauk SSSR, 81: 3 131.
Levenstein, R. Ya., 1972. Ecology and zoogeography of some Polychaeta representatives of the abyssal Pacific. Proceedingsof the Royal Society of Edinburgh, Section B, 73: 171 181.
Levinsen, G. M. R., 1883. Systematisk-geografisk Oversigt over de nordiske Annulata, Gephyrea, Chaetognathi og Balanoglossi. 2. Videnskabelige Meddelelser fra Dansk naturhistorisk Førening i Kjøbenhavn, 1883: 92 350.
Licher, F. and W. Westheide, 1997. Review of the genus Sigambra (Polychaeta: Hesionidae), redescription of S. bassi (Hartman, 1947), and descriptions of two new species from Thailand and China. Steenstrupia, 23: 1 20.
Light, W. J., 1977. Spionidae (Annelida: Polychaeta) from San Francisco Bay, California: a revised list of nomenclaturalchanges, new records, and comments on related species from the northeastern Pacific Ocean. Proceedings of the Biological Society of Washington, 90: 66 88.
Light, W. J., 1991. Systematic revision of the genera of the polychaete subfamily Maldaninae Arwidsson. Ophelia, Supplement 5: 133 146.
Linnaeus, C., 1758. Systema naturae. Per regna tria naturae, secundum classes, ordines, genera, species, cum characteribus,
differentiis synonymis locis, 10th edition. 824 pp. Laurentii Salvii, Holmiae.
Linnaeus, C., 1767. Systema naturae. Per regna tria naturae, secundum classes, ordines, genera, species, cum characteribus,
differentiis synonymis locis, 12th edition. 1327 pp. Holmiae.
Lovell, L. L., 2002. Paraonidae (Annelida: Polychaeta) of the Andaman Sea, Thailand. Phuket Marine Biological Center
Special Publication, 24: 33 56.
Lützen, J., 1961. Sur une nouvelle espèce de polychète Sphaerodoridium commensalis, n. gen., n. spec., (Polychaeta Errantia,
famille des Sphaerodoridae) vivant en commensal de Terebellides stroemi Sars. Cahiers de Biologie Marine, 2:
409 416.
Maciolek, N. J., 1985. A revision of the genus Prionospio Malmgren, with special emphasis on species from the Atlantic
Ocean, and new records of species belonging to the genera Apoprionospio Foster and Paraprionospio Caullery (Polychaeta, Annelida, Spionidae). Zoological Journal of the Linnean Society, 84: 325 383.
Mackie, A. S. Y., 1987. A review of species currently assigned to the genus Leitoscoloplos Day, 1977 (Polychaeta: Orbiniidae), with descriptions of species newly referred to Scoloplos Blainville, 1828. Sarsia, 72: 1 28.
Mackie, A. S. Y. and S. J. Chambers, 1990. Revision of the type species of Sigalion, Thalanessa and Eusigalion (Polychaeta:

Deep-sea Benthic Polychaetes off Northern Honshu

189

Sigalionidae). Zoologica Scripta, 19: 39 56.
Mackie, A. S. Y. and F. Pleijel, 1995. A review of the Melinna cristata species group (Polychaeta: Ampharetidae) in thenortheastern Atlantic. Mitteilungen aus den Hamburgischen Zoologischen Museum und Institut, 92: 103-124.
Maekawa, N. and I. Hayashi, 1989. Onuphid polychaetes from Wakasa Bay, Sea of Japan. Memoirs of the College of Agriculture, Kyoto University, (134) : 61 93.
Maekawa, N. and I. Hayashi, 1999. Taxonomic study on the genus Onuphis (Polychaeta, Onuphidae) from Japan and adjacentseas, with description of six new species. Bulletin of the National Science Museum, Series A (Zoology), 25: 163 214.
Malmgren, A. J., 1865. Nordiska Hafs-Annulater. Öfversigt af Kongliga Vetenskaps-Akademiens Förhandlingar, Stockholm,
21: 51 110, 181 192.
Malmgren, A. J., 1866. Nordiska Hafs-Annulater. Öfversigt af Kongliga Vetenskaps-Akademiens Förhandlingar, Stockholm,
22: 355 410.
Malmgren, A. J., 1867a. Spetsbergens, Grönlands, Islands och den Scandinaviska halföns hittills kända Annulata polychaeta.
127 pp. Thesis. J. C. Frenckell, Helsingfors.
Malmgren, A. J., 1867b. Annulata Polychaeta Spetsbergiae, Grönlandiae, Islandiae et Scandinaviae hactenus cognita. Öfversigt af Kongliga Vetenskaps-Akademiens Förhandlingar, Stockholm, 24: 127 235, pls. 2 15. [Identical to 1867a.]
Marenzeller, E. von, 1879. Südjapanische Anneliden I. (Amphinomea, Aphroditea, Lycoridea, Phyllodocea, Hesionea, Syllidea, Eunicea, Glycerea, Sternaspidea, Chaetopterea, Cirratulea, Amphictenea). Denkschriften der Mathematischnaturwissenschaftlichen Classe der Kaiserlichen Akademie der Wissenschaften, Wien, 41: 109 154.
Marenzeller, E. von, 1884. Südjapanische Anneliden II. (Ampharetea, Terebellacea, Sabellacea, Serpulacea). Denkschriftender Mathematisch-naturwissenschaftlichen Classe der Kaiserlichen Akademie der Wissenschaften, Wien, 49: 197 224.
Marenzeller, E. von, 1902. Südjapanische Anneliden III. Aphroditea, Eunicea. Denkschriften der Mathematisch-naturwissenschaftlichen Classe der Kaiserlichen Akademie der Wissenschaften, Wien, 72: 563 582.
McIntosh, W. C., 1879. On the Annelida obtained during the cruise of HMS. Valorous to Davis Strait in 1875. Transactionsof the Linnean Society of London, New Series, 1: 499 511.
McIntosh, W. C., 1885. Report on the Annelida Polychaeta collected by the H.M.S. Challenger during the years 1873 76.
Report on the scientific results of the Voyage of H.M.S. Challenger during the years 1873 76 under the command of the
Captain George S. Nares, R. N., F. R. S. and the Late Captain Frank Tourle Thomson, R. N., Zoology, 12: 1 554.
McIntosh, W. C., 1914. Notes from the Gatty Marine Laboratory, St. Andrews, No. 36. 2. On some of the species of Prionospio Malmgren. 3. On the British Amphictenidae. 4. On the British Ampharetidae. Annals and Magazine of Natural
History, Series 8, 13: 77 110.
McIntosh, W. C., 1922. Notes from the Gatty Marine Laboratory, St. Andrews, No. 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.
Mesnil, F. and M. Caullery, 1898. Étude de morphologie externe chez les Annélides. La familie nouvelle des Levinseniens.
Revision des Ariciens-affinites des deux familles. Les Aspitobranchiens. Bulletin Biologique de la France et de la Belgique, 31: 126 150.
Mesnil, F. and P. Fauvel, 1939. Polychètes sédentaries de l’expédition des Siboga. Maldanidae, Cirratulidae, Capitellidae,
Sabellidae et Serpulidae. Siboga-Expedition, 24.2: 1 42.
Miura, T., 1986. Japanese polychaetes of the genera Eunice and Euniphysa: Taxonomy and branchial distribution patterns.
Publications of the Seto Marine Biological Laboratory, 31: 269 325.
Monro, C. C. A., 1930. Polychaete worms. Discovery Reports, 2: 1 222.
Monro, C. C. A., 1936. Polychaeta worms II. Discovery Reports, 12: 59 197.
Moore, J. P., 1902. Descriptions of some new Polynoidae, with a list of other Polychaeta from North Greenland waters. Proceedings of the Academy of Natural Sciences of Philadelphia, 54: 258 278, pls. 13 14.
Moore, J. P., 1903. Polychaeta from the coastal slope of Japan and from Kamchatka and Bering Sea. Proceedings of the
Academy of Natural Sciences of Philadelphia, 55: 401 490, pls. 23 27.
Moore, J. P., 1904. New Polychaeta from California. Proceedings of the Academy of Natural Sciences of Philadelphia, 56:
484 502.
Moore, J. P., 1905. New species of Polychaeta from the North Pacific, chiefly from Alaskan waters. Proceedings of the Academy of Natural Sciences of Philadelphia, 57: 525 554, pls. 34 36.
Moore, J. P., 1906. Additional new species of Polychaeta from the North Pacific. Proceedings of the Academy of Natural
Sciences of Philadelphia, 58: 217 260, pls. 10 12.
Moore, J. P., 1907. Description of new species of spioniform annelids. Proceedings of the Academy of Natural Sciences ofPhiladelphia, 59: 195 207.
Moore, J. P., 1908. Some polychaetous annelids of the northern Pacific coast of North America. Proceedings of the Academyof Natural Sciences of Philadelphia, 60: 321 364.
Moore, J. P., 1909. The polychaetous annelids dredged by the U. S. S. Albatross off the coast of Southern California in 1904.
I. Syllidae, Sphaerodoridae, Hesionidae and Phyllodocidae. Proceedings of the Academy of Natural Sciences of Phila-

190

Minoru Imajima

delphia, 61: 321 351, pls. 15 16.
Moore, J. P., 1910. The polychaetous annelids dredged by the U. S. S. Albatross off the coast of Southern California in 1904.
II. Polynoidae, Aphroditidae and Sigalionidae. Proceedings of the Academy of Natural Sciences of Philadelphia, 62:
328 402, pls. 28 33.
Moore, J. P., 1911. The polychaetous annelids dredged by the U. S. S. Albatross off the coast of Southern California in 1904.
III. Euphrosynidae to Goniadidae. Proceedings of the Academy of Natural Sciences of Philadelphia, 63: 234 318, pls.
15 21.
Moore, J. P., 1923. The polychaetous annelids dredged by the U. S. S. Albatross off the coast of Southern California in 1904.
Spionidae to Sabellariidae. Proceedings of the Academy of Natural Sciences of Philadelphia, 75: 179 259, pls. 17 18.
Müller, O. F., 1776. Zoologiae Danicae prodromus, seu animalium daniae et Norvegiae indigenarum characters, nomine etsynonyma imprimis popularium. 274 pp. Havniae.
Nilsen, R. and T. Holthe, 1985. Arctic and Scandinavian Oweniidae (Polychaeta) with comments on the phylogeny of thefamily. Sarsia, 70: 17 32.
Nilsson, D., 1928. Neue und alte Amphicteniden. Göteborgs Kongliga Vetenskaps och Vitterhets-Samhälles, Handlingar,
Series 4, 33: 1 96.
O’Connor, B. D. S., 1987. The Glyceridae (Polychaeta) of the North Atlantic and Mediterranean, with descriptions of two
new species. Journal of Natural History, 21: 167 189.
Okuda, S., 1934. Two species of the sedentary polychaete Pectinaria. Annotationes Zoologicae Japonenses, 14: 321 326.
Okuda, S., 1936. Polychaetous annelids from Toyama Bay and its adjacent waters. I. Polychaeta sedentaria. Bulletin of the
Biogeographical Society of Japan, 6: 147 157.
Okuda, S., 1937a. Some ariciid worms from Japan. Annotationes Zoologicae Japonenses, 16: 99 105.
Okuda, S., 1937b. Spioniform polychaetes from Japan. The Journal of the Faculty of Science, Hokkaido Imperial University,
Series 6, Zoology, 5: 217 254.
Okuda, S., 1937c. Annelida Polychaeta in Onagawa Bay and its vicinity. I. Polychaeta sedentaria. Science Reports of the
Tohoku Imperial University, Fourth Series, Biology, 12: 45 69, pl. 2.
Okuda, S., 1938a. Polychaetous annelids from the vicinity of the Mitsui Institute of Marine Biology. Japanese Journal of
Zoology, 8: 75 105.
Okuda, S., 1938b. The Sabellariidae of Japan. The Journal of the Faculty of Science, Hokkaido Imperial University, Series 6,
Zoology, 6: 235 253.
Okuda, S., 1939. Annelida Polychaeta in Onagawa Bay and its vicinity. II. Polychaeta errantia with some addenda of Polychaeta sedentaria. Science Reports of the Tohoku Imperial University, Fourth Series, Biology, 14: 219 244.
Okuda, S., 1943. Occurrence of a freshwater polychaete in central China. The Journal of the Shanghai Science Institute, New
Series, 2: 99 103.
Orensanz, J. M., 1973. Los anélidos poliquetos de la Provincia Biogeográfica Argentina. IV. Lumbrineridae. Physis Sección
A, Buenos Aires, 32: 343 393.
Orensanz, J. M., 1974. Los anélidos poliquetos de la Provincia Biogeográfica Argentina. VI. Arabellidae. Physis Sección A,
Buenos Aires, 33: 381 408.
Orensanz, J. M., 1990. The eunicemorph polychaete annelids from Antarctic and Subantarctic seas. Biology of the Antarctic
Seas XXI. Antarctic Research Series, 52: 1 183.
Parapar, J., 2001. Revision of five species referred to Myriochele and Galathowenia (Polychaeta: Oweniidae) from the Antarctic Seas based upon type material. Proceedings of the Biological Society of Washington, 114: 403 413.
Paxton, H., 1986. Generic revision and relationships of the family Onuphidae (Annelida: Polychaeta). Records of the Australian Museum, 38: 1 74.
Petersen, M. E., 2000. 11. Family Sternaspidae Carus, 1863. In: Blake, J. A., B. Hilbig and P. V. Scott (eds.), Taxonomic Atlasof the benthic fauna of the Santa Maria Basin and Western Santa Barbara Channel, Volume 7. The Annelida Part 4.
Polychaeta: Flabelligeridae to Sternaspidae, pp. 311 336. Santa Barbara Museum of Natural History, Santa Barbara,
California.
Pettibone, M. H., 1953. Some scale-bearing polychaetes of Puget Sound and adjacent waters. 89 pp. University of Washington Press, Seattle.
Pettibone, M. H., 1954. Marine polychaete worms from Point Barrow, Alaska, with additional records from the North Atlantic and North Pacific. Proceedings of the United States National Museum, 103: 203 356.
Pettibone, M. H., 1956. Marine polychaete worms from Labrador. Proceedings of the United States National Museum, 105:
531 584.
Pettibone, M. H., 1957. North American genera of the family Orbiniidae (Annelida: Polychaeta), with descriptions of new
species. Journal of the Washington Academy of Sciences, 47: 159 167.
Pettibone, M. H., 1963. Marine polychaete worms of the New England Region. I. Aphroditidae through Trochochaetidae.
Bulletin of the United States National Museum, 227: 1 356.
Pettibone, M. H., 1966. Revision of the Pilargidae (Annelida: Polychaeta), including descriptions of new species andDeep-sea Benthic Polychaetes off Northern Honshu

191

redescription of the pelagic Podarmus ploa Chamberlin (Polynoidae). Proceedings of the United States National Museum, 118: 155 208.
Pettibone, M. H., 1970. Two new genera of Sigalionidae (Polychaeta). Proceedings of the Biological Society of Washington,
83: 365 386.
Pettibone, M. H., 1971. Revision of some species referred to Leptonereis, Nicon, and Laeonereis (Polychaeta: Nereididae).
Smithsonian Contributions to Zoology, (104): 1 53.
Pettibone, M. H., 1989. Revision of the aphroditoid polychaetes of the family Acoetidae Kinberg (=Polyodontidae Augener)
and reestablishment of Acoetes Audouin and Milne-Edwards, 1832, and Euarche Ehlers, 1887. Smithsonian Contributions to Zoology, (464): 1 138.
Quatrefages, A. de, 1865. Histoire naturelle des Annéles marins et d'eau dorce. Annélides et Gephyriens. Volume 1, 588 pp.
Librarie Encyclopédique de Rôret, Paris.
Ranzani, C., 1817. Beschreibung einer neuen Gattung Thalassema. Isis von Oken, Jena, 1: 1457 1461.
Rathke, H., 1843. Beiträge zur Fauna Norwegens. Nova Acta Academiae Caesarea Leopoldino-Carolinae Germanicum Naturae Curiosorum, 20: 1 264.
Reish, D. J., 1968. A biological survey of Bahia de los Angeles, Gulf of California, Mexico. II. Benthic polychaetous annelids. Transactions of the San Diego Society of Natural History, 15: 67 106.
Rullieŕ, F., 1963. Une petite collection d’Annélides polychètes de l’Ile Maurice. Bulletin de la Société Zoologique de France,
87: 471 481.
Rullieŕ, F., 1965. Contribution à la faune des Annélides polychètes du Dahomey et du Togo. Cahiers ORSTOM -Océanographie, Paris, 3: 5 66.
Salazar-Vallejo, S. I., 1986. Pilargidae (Annelida: Polychaeta) de Mexico: lista de especies, nueva especie y biografia. Cahiers de Biologie Marine, 27: 192 210.
Salazar-Vallejo, S. I., 1990. Redescriptions of Sigambra grubii Müller, 1858 and Hermundura tricuspis Müller, 1858 from
Brazil and designation of neotypes (Polychaeta: Pilargidae). Journal of Natural History, 24: 507 517.
Saphronova, M. A., 1991. Redescription of some species of Scionella Moore, 1903, with a review of the genus and comments
on some species of Pista Malmgren, 1866 (Polychaeta: Terebellidae). Ophelia, Supplement 5: 239 247.
Sars, M., 1851. Beretning om en i sommeren 1849 foretagen zoologisk Reise i Lofoten og Finmarken. Nyt Magazin for
Naturvidenskaberne, Oslo, 6: 121-211.
Seidler, H. J., 1924. Beiträge zur Kenntnis der Polynoiden, I. Archiv für Naturgeschichte, Berlin, 89A: 1 217.
Söderström, A., 1920. Studien über die Polychätenfamilie Spionidae. Inaugural-Dissertation. 286 pp. Almqvist and Wiksells,
Uppsala.
Southern, R., 1914. Clare Island Survay. Part 47, Archiannelida and Polychaeta. Proceedings of the Royal Irish Academy, 31:
1 160.
Southern, R., 1921. Polychaeta of the Chilka Lake and also of fresh and brackish waters in other parts of India. Memoirs of
the Indian Museum, 5: 563 659, pls. 19 31.
Støp-Bowitz, C., 1948. Polychaeta from the Michael Sars North Atlantic deep-sea expedition 1910. Report on the Scientific
Results of the Michael Sars North Atlantic Deep-Sea Expedition, 5: 1 91.
Sun, R. P. and D. J. Yang, 1987. Studies of Orbiniidae (Polychaeta) from the Yellow Sea and the East China Sea. Studia
Marina Sinica, (28): 151 168. (In Chinese.)
Takahashi, K., 1938. Polychaetous annelids of Izu Penisula. I. Polychaeta, collected by the “Misago” during the zoologicalsurvey around the Izu Peninsula. Science Reports of the Tokyo Bunrika Daigaku, Section B, 3: 193-221.
Tauber, P., 1879. Annulata Danica. En Kritisk revision af de i Danmark fundne Annulata Chaetognatha, Gephyrea, Balanoglossi, Discophoreae, Oligochaeta, Gymnocopa og Polychaeta. 144 pp. Reitzel, Kjøbenhavn.
Tebble, N. and S. Chambers, 1982. Polychaetes from Scottish waters. A. Guide to Identification, Part 1. Family Polynoidae.
Royal Scottish Museum Studies, 1982: 1 73.
Théel, H. J., 1879. Les Annélides polychètes des Mers de la Nouvelle-Zemble. Konglia Svenska Vetenskaps-Akademiens
Handlinger, Stockholm, 16: 1 75.
Treadwell, A. L., 1906. Polychaetous annelids of the Hawaiian Islands, collected by the steamer Albatross in 1902. Bulletinof the U. S. Fisheries Commission, 23: 1145 1181.
Treadwell, A.L., 1923. Polychaetous annelids from Lower California with descriptions of new species. American Museum
Novitates, 74: 1 11.
Treadwell, A.L., 1942. Polychaetous annelids from Lower California and the Philippine Islands in the collections of theAmerian Museum of Natural History. American Museum Novitates, 1172: 1 5.
Uchida, H., 1968. Polychaetous annelids from Shakotan (Hokkaido) I. The collection in 1967. The Journal of the Faculty of
Science, Hokkaido University, Series 6, Zoology, 16: 595 612.
Uebelacker, J. M.,1984a. Chapter 41: Family Lumbrineridae Malmgren, 1867. In: Uebelacker J. M. and P. G. Johnson (eds.),
Taxonomic guide to the polychaetes of the northern Gulf of Mexico, Volume 6. 45 pp. Barry A. Vittor and Associates,
Inc., Mobile, Alabama.

192

Minoru Imajima

Uebelacker, J. M.,1984b. Chapter 51: Family Ampharetidae Malmgren, 1867. In: Uebelacker J. M. and P. G. Johnson (eds.),
Taxonomic guide to the polychaetes of the northern Gulf of Mexico, Volume 7. 32 pp. Barry A. Vittor and Associates,
Inc., Mobile, Alabama.
Uschakov, P. V., 1950. Bristle-worms (Polychaeta) from the Okhotsk Sea. Issledovaniya Dal'novostochnykh Morei SSSR, 2:
140 234, 2 pls. (In Russian.)
Uschakov, P. V., 1955. Polychaeta of the far eastern seas of the U.S.S.R. Key to the Fauna of the U.S.S.R. No. 56, 445 pp.
Akademii Nauk SSSR, Moscow. (In Russian.)
Uschakov, P. V., 1965. Polychaeta of the far eastern seas of the U.S.S.R. Key to the Fauna of the U.S.S.R. No. 56, 419 pp.
Zoological Institute of the Academy of Sciences of the U.S.S.R., Jerusalem. (Translated by the Israel program for scientific translations.)
Uschakov, P. V., 1958. Novye i interesnye vidy mnogoschchetinkovykh chervei (Polychaeta) iz raiona iuzhnogo Sakhalina iiuzhnkh Kuril’skykh Ostrovov. Issledovaniya Dal'novostochnykh Morei SSSR, 5: 78 89.
Uschakov, P. V., 1972. Fauna of the USSR. Polychaetes. Volume 1. Polychaetes of the suborder Phyllodociformia of the polarbasin and northwestern part of the Pacific (Family Phyllodocidae, Alciopidae, Tomopteridae, Typhloscolecidae, andLacydoniidae). Akademii Nauk SSSR, New Series (102): 1 272. (In Russian.)
Uschakov, P. V., 1982. Fauna of the USSR, Polychaetes, Volume 2, Part 1. Polychaetes of the suborder Aphroditiformia of the
Arctic Ocean and the northwestern part of the Pacific (Familiy Aphroditidae and Polynoidae). Akademii Nauk SSSR,New Series (126): 1 272. (In Russian.)
Uschakov, P. V. and B.-L. Wu, 1962a. Polychaete worms of the Yellow Sea. VI. Supplement of erratic worms (Polychaeta,
Errantia). Studia Marina Sinica, (2): 110 133, 4 pls. (In Chinese and Russian.)
Uschakov, P. V. and B.-L. Wu, 1962b. Polychaete worms of the Yellow Sea. II. Fam. Chrysopetalidae, Glyceridae, and Nephthydidae (Polychaeta, Errantia). Studia Marina Sinica, (1): 1 28, 4 pls. (In Chinese and Russian.)
Uschakov, P. V. and B.-L. Wu, 1962c. Polychaete worms of the Yellow Sea. IV. The polychaetous annelids of the families
Syllidae, Hesionidae, Pilargiidae, Amphinomidae and Eunicidae (Polychaeta, Errantia). Studia Marina Sinica, (1):57 85, 3 pls. (In Chinese and Russian.)
Uschakov, P. V. and B.-L. Wu, 1979. Polychaeta Errantia of the Yellow Sea. Studia Marina Fauna, 3: i-viii, 1 137.
Verrill, A. E., 1873. Results of recent dredging expeditions on the coast of New England. American Journal of Science, Series
3, 5: 98 106.
Verrill, A. E., 1881. New England Annelida. Part 1. Historical sketch, with annotated lists of the species hitherto recorded.
Transactions of the Connecticut Academy of Arts and Science, 4: 285 324, 12 pls.
Verrill, A. E., 1885. Notice of recent additions to the marine Invertebrata of the northeastern coast of America, with descriptions of new genera and species and critical remarks on others. Proceedings of the United States National Museum, 8:
424 448.
Webster, H. E. and J. E. Benedict, 1887. The Annelida chaetopoda from Eastport, Maine. Reports of the United States FishCommission for 1885: 707 755, 8 pls.
Wesenberg-Lund, E., 1948. Maldanidae (Polychaeta) from West Greenland waters. Meddelelser om Grønland Kommissionen
for Videnskabelige Undersøgel Grønland, 134: 1 58.
Wesenberg-Lund, E., 1950. Polychaeta. The Danish Ingolf-Expedition, 4: 1 92, 10 pls.
Williams, S. J., 1987. Taxonomic notes on some Ampharetidae (Polychaeta) from Southern California. Bulletin of the Biological Society of Washington, (7): 251 258.
Wollebaek, A., 1912. Nordeuropaeiske Annulata Polychaeta. I. Ammocharidae, Amphictenidae, Ampharetidae, Terebellidae
og Serpulidae. Videnskapsselskapets Skrifter, 1. Mathematik-Naturvidenskabelig Klasse, 1911: 1 144, pls. 51.
Wu, B. L., R. Sun and M. Chen, 1980. Zoogeographical studies on Polychaeta from the Xisha Islands and its adjacent waters.
Acta Oceanogia Sinica, 2: 111 130. (In Chinese with English summary.)
Yang, D. J. and R. P. Sun, 1988. Polychaetous annelids commonly seen from the Chinese waters. 352 pp. Agricultural Press.,
Beijing. (In Chinese.)Zaks, I. G., 1922. A new polychaete species belonging to the family Ammocharidae: Myriochele oculata n. sp. Trudy Petrogradskogo obschcestva Estestvoispytatelei, 53: 171 174, figs. 1 3. (In Russian.)