New Records of Semiaquatic Species Marionina (Clitellata, Enchytraeidae) from Japan, with a Description of Marionina Biwaensis Sp

T. TORII

Turk J Zool 2012; 36(1): 15-24 © TÜBİTAK Research Article doi:10.3906/zoo-1003-111

New records of semiaquatic species Marionina (Clitellata, Enchytraeidae) from Japan, with a description of Marionina biwaensis sp. nov.

Takaaki TORII* IDEA Consultants Inc., 1334-5, Riemon, Yaizu-shi, Shizuoka Pref., 421-0212, JAPAN

Received: 07.03.2010

Abstract: Marionina biwaensis sp. nov. is described from Lake Biwa. Th is new species diff ers from all other Marionina species: lacking lateral chaetae completely and ventral chaetae in II, segment number 38-43, origin of dorsal blood vessel XV or XVI, spermathecal ampullae with irregular protuberances and chaetal distribution 1 per postclitellar bundles. In addition, the descriptions of M. coatesae, M. nevisensis, and M. riparia are augmented and recorded here for the fi rst time in Japan.

Key words: Clitellata, Enchytraeidae, Marionina, Marionina biwaensis sp. n., Japan

Introduction Lumbricillus lineatus (Müller, 1774); and Fridericia Th ere have been many studies of Enchytraeidae perrieri (Vejdovsky, 1878). Unidentifi ed Marionina in Asia. In China, for example, a total of 56 species species have also been recorded from riverbanks from 16 genera of Enchytraeidae have been hitherto (Torii and Ohtaka, 2007) and moist soils (Nakamura, recorded (Wang and Cui, 2007; Chen and Xie, 2008; 1980, 1986; Fujita, 1991). It is quite probable that Chen and Xie, 2009). many undescribed species are currently present in Japan. In Japan, 18 species from 8 genera have been previously recorded. Among these studies, published Th e species richness of Enchytraeidae is records mainly concern those from terrestrial considered to be greatly underestimated. Due to habitats (Nakamura, 1980, 1984, 1986, 1993; taxonomic diffi culties, a lack of modern identifi cation Schmelz et al., 2000; Nakamura et al. 2001, 2006); guides for most taxa, and few trained and practicing only a few are recorded from aquatic or semiaquatic systematists, enchytraeids have long been neglected sites (Yamaguchi, 1953; Coates and Ellis, 1981; and are rarely identifi ed, even to the genus (Martin Torii and Ohtaka, 2007). Of these aquatic and et al., 2008). Th is paper presents 1 new species of semiaquatic fi ndings, only 4 species of enchytraeids Marionina and gives new descriptions of 3 further have been identifi ed: Lumbricillus annulatus Eisen, species of Marionina, all recorded in Japan for the 1904; Lumbricillus nipponicus (Yamaguchi, 1937); fi rst time.

* E-mail: [email protected]

15 New records of semiaquatic species Marionina (Clitellata, Enchytraeidae) from Japan, with a description of Marionina biwaensis sp. nov.

Materials and methods duct, ne: nephridium, oe: esophagus, ph: pharyngeal All specimens were collected in Japan from pad, pg: pharyngeal glands, ppb: postpharyngeal the Shiga, Aichi, Shizuoka, Kanagawa, and bulb, sa: spermathecal ampulla, sf: sperm funnel, sv: Chiba prefectures. Living and fi xed specimens seminal vesicle, vd: vas deferens. were examined. Specimens were fi xed in either 10% formalin or 70% ethanol, sometimes aft er Results anaesthetization in low concentrations of ethanol. Specimens were dehydrated in a graded ethanol Genus Marionina Michaelsen, 1890 series, cleared in methyl salicylate, and mounted Marionina biwaensis sp. nov. whole onto slides in Canada balsam. Unless otherwise specifi ed in the descriptions, measurements refer to (Figure 1a-1h) whole-mounted specimens. Specimens examined in Type material. Holotype: 1 fully mature specimen, this study were deposited in the National Museum of Lake Biwa, Ohura, Nishiasai-cho, Shiga Prefecture, Nature and Science, Tokyo, Japan (NSMT). Japan, 35°27ʹ23ʺN, 136°06ʹ29ʺE, 14 October 2007, Abbreviations in the fi gures: (NSMT-An-404). Paratype: 6 mature and 3 immature specimens, same data as holotype, (NSMT-An-405). b: brain, c: coelomocyte, cc: chloragogen cell, co: collar of sperm funnel, dv: dorsal blood vessel, ecd: Other material: 5 mature specimens, same data as spermathecal ectal duct, end: spermathecal ental holotype.

(a) (h) (b) (c)

ph 20 μm

dv ppb sf (e) oe co pg

sa 100 μm 100 μm (f) (d) ecd

c ne (g) cc 50 μm 100 μm 100 m μ sa

10 μm

Figure 1. Marionina biwaensis sp. nov. from Lake Biwa, Japan (NSMT-An-404), all fi gures from living specimen: a) anterior body region (7 segments) of a mature specimen, body interior, dorsal view, almost coelomocytes omitted; b) same, lateral view (8 segments); c) segment XI-XIII, ventral view of a mature specimen, almost coelomocytes omitted; d) chaetae, bundle of IV, whole mount, side view; e) sperm funnel; f) spermatheca; g) same, dorsal view; h) coelomocyte.

16 T. TORII

Description of new materials. Body mostly conspicuous canal, funnel comprising one-third of transparent and partly opaque, due to intensely white anteseptal part. Whole anteseptale about one-third spots caused by accumulations of coelomocytes. or one-fourth of the total length of nephridium. Length 7-9 mm in vivo, 5-7 mm in fi xation; width Postseptale without a distinct transition to eff erent at clitellum 110-130 μm. Segments 38-43 (holotype: duct. Postclitellar nephridia from 14/15 onward. 41). Chaetae straight, ental hook distinct (Figure Testis paired in XI (10/11). Seminal vesicle well 1d), preclitellar chaetae 32.5-37.5 μm long (mostly developed, single, extending usually from 10/11 to 33-35 μm), 2.7-3.1 μm wide, postclitellar chaetae 11/12. Sperm funnel (Figures 1c and 1e) 110-130 μm 27.5-33 μm long (mostly 28-31 μm), 2.0-2.4 μm long and 40-70 μm wide, collar narrower than funnel wide. Chaetal formula 0-0:0,2(1)-1. Lateral chaetae body. Vas deferens narrowing from glandular part to absent, ventral chaetae absent in II. Ventral chaetae male pore, moderately long; loose or tight irregularly absent in XI, XII, and XIII in mature specimens. coiled in XII. Th ick bundles of fi liform spermatozoa Preclitellar ventral chaetae of nearly equal length. present in front of sperm funnel. Penial bulb Postclitellar chaetae slightly thinner and shorter than extension approximately one-third of body width, in preclitellar region. Chaetae equal-sized within 35-50 μm long. Vas deferens runs along anterolateral a bundle. Prostomium rounded. Head pore at 0/1. part of bulb. Bursal slit 23-25 μm long. Copulatory Epidermal gland cells inconspicuous. glands absent. No egg sac. One or two oocytes in XIII. Clitellum extending over XII–1/2XIII, cells not Paired spermathecal pores midlateral at 4/5, tall, with slightly irregular shape and distribution. spermathecae attached to esophagus in posterior of Brain (Figures 1a and 1b) in I–II, dorsoventrally V (Figures 1a and 1b). Ampulla (Figures 1f and 1g) compressed and narrow, anterior part convex, more or less oval with some small, irregular lobes posterior part deeply incised, approximately twice along ectolateral edge, but only oval or marginally as wide as anterior part, length 70-100 μm, posterior lobed in submature specimens; ental duct short and width 50-65 μm. Dorsal blood vessel originating not separate from unlobed ental part of ampulla. near 15/16 or 16/17; anteriorly bifurcating behind Sperm head penetrate in small lobes, tails extending pharyngeal pad in III, branches extending near side into cavity of ampulla. Ectal duct length less than of brain and uniting as ventral vessel (Figure 1a); ampulla plus ental duct; ectal duct surrounded by blood colorless. Pharyngeal pad slightly thickened. small, irregular gland cells; a few glands near the No esophageal appendages. Pharyngeal glands 3 ectal pore longer than the rest. pairs, the anterior 2 pairs merging dorsally at 4/5 and Habitat. Th e present specimens were collected 5/6, respectively, the third pair elongate and without from freshwater sites (northern basin of Lake Biwa), dorsal connection at 6/7, ventral lobes in IV (small) near-shore shallows, and the sandy substrate zone. and V. Paired postpharyngeal bulbs in III. Transition Th e species was the dominant oligochaete in the of esophagus to intestine gradual, no esophageal or intestinal diverticula. surface layer (5-10 cm) of interstitial sand and gravel at the locality. Coelomocytes (Figure 1h) fl attened oval, nucleate, approximately 14-18 μm long in vivo, 1.0- Etymology. Th e name “biwaensis” was given aft er 1.5 times as long as wide, fi lled with distinct, globular the type locality of the species, Lake Biwa, Japan. vesicles, but sometimes showing a granulated Remarks. M. biwaensis resembles the Hungarian cytoplasm aft er fi xations. In living specimens the M. scintillans Boros and Dózsa-Farkas, 2008; the vesicular cytoplasm appears whitish under incident Chinese M. seminuda Xie and Rota, 2001; the light. Chloragocytes dense from V or VI, slightly Holarctic M. subterranea (Knöllner, 1935); and the smaller than coelomocytes. Preclitellar nephridia 2 northern European M. glandulifera (Jansson, 1960). or 3 pairs at 7/8 to 9/10 or 8/9 to 9/10, 80-100 μm Th ese species also lack lateral chaetae and have long, sometimes 1 or both nephridia absent at 9/10. opaque coelomocytes, a pharyngeal pattern of the Anteseptal part of nephridium large, with funnel and dorsal blood vessel bifurcation, and a posteriorly

17 New records of semiaquatic species Marionina (Clitellata, Enchytraeidae) from Japan, with a description of Marionina biwaensis sp. nov.

incised brain (Knöllner, 1935; Jansson, 1960; Xie Coates and Stacey, 1993: 411, Figure 12; Healy and and Rota, 2001; Boros and Dózsa-Farkas, 2008). Coates, 1997: 93; Rota et al., 2003: 509-510. Th e diff erences between M. biwaensis and M. Material examined. Seven mature specimens, scintillans, M. seminuda, M. subterranea, and M. Sagami River, Hiratsuka-shi, Kanagawa Prefecture, glandulifera are as follows (see also Table): segment Japan, 35°19ʹ17ʺN, 139°21ʹ59ʺE, 2 November 2008 number (38-43 in M. biwaensis, fewer in the other (one of them has been registered as NSMT-An-406); 4 species), origin of dorsal blood vessel (XV or XVI 5 mature specimens, Kiso River, Gomyo-cho, Yatomi- in M. biwaensis, XII or XIII in the other 4 species), shi, Aichi Prefecture, Japan, 35°07ʹ16ʺN, 136°42ʹ16ʺE, morphology of the spermathecal ampullae (with 11 September 2008; 3 mature specimens, Kikuchi irregular protuberances in M. biwaensis; without River, Yokoshima-cho, Tamana-shi, Kumamoto protuberances, lobes, or diverticula in the other 4 Prefecture, Japan, 32°53ʹ27ʺN, 130°32ʹ14ʺE, 15 June species), and chaetal distribution (1 per postclitellar 2008. bundles in M. biwaensis, 2 per postclitellar bundles in Brief description of new materials. Dimensions the other species). in preserved materials: length 4.3-6.5 mm, width Japanese name. Biwa-himemimizu. at clitellum 150-220 μm. Segments 35-45. Chaetae Marionina coatesae Erséus, 1990 straight with ental hook, preclitellar chaetae 40-55 μm long, 3.4-3.7 μm wide, postclitellar chaetae 54-57 (Figures 2a and 2b) μm long, 3.5-3.7 μm wide. Two chaetae per bundle Marionina coatesae Erséus, 1990: 318-319, Figure throughout body. Ventral chaetae of XII missing. 26A-G; Erséus et al., 1990: 117-118, Figure 6A-B; Dorsal blood vessel originating posterior of XIII;

Table. Comparison of marionine species lacking all lateral chaetae and “marionine” type of anterior dorsal blood vessels.

M. scintillans M. seminuda M. subterranea M. glandulifera M. biwaensis sp. nov. Boros and Dózsa-Farkas, Xie et Rota, 2001 (Knöllner, 1935) (Jansson, 1960) 2008

Body length (mm) 7-11 (in vivo), 5-7 2.0-2.9 (in vivo), 1.6-2.2 3-5 (in vivo), 2-3 3-3.2 1.5-2.3

Segment number 38-43 19-23 23-27 28-30 24-26

Chaetal length anterior (μm) 32.5-37.5 20-22 25-30 15-20 20-25 posterior (μm) 27.5-33 28-32 22-27 20-25

Chaetal distribution 0-0:0,2(1)-1 0-0:(0),2-2 0-0:0,2-(1),2 0-0:2-2 0-0:2-2

Dorsal blood vessel origin XV or XVI XII XIII XIII XIII

2 pairs merging dorsally 3 pairs, anterior 2 pairs 3 pairs, anterior 2 pairs Pharyngeal gland and small secondary 3 pairs merging dorsally 3 pairs merging dorsally merging dorsally merging dorsally glands in VI

Postpharyngeal bulb Present Present Present Absent Present

Seminal vesicle Single Absent 1 pair ? or single* ? or single*

With irregular Without protuberances, Without protuberances, Without protuberances, Without protuberances, Spermathecal ampulla protuberances lobes and diverticula lobes and diverticula lobes and diverticula lobes and diverticula

Along length of duct, Along length of duct, Along length of duct, 1 Glandular cells around 1 or 2 larger glands larger glands originating or 2 glands originating Only at the ectal pores Only at the ectal pores spermathecal duct originating around ectal around ectal pores around ectal pores pores

Boros and Dózsa-Farkas, Kossmagk-Stephan, 1983; Kossmagk-Stephan, 1983; References Present study Xie and Rota, 2001 2008 *Coates, 1983 *Coates, 1983

18 T. TORII

rings scattered in wall and loose bundles in ampullar (a) (b) lumen. Ectal duct surrounded by glandular lobes of b varying and irregular size; some glands near the ectal pore longer than the rest. ph Habitat. Th e present specimens were collected sa from the estuary region of interstitial sand and gravel bottom. Remarks. Th is is the fi rst record of Marionina oe coatesae from Japan. Healy and Coates (1997) saw a copulatory gland in only one specimen, but most Japanese materials have a copulatory gland. Other ne morphological characters of the present specimens agreed closely with those in the original description (Erséus, 1990) and redescription (Healy and Coates, c sv 1997). Distribution. Hong Kong (Erséus, 1990), sf northern China (Erséus et al., 1990), Japan (present study), and western Australia (Coates and Stacey, 1993; Rota et al., 2003). Japanese name. Kôtesu-himemimizu. Marionina nevisensis Righi and Kanner, 1979 100μm (Figures 3a-3d) 100μm Marionina achaeta nevisensis Righi and Kanner, Figure 2. Marionina coatesae from Sagami River, Japan: a) 1979: 65-66, Figures 52-57. anterior body region (13 segments) of a mature specimen, from whole mount, body interior, dorsal Marionina nevisensis Coates, 1983: 825-826, view, almost coelomocytes and vessel systems omitted; Figures 6-8; Coates, 1990: 32-33, Figure 10; Erséus, b) same, anterior 6 segments. 1990: 320-321, Figures 27A-B; Erséus et al., 1990: 118-119, Figure 7; Coates and Stacey, 1993: 411-413, Figure 13; Healy and Coates, 1997: 93-95. anteriorly bifurcating behind pharyngeal pad in IV, branches extending near side of brain and uniting Material examined. Th ree mature specimens, as ventral vessel; blood colorless. Pharyngeal glands Lake Biwa, fl ow region of Ado River, Adogawa-cho, 3 pairs, the anterior 2 pairs merging dorsally at 4/5 Shiga Prefecture, Japan, 35°18ʹ57.2ʺN, 136°04ʹ31.1ʺE, and 5/6, respectively, the third pair elongate and 23 August 2006 (these specimens have been registered without dorsal connection at 6/7, ventral lobes in V as NSMT-An-407 with one other species on the and VI. Postpharyngeal bulbs not visible. Transition same slide); 5 mature specimens, Takatoki River, of esophagus to intestine gradual. Coelomocytes head works, Shiga Prefecture, Japan, 35°29ʹ38.4ʺN, fl attened oval, nucleate, not granulated. Seminal 136°15ʹ14.5ʺE, 18 January 2008. vesicle developed, unpaired laterally, sometimes Brief description of new materials. Body mostly extending forward into IX. Sperm funnel 2 to 3 times transparent, with intensely white spots caused as long as wide, collar width equal to funnel. Vas by accumulations of coelomocytes. Appearance deferens narrowing from glandular part to male pore, nematode-like because of elongated and nonchaetal moderately long; loose or tight irregularly coiled habitus. Preclitellar cuticle 3-5 μm thick. Live length in XII. Penial bulb small and compact. Copulatory 7-11 mm, fi xed length 5-7 mm. Dimensions in gland present in XIV. Spermathecal ampulla preserved material: width at clitellum 110-130 μm. irregularly round or oval. Sperm arranged in distinct Segments 35-38. No chaetae. Dorsal blood vessel

19 New records of semiaquatic species Marionina (Clitellata, Enchytraeidae) from Japan, with a description of Marionina biwaensis sp. nov.

(a) (b)

sa b

oe sa ph

20μm

ne dv

(d) pg sa sv co oe

sf sf

200μm

50μm 100μm

Figure 3. Marionina nevisensis from Lake Biwa, Japan: a) anterior body region (12 segments) of a mature specimen, from whole mount, body interior, dorsal view, almost coelomocytes omitted; b) same, anterior 6 segments; c) spermatheca, from living specimen; d) sperm funnel, from living specimen. originating in posterior of XIV; anteriorly bifurcating Collar width equal to funnel. Vas deferens moderately behind pharyngeal pad in IV, branches extending long; loose or tight, irregularly coiled in XII. Penial near side of brain and uniting as ventral vessel; blood bulb small and compact, 37-42 μm wide. Vas deferens colorless. Pharyngeal glands 3 pairs, the anterior 2 runs along anterolateral part of bulb. Copulatory pairs merging dorsally at 4/5 and 5/6, respectively, glands absent. Spermathecal ampulla irregularly round the third pair elongate and without dorsal connection or oval. Sperm as loose bundles in ampulla. Ectal duct at 6/7. Transition of esophagus to intestine gradual. surrounded by small, irregular gland cells; some glands Coelomocytes fl attened oval, nucleate, their granular near the ectal pore longer than the rest. cytoplasm appears whitish, fi lled with distinct, globular Habitat. Th e present specimens were collected vesicles. Testes paired in XI (10/11). Seminal vesicle from freshwater sites and were present in the surface developed, unpaired, sometimes extending forwards layer (5-10 cm) of the interstitial sand and gravel into VII. Sperm funnel 4 to 6 times as long as wide. bottom.

20 T. TORII

Remarks. Th is is the fi rst record of Marionina Distribution. Palearctic: Caribbean, Hong Kong, nevisensis from Japan. In total, 3 species of Marionina northern and southern China, Canada (Coates, are known as being without all chaetae: M. achaeta 1990), Japan (present study), Australia (Coates, 1990; Lasserre, 1964; M. arenaria Healy, 1979; and M. Coates and Stacey, 1993; Healy and Coates, 1997); all nevisensis Righi and Kanner, 1979. Th ey are known specimens collected from marine regions except in from upper intertidal and supralittoral beach Japan. habitats (Coates, 1983). In Coates (1983), glandular Japanese name. Subesube-himemimizu. proliferations were present on the ventral nerve cord at the ganglia of XIII, XIV, and frequently XV, but Marionina riparia Bretscher, 1899 these are not present or not visible in the Japanese (Figures 4a-4f) materials. Other taxonomic characters examined in See Rota (1994) for synonymy. the present Japanese specimens agree well with the previous descriptions (Coates, 1983, 1990), but the Material examined. One mature specimen, Tega freshwater habitat of the present specimens contrasts bog, Tobarishinden, Kashiwa-shi, Chiba Prefecture, with all previous records of the species from marine Japan, 35°51ʹ21.8ʺN, 140°00ʹ24.6ʺE, 24 May 2006 or brackish habitats. (this specimen has been registered as NSMT-

(a) (b) b ph b (c) (d)

oe ecd pg

sa sa oe end sf c 50μm (e)

100μm

(f) vd sf

200μm 50μm

20μm

Figure 4. Marionina riparia from Lake Ippeki, Japan: a) anterior body region (13 segments) of a mature specimen, from whole mount, body interior, dorsal view, almost coelomocytes and vessel systems omitted; b) same, anterior 7 segments; c) spermatheca, from live specimen, sperm fi brous pattern; d) same, sperm mass pattern; e) sperm funnel, from living specimen; f) chaetae, bundle of IV, whole mount, side view.

21 New records of semiaquatic species Marionina (Clitellata, Enchytraeidae) from Japan, with a description of Marionina biwaensis sp. nov.

An-408); 1 mature specimen, Lake Ippeki, Yoshida, Christensen, 1959; Rota and Healy, 1994), except for Ito-shi, Shizuoka Prefecture, Japan, 34°55ʹ28.6ʺN, the cutaneous glands, which are not so colored, and 139°06ʹ41.6ʺE, 23 March 2008; 10 mature specimens, for the sperm in the spermathecal ampulla, formed as Tenryu River, Ryuyoh-cho, Shizuoka Prefecture, fi bers (Figure 4c) and not as coils (Figure 4d) in some Japan, 34°39ʹ39.5ʺN, 137°47ʹ51.2ʺE, 9 February fi xed specimens. 2008; 4 mature specimens, Lake Biwa, Hayasaki, Distribution. Palearctic: Great Britain, Ireland, Nagahama-shi, Shiga Prefecture, Japan, 35°25ʹ10.2ʺN, Sweden, Finland, Denmark, Germany, Spain, France, 136°12ʹ00.2ʺE, 25 May 2006. Switzerland, Austria, Italy, Poland, Czechoslovakia, Brief description of new materials. Body slightly Romania, Lebanon (Rota and Healy, 1994), reddish. Live length 9-11 mm. Dimensions in China (Wang et al., 1999), Japan (present study), preserved material: length 6-8 mm, width at clitellum and northern Africa: Morocco, Algeria, Tunisia 200-220 μm. Segment 25-34. Chaetae sigmoid, (Martinez-Ansemil and Giani, 1987). preclitellar chaetae 43-47 μm long, 2.9-3.1 μm wide. Japanese name. Mizu-himemimizu. Postclitellar chaeta 42-45 μm long, 2.8-2.3.0 μm wide. Chaetal formula 3,4-3,4:3,4,5-3,4. Clitellum over XII–1/2XIII, cells tall, with irregular shape and Discussion distribution. Dorsal blood vessel originating near Enchytraeidae are diversifi ed in various 13/14; anterior blood system “lumbricilline” (Coates environments, especially in terrestrial and and Ellis, 1981). Blood light-reddish. Pharyngeal semiaquatic regions in central Japan. In this paper, glands 3 pairs, either separate or with a very narrow 4 species of Marionina from semiaquatic habitats are connection dorsally, at 4/5 and 5/6, 6/7, respectively. described, although some hitherto undetermined Transition of esophagus to intestine gradual. enchytraeid species belonging to other genera were Coelomocytes fl attened oval, nucleate, fi nely found as well. Enchytraeidae oft en appear as the granulated. Testis paired in XII (10/11). No seminal dominant family and most common oligochaetes vesicle. Sperm funnel barrel-shaped, narrower in the surface layer of interstitial sand and gravel in toward the end, 2 to 3 times as long as wide. Collar semiaquatic regions, a habitat susceptible to drastic rather indistinct, narrower than funnel. Vas deferens changes due to climate change, aridifi cation, and moderately long, loose or tight, irregularly coiled water pollution. Further studies of enchytraeid in XII. Spermathecal ectal duct with 1 or 2 glands biodiversity in semiaquatic regions are therefore very at ectal pore. Ental duct of spermatheca attached to important for environmental assessments in Japan. esophagus in posterior of V. Th e connection between It is noteworthy that M. nevisensis is fi rst spermatheca and esophagus rather thin. Ampulla recorded from Japan in a freshwater region. M. fusiform or oval with thin walls. Sperm arranged nevisensis has been known from a marine littoral in parallel in ampullar lumen, or arranged in 1 to 4 habitat on the Pacifi c coast. Timm and Vvedenskaya rings at the basis of ampulla. (2006) reported a similar case on the Kamchatka Peninsula. Additionally, 4 species of marine-origin Habitat. Th e present specimens were collected Enchytraeidae (Lumbricillus pagenstecheri (Ratzel, from freshwater sites (Tega bog, Lake Biwa, and 1869), Lumbricillus arenarius (Michaelsen, 1889), Lake Ippeki) and brackish sites (Aburaga-fuchi), Marionina charlottensis Coates, 1980, and Marionina salinity 0.1-0.5%. Th e specimens were present in the klaslisharum Coates, 1983) were recorded in interstitial sand and silt bottom, 1-40 cm of water freshwater Lake Kurilskoe. It is quite probable that depth. the Japanese population is in the middle stages of Remarks. Th is is the fi rst record of Marionina diff erentiation due to geographical isolation, although riparia from Japan. Th e present materials agree well the Japanese and the other regional populations are with the previous descriptions (e.g. Nielsen and of the same lineage.

22 T. TORII

In this paper, Marionina biwaensis sp. nov. is Acknowledgements described as characterized by the complete lack I am grateful to Dr. Rüdiger M. Schmelz of lateral chaetae and ventral chaetae in II, and (Universidad de A Coruña, Fac. Ciencias, Spain) comparison of marionine species lacking all lateral and Dr. Akifumi Ohtaka (Hirosaki University, chaetae and “marionine” types of anterior dorsal Aomori, Japan) for reading and making suggestions blood vessels (Table). Together with M. coatesae about this manuscript. I would also like to thank and M. nevisensis, M. biwaensis belongs to a group Dr. Yoshikazu Takashima (Hokkaido) for providing of species distinguished by a pharyngeal bifurcation me with valuable information about references, and of the dorsal blood vessel. Th is group may be anonymous reviewers for their eff ort and valuable monophyletic within Marionina. suggestions to improve the manuscript. Th anks are M. coatesae, M. nevisensis, and M. riparia are also due to Ms. Masako Masuda (Japan) for assistance recorded for the fi rst time in Japan. in the fi eld.

References

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