Revision of the Scaleworm Genus Eulagisca Mcintosh (Polychaeta: Polynoidae) with the Erection of the Subfamily Eulagiscinae and the New Genus Pareulagisca

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 110(4):537-551. 1997.

Revision of the scaleworm genus Eulagisca Mcintosh (Polychaeta: Polynoidae) with the erection of the subfamily Eulagiscinae and the new genus Pareulagisca

Marian H. Pettibone

Department of Invertebrate Zoology, National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560, U.S.A.

Abstract.—A new subfamily of Polynoidae, Eulagiscinae, is established for Eulagisca Mcintosh, to include the type species E. corrientis, E. gigantea Mon- ro, E. puschkini Averincev, and two new species, E. uschakovi and E. macnabi, all from the Antarctic and Subantarctic regions, and the new genus Pareula- gisca for P. panamensis (Hartman), new combination, from the central Pacific Panama region.

Mcintosh (1885) described Eulagisca rows of papillae; prominent conical facial corrientis from deep water in the South At- tubercle on the upper Up, ventral to the cera- lantic and South Indian Oceans. Monro tophore of the median antenna; tentaculo- (1939) described some large polynoids phores of segment I, lateral to the prosto- from the deep Antarctic as E. gigantea. mium, with projecting digitiform acicular Hartman (1939) added E. panamensis from lobe and group of long setae; and 15 pairs shallow water of Pacific Panama, and Av- of elytra (16 pairs in P. panamensis). erincev (1972) added E. puschkini from In addition to the collections in the Na- shallow water in the Antarctic Davis Sea. tional Museum of Natural History, Smith- The prostomia resemble those of the mem- sonian Institution (USNM), the specimens bers of the subfamilies Lepidonotinae Wil- covered herein are deposited in the following Natural History ley, 1902 and Lepidastheniinae Pettibone, Museums: Museum, London Natural History Museum 1989, by having the lateral antennae insert- (BMNH); of Los Angeles County (LACM-AHF): ed on anterior extensions, on the same level Zoological Institute Academy of Sciences, as the ceratophore of the median antenna, Leningrad (ZIASL); Zoological Museum, or subterminally, without distinct cerato- Hamburg (ZMH): Zoologisk Museum, phores. The parapodia differ from those of Universitet i Oslo (ZMUO). both subfamilies by having the notopodia and neuropodia well developed, with projecting digitiform acicular lobes, rather than Family Polynoidae Kinberg, 1856 Eulagiscinae, new subfamily projecting acicular lobes lacking. The elytra of some specimens, identified as E. gigan- Diagnosis.—Body elongate, with seg- tea, and some new material, showed some ments moderate in number (up to 41). El- differences and are referred to two new spe- ytra and bulbous elytrophores 15 (Eulagis- cies: E. uschakovi and E. macnabi. ca) or 16 (Pareulagisca) pairs, on segments Some distinctive characters of species of 2, 4, 5, 7, alternate segments to 23, 26, 29, Eulagisca, and not shown by E. panamen- 32 (33). Elytra large, oval, with or without sis, referred herein to the new genus Pareu- papillae, microtubercles, macrotubercles lagisca, include the following: long palps and globular vesicles or papillae. Dorsal of the prostomium with six longitudinal cirri, on non-elytrigerous segments, with 538 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON cylindrical cirrophores posterodorsal to no- alternate segments to 23, 26, 29, 32, 33. topodia, with distal styles; dorsal tubercles, Elytra without tubercles and border or surface papillae (Fig. in line with elytrophores, bulbous or indis- 5G). Prostomium with long palps, without 6 longitudinal tinct. Prostomium bilobed, with paired rows of papillae (Fig. 5A). Tentaculo- palps and three antennae; median antenna phore of segment I without projecting with large ceratophore in anterior notch of acicular lobe, with single seta (Fig. 5A); prostomium, with distal style; lateral anten- facial tubercle weakly developed. Noto- nae inserted terminally or subterminally on setae of two kinds: 1) stout, acicular, anterior extensions of prostomium, without smooth or with faint spinous rows (Fig. distinct ceratophores; two pairs of eyes. 5D); and 2) slender, capillary, smooth or First or tentacular segment not visible dor- with few spines (Fig. 5E) Pareulagisca, n.gen.: P. panamensis sally; tentaculophores lateral to prostomi- (Hartman 1939), n. comb. (Fig. 5A-G) um, each with dorsal and ventral tentacular - Elytra 15 pairs, on segments 2, 4, 5, 7, cirri, similar to median antenna, with or alternate segments to 23, 26, 29, 32. El- digitiform acicular lobe nu- without and ytra variable. Prostomium with long merous or single setae; with or without dis- palps with 6 longitudinal rows of papil- tinct facial tubercle on upper lip. Second or lae (Fig. 2A) Tentaculophores of seg- buccal segment with nuchal fold, first pair ment I with projecting digitiform acicu- of elytra and elytrophores, biramous para- lar lobe and group of long setae; with podia, and ventral buccal cirri, longer than prominent conical facial tubercle (Fig. 2A). Notosetae of one kind, stout, with following ventral cirri. Biramous parapodia long spinous regions, tapering to blunt with both noto- and neuropodia with prom- tips (Figs. 1C, 2C, 3C, 4D) inent projecting digitiform acicular lobes Eulagisca Mcintosh 1885 2 and numerous noto- and neurosetae. Noto- 2. Neurosetae tapering to sharp tips (Fig. setae of one kind, stout (Eulagisca) or two 3D; Averincev 1972, pi. 13:9-11) 3 kinds: stout and slender, capillary (Pareu- - Neurosetae tapering to slightly hooked lagisca). Ventral cirri short, tapering. Phar- tips (Figs. ID, 2D, 4E) 4 Elytra with sharply pointed tubercles and ynx with two pairs of jaws and eleven pairs 3. globular vesicles with few to numerous of dorsal and ventral papillae (not examined terminal spines (Fig. 3E) in Pareulagisca). Pygidium with pair of ... E. uschakovi, new species (Fig. 3A-E). anal cirri. - Elytra without border or surface papillae; Remarks.—The prostomia of the Eulag- with numerous microtubercles on anterior iscinae agree with the Lepidastheniinae and part (Averincev 1972, pi. 13:8)

the Lepidonotinae in having the lateral an- . . E. puschkini Averincev, 1972 (pi. 13:8-12). Elytra with border papillae (Figs. IE, 4F) tennae inserted terminally or subterminally, 4. 5 without distinct ceratophores, and disagree- - Elytra without border papillae; surfaces ing with the Harmothoinae having distinct with conical microtubercles, larger conical ceratophores and inserted ventrally. The nu- tubercles, and globular vesicles (Fig. 2F) chal folds on the second or buccal segment .... E. gigantea Monro, 1939 (Fig. 2A-F) in Eulagiscinae are distinctive. The bira- 5. Surface of elytra with conical papillae on mous parapodia, with both rami having anterior part (Fig. IE) prominent digitiform acicular lobes and nu- . . E. corrientis Mcintosh, 1885 (Fig. 1A-E) - Surface of elytra with conical microtuber- merous noto- and neurosetae in Eulagisci- cles and larger spiny tubercles (Fig. 4F) nae, differ from the other subfamilies. .... E. macnabi, new species (Fig. 4A-F).

Genus Eulagisca Mcintosh, 1885 Key to the Genera and Species of the — Subfamily Eulagiscinae Type species. Eulagisca corrientis Mc- intosh, 1885, by monotypy. Gender, femi-

1. Elytra 16 pairs, on segments 2, 4, 5, 7, nine. VOLUME 110, NUMBER 4 539

Diagnosis. —Body rather large, wide, (1921:45), Mcintosh's choice of Eulagisca flattened, slightly tapering anteriorly and for E. corrientis was a poor one, since it more so posteriorly, with segments up to differs in so many ways from Lagisca, in- 41. Elytra and bulbous elytrophores 15 cluding the type of the prostomium, para- pairs, on segments 2, 4, 5, 7, alternate seg- podia, and elytra. ments to 23, 26, 29, 32. Elytra large, covering dorsum, easily detached, with or with- Eulagisca corrientis Mcintosh, 1885 out microtubercles, macrotubercles, bul- Fig. 1 bous vescicles, and fringes of papillae. Dor- Eulagisca corrientis Mcintosh, 1885:91, sal cirri with cylindrical cirrophores and Fig. 1, pi. 13:fig. 4, pi. 7A:figs. 3, 4, long styles on non-elytrigerous segments; Not Eulagisca corrientis.—Benham, 1921: dorsal tubercles bulbous. Prostomium bi- 43, pi. 6:figs. 36-38, pi. 7:figs. 39-42. lobed, with paired long palps each with six [?] Eulagisca corrientis.—Monro, 1930:48, longitudinal rows of papillae, and three an- fig. lla-e (no elytra); 1936:90 (no ely- tennae; median antenna with ceratophore in tra). anterior notch of prostomium, with long Not Harmothoe corrientis (?). —Augener, style; lateral antennae inserted on anterior 1932:19. (= Harmothoe sp.). extensions of prostomium, on same level as Not Eulagisca corrientis.—Stiller, 1996:30, median antenna, without distinct cerato- pi. 4:figs. 1-5. phores; eyes 2 pairs, large, laterally on middle of prostomium, nearly contiguous. First Material examined.—South Atlantic: off or tentacular segment indistinct dorsally; east coast of South America, south of Buen- tentaculophores lateral to prostomium, each as Aires, 37°17'S, 53°52'W, 1097 m, green with projecting digitiform acicular lobe, sand, Challenger Sta. 320, 14 Feb 1876, group of setae, and long dorsal and ventral syntype (BMNH 1885.12.1.71). South In- tentacular cirri. Prominent conical facial tu- dian Ocean: Between Kerguelen and Heard bercle ventral to ceratophore of median an- Islands, 52°04'S, 71°22'E, 274 m, coarse tenna. Second segment with semilunar nu- gravel, Challenger Sta. 150, 2 Feb 1874, chal pad and nuchal fold covering posterior syntype (BMNH 1885.12.1.83). part of prostomium, first pair of elytro- Remarks. —The species is somewhat phores, biramous parapodia, and long ven- doubtful, since it was based on two syn- tral buccal cirri lateral to ventral mouth. types, a smaller complete specimen but Parapodia biramous, with numerous golden without elytra, from Challenger Sta. 320, setae. Notopodia well developed, with and an anterior fragment of a larger speci- prominent digitiform acicular lobes; neu- men with two elytra, from Sta. 150. There ropodia with long digitiform presetal acic- is a possibility that the two syntypes belong ular lobes and shorter rounded postsetal to different species but the type material lobes. Notosetae numerous, stouter than does not allow one to clarify this point. neurosetae, with long spinous regions, ta- Monro (1930:48) examined Mcintosh's pering to blunt tips; neurosetae more nu- type specimens and referred some speci- merous and finer than notosetae, with long mens from off the South Shetlands to E. spinous regions, tapering to rather long bare corrientis, but without elytra and thus tips. Ventral cirri short, subulate. Pharynx doubtful. Monro (1936:90) added a record with two pairs of chitinous jaws and 11 from off the Falkland Islands, but the spec- pairs of dorsal and ventral papillae. Pygid- imen also lacked elytra. Stiller (1996:30) ium with pair of anal cirri. Nephridial pa- added E. corrientis from the eastern Wed- pillae short, cylindrical, beginning on seg- dell Sea. It does not agree with the two syn- ment VI. types in the BMNH. Remarks. —As indicated by Benham The specimens described as E. corrientis —

540 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON by Benham (1921:43) appear to be a mix- er rounded postsetal lobes (Fig. IB; Mcin- ture of Eulagisca species and need to be tosh 1885: fig. 1). Notosetae numerous, reexamined. The specimen that Augener forming radiating bundle of four lengths, (1932:19) questionably referred to Har- slightly stouter than neurosetae, short, mothoe corrientis was examined from the slightly curved and longer, straight, with

Olso Museum (ZMUO); it is in poor con- long spinous regions, tapering to short bare dition, the prostomium is harmothoid and tips (Fig. 1C; Mcintosh 1885: pi. 7A fig. different from that of Eulagisca; it is re- 3). Neurosetae numerous, forming fan- ferred to Harmothoe sp. shaped bundle, with long spinous regions, Description.—Incomplete syntype with tapering to slightly hooked bare tips; upper 19 segments, 29+ mm long and 24 mm ones more slender and upper few with del- wide with setae. Smaller complete syntype icate secondary tooth (Fig. ID; Mcintosh with 35 segments, 34 mm long and 16 mm 1885: pi. 7A fig. 4). Ventral cirri slender, wide with setae. Dorsum with brown pig- with few papillae, tapering, extending mentation. Elytra and bulbous elytrophores slightly beyond basal part of neuropodia

15 pairs, in usual arrangement (Mcintosh (Mcintosh 1885: fig. 1). 1885: pi. 13, fig. 4). Elytra subreniform, Distribution.—South Atlantic and South surface with soft conical papillae and long Indian Oceans, in 274-1097 meters. delicate clavate papillae near posterior and lateral borders (Fig. IE). Dorsal cirri with Eulagisca gigantea Monro, 1939 cylindrical cirrophores and papillate styles Fig. 2 extending beyond setae; dorsal tubercles Eulagisca gigantea Monro, 1939:103 nodular. (part), Fig. 4a-d, f-h (not e), Sta. 30 Bilobed prostomium wider than long, only.—Uschakov, 1962:174 (part), Sta. with long papillate palps and three anten- 283, pi. 9E, G, H (not F); 1966:174 nae; median antenna with large ceratophore (part), Sta. 282, pi. 9E, G, H (not F) (En- in anterior notch, with style missing; lateral glish translation). —Hartman, 1967:21 antennae inserted on anterior extensions of (part, not Sta. 1003).—Not Averincev, prostomium, with rather long styles with 1972:131 (=E. uschakovi, new spe- subterminal enlargements and filamentous cies).—Not Desbruyeres, 1976:85, pi. 1: tips; two pairs of large closely approximat- figs, a-c (= Eulagisca, new species?). ed eyes on lateral sides; tentaculophores lat- Hartmann-Schroder & Rosenfeldt, 1990: eral to palps and prostomium, each with 91 (part), pi. 1:3-5 (not 1, 2).—Not Still- projecting digitiform acicular lobe and four er, 1996:30, pi. 10:figs. 1-5. (=E. us- setae directed anteriorly, and dorsal and chakovi, new species). ventral tentacular cirri, similar to lateral antennae; prominent conical facial tubercle Material examined.—Antarctic: Off ventral to ceratophore of median antenna Princess Elizabeth Land, 66°48'S, 71°24'E, (Fig. 1A; Mcintosh 1885: pi. 13, fig. 4). 540 m, British, Australian, New Zealand, Segment II with large semilunar nuchal Antarctic Research Expedition Sta. 30, 27 fold, first pair of large elytrophores, bira- Dec 1929, lectotype (BMNH 1941.3.3.16, mous parapodia and long ventral buccal cir- largest of three syntypes of E. gigantea). ri (Fig. 1A; Mcintosh: pi. 13 fig. 4). Drake Passage, between South America Biramous parapodia with notopodia and and Antarctica, 56°06'S, 66°19'W, 384-494 neuropodia about equal in size, both with m, Eltanin Sta. 740, 18 Sep 1963, 1 spec- projecting acicular lobes; notopodia round- imen (USNM 57466, ident. by Hartman ed basally, with projecting acicular lobes on 1967). South Shetland Islands, 61°25'S, lower sides; neuropodia subcorneal with 56°30'W, 300 m, Eltanin Sta. 993, 13 Mar projecting presetal acicular lobes and short- 1964, 1 specimen (USNM 57467, ident. by VOLUME 110, NUMBER 4 541

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Fig. 1. Eulagisca corrientis, A-D, syntype from Sta. 320 (BMNH 1885.12.1.71): E, syntype from Sta. 150 (BMNH 1885.12.1.83): A, Dorsal view of anterior end, right tentaculophore damaged; missing: styles of median and lateral antennae, palps, and tentacular cirri; B, Right elytrigerous parapodium, anterior view, aciculae dotted, style of ventral cirrus missing; C, Tips of long and short notosetae; D, Middle and upper neurosetae, with detail of tip, E, Right elytron, with detail of papillae. Scales =1.0 mm for A; 0.5 mm for B; 0.1 mm for C,D; E (not to scale).

Hartman 1967). South Orkney Islands, vesicles as on Fig. 4c, d and not Fig. 4e by 60°50'S, 42°55'W, 298-302 m, Eltanin Sta. Monro (1939), was selected as the lectoty-

1082, 14 Apr 1964, 1 specimen (USNM pe. The next largest syntype from Sta. 107, 57469, ident. by Hartman 1967). 59°53'S, with elytral tubercles as shown on Fig. 4e, 32°19'W, 523-671 m, Islas Orcada Cruise was selected as the holotype of E uschak- 1876, USARP Sta. 130, 25 Feb. 1976, 1 ovi, new species. The smallest syntype from specimen (USNM 74575). Sta. 39, which lacked elytra, was selected Remarks.—The type material of Eula- as a paratype of E. uschakovi, based on the gisca gigantea Monro (1939) consists of similarity of the neurosetae. three syntypes. They proved to be a mixture The two specimens from the Continental of at least two species. The largest syntype plateau off Kerguelen, in 172-262 m, de- from Sta. 30, with the elytral tubercles and scribed and figured by Desbruyeres (1976) 542 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON as E. gigantea, differ: the elytra have a neurosetae, with short clavate papillae. Dor- well-developed lateral fringe of long papil- sal tubercles bulbous, posterior ones on seg- lae and the surface is covered with conical ments 33-38 projecting posteriorly. spiny vesicles (pi. 1A-C), thus differing Bilobed prostomium wider than long, from E. gigantea, and probably indicating median antenna with large ceratophore in a new species. The specimens were not anterior notch, with style missing (probably available for examination to complete the long, similar to but longer than lateral an- description. tennae); lateral antennae inserted on ante- One of the specimens from Elephant Is- rior extensions of prostomium, without dis- land, Antarctica, in 280-432 m, referred to tinct ceratophores, with styles rather long, E gigantea by Hartmann-Schroder and Ro- finely papillated, with subterminal enlarge- senfeldt (1990:91, pi. 1; 3-5, not 1, 2), ments and filamentous tips; palps long, agree with E. gigantea, based on the photos stout, tapering, with six longitudinal rows of the elytra; the other specimen is referred of papillae; two pairs of rather large eyes to E. uschakovi, new species. Stiller (1996: with lenses on posterior half of prostomi- 30, pi. 10:figs. 1-5) reported E. gigantea um; tentaculophores (segment I) lateral and from the eastern Weddell Sea. Based on the anterior to prostomium, with prominent description and figures, it is added to E. us- digitiform acicular lobes, group of long se- chakovi, new species. tae (up to 10) directed anteriorly, and long Additional records of E. gigantea, with dorsal and ventral tentacular cirri, similar to elytra lacking, are omitted. They must be but longer than lateral antennae; with prom- referred to Eulagisca sp. inent conical facial tubercle below cerato- Description.—Lectotype with 38 seg- phore of median antenna (=subtentacular ments, 148 mm long, 44 mm wide with se- cirrus by Monro) (Fig. 2A; Monro 1939: tae. Specimen from South Shetland Island fig. 4a). Segment II with prominent semi- (USNM 57467) 68 mm long, 46 mm wide lunar nuchal pad and conical fleshy nuchal with setae, 38 segments. Specimen from fold covering posterior part of prostomium South Orkney Island (USNM 57469) 155 (latter may be pulled back), first pair of mm long, 94 mm wide with setae, 38 seg- large elytrophores, biramous parapodia, and ments. Body large, flattened, with long par- long ventral buccal cirri, inserted basally, apodia (parapodia and setae longer than similar to tentacular cirri (Fig. 2A; Monro body width). Body colorless except for trac- 1939: fig. 4a). es of brown transverse bands, with golden Biramous parapodia with small subcorn- setae. eal notopodia with projecting acicular Fifteen pairs of elytra, on bulbous ely- lobes; larger neuropodia with subtriangular trophores, large, thick, subreniform, cover- presetal acicular lobes and shorter subcon- ing dorsum, darker along posterior border, ical postsetal lobes (Fig. 2B; Monro 1939: without border papillae (Fig. 2F; Monro fig. 4f). Notosetae numerous, forming ra- 1939: fig. 4b). Surface of anterior part of diating bundle, about as stout as neurosetae, elytra with numerous conical microtuber- of three lengths, some shorter, curved, some cles, middle part with larger sharply pointed about as long as neurosetae, stout, acicular, conical tubercles and posterior part with with long spinous regions, tapering to bare curved conical tubercles (rarely 2-pronged), entire tips (Fig. 2B, C; Monro 1939: fig. 4g; some smaller spines, and soft globular ves- Uschakov 1962: pi. 9G). Neurosetae nu- icles, dotted yellowish distally (Fig. 2F; merous, forming fan-shaped bundle, with Monro 1939: fig. 4c, d, not e; Uschakov, long spinous regions and rather long, slight- 1966: pi. 9E, not F). Dorsal cirri with cy- ly hooked entire bare tips; upper ones more lindrical cirrophores, bulbous on posterior slender, with longer spinous regions (Fig. basal parts, with styles extending to end of 2D, E; Monro 1939: fig. 4h; Uschakov —

VOLUME 110, NUMBER 4 543

Fig. 2. Eulagisca gigantea, A. Specimen from South Shetland Island (USNM 57467); B-F, lectotype (BMNH 1941.3.3.16): A, Dorsal view of anterior end, pharynx partially extended; missing; right palp, median and right lateral antennae, right dorsal and ventral tentacular cirri, left parapodium of segment 2 not shown; B, Right elytrigerous parapodium, anterior view; C, Notoseta from same; D, Middle neuroseta from same; E, Upper neuroseta from same; F, Elytron, with detail of microtubercles, larger tubercles, and soft globular vesicles. Scales

= 2.0 mm for A; 2.0 mm for B; 0.1 mm for C-E; 5.0 mm for F.

1966: pi. 9H). Ventral cirri short, subulate, 1962:174 (part: Sta. 232, pi. 9F only; extending slightly beyond basal part of neu- 1966:174 (part: Sta. 232, pi. 9F only (En- ropodia (Fig. 2B). glish translation). —Hartman, 1967:21 Distribution. —Antarctic, Off Princess (part: Sta. 1003 only).—Averincev, 1972: Elizabeth Land, Davis Sea, Drake Passage, 131, pi. 13:1-7.—Hartmann-Schroder & Scotia Sea, South Shetlands, South Ork- Rosenfeldt, 1990:91 (part, pi. 1:1, 2, not neys, in 200-900 meters. 3-5). Not Monro, 1939 (lectotype). Stiller, 1996:30, pi. 10: figs. 1-5. Eulagisca uschakovi, new species Material examined. —Antarctic: Off Fig. 3 MacRobertson Land, 66°45'S, 62°63'E, 219 Eulagisca gigantea Monro, 1939:103 (part: m, British, Australian, New Zealand Ant- Sta. 39 & 107, Fig. 4e only).—Uschakov, arctic Research Expedition Sta. 107, 16 Feb 544 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

1931, holotype (BMNH 1941.3.3.18, as Stiller 1996, pi. 10 fig. 3). Dorsal cirri with next longest syntype of E. gigantea). Off large cylindrical cirrophores, inflated basal- Enderby Land, 66°10'S, 49°41'E, 300 m, ly, with long papillate styles extending be- B.A.N.A.R.E. Sta. 39, 17 Feb 1930, para- yond setae; dorsal tubercles nodular (Fig. type (BMNH 1941.3.3.17, as smallest syn- 3B; Averincev 1972: pi. 13:2). type of E. gigantea). Bilobed prostomium wider than long; Bransfield Strait, 62°41'S, 54°43'W, 210- median antenna with large ceratophore in 220 m, Eltanin Sta. 1003, 15 Mar 1964, anterior notch, with long, finely papillated paratype (USNM 57468, as E. gigantea by style with subterminal enlargement and fil- Hartman, 1967). Off Palmer Archipelago, amentous tip; lateral antennae inserted on 62°09'S, 57°49'W, 220-229 m, Eastwind anterior extensions of prostomium, without Sta. 66, 19 Feb 1966, 2 paratypes (USNM distinct ceratophores, styles similar to but 35289). Scotia Sea, 61°19'S, 44°26'W, 274- shorter than median antenna; palps long, 280 m, Islas Orcada Sta. 123, 22 Feb 1976, stout, tapering, with six longitudinal rows

1 specimen (USNM 74576). of papillae; two pairs of rather large eyes Ross Sea, 75°50'S, 173°08'W, 476 m, El- on posterior half of prostomium; tentacu- tanin Sta. 38, 31 Jan 1968, 2 paratypes lophores (segment I) lateral and anterior to (USNM 58350). prostomium, with digitiform acicular lobes, Types.—Holotype (BMNH 1941.3.318, group of long setae directed anteriorly, and next largest syntype of E. gigantea)'. 110 long dorsal and ventral tentacular cirri, sim- mm long, 70 mm wide with setae, 39 seg- ilar to but longer than antennae; with prom- ments, with elytra. Paratype (BMNH inent conical facial tubercle below ceratop- 1941.3.3.17, smallest syntype of E. gigan- hore of median antenna (Fig. 3A; Averin- tea): 56 mm long, 36 mm wide with setae, cev 1972:pl. 13:1; Stiller 1996:pl. 10, fig. 37 segments, without elytra but neurosetae 1). Segment II with prominent semilunar agree with holotype. Two paratypes from nuchal pad and conical nuchal fold cover- Ross Sea (USNM 58350): 140-190 mm ing posterior part of prostomium, first pair long, 85-100 mm wide with setae, 39 seg- of large elytrophores, biramous parapodia, ments, with elytra. Paratype from Palmer and long ventral buccal cirri, inserted ba- Archipelago (USNM 58351): 81 mm long, sally, similar to tentacular cirri (Fig. 3A; 48 mm wide with setae, 38 segments, with Averincev 1972:pl. 13:1; Stiller 1996: pi. elytra (figured). 10, fig. 1). Description.—Body brownish on mid- Biramous parapodia with smaller and dorsum, on bases of elytrophores and dorsal shorter notopodia and larger neuropodia, tubercles. Dorsum with ciliated bands (ca both with projecting digitiform acicular 14 per segment), continuing on elytro- lobes (Fig. 3B). Notosetae numerous, short phores and dorsal tubercles (Averincev to longer, slightly stouter than neurosetae, 1972: pi. 13:1). with long spinous regions and short bare Elytra large, thin (compared with E. gi- tapered tips (Fig. 3C; Averincev 1972: pi. gantea), subreniform, splashed with brown- 13:4; Stiller 1996: pi. 10, fig. 4). Neurosetae ish pigmentation; surfaces with numerous numerous, with long spinous regions, all ta- microtubercles on anterior part, sharply pering to long, slender, bare, pointed tips pointed tubercles on middle part, and pos- (Fig. 3D; Averincev 1972: pi. 13:3; Stiller terior part with additional globular vesicles 1996: pi. 10, fig. 5). Ventral cirri short, ta- with few to numerous terminal spines, and pered, papillate (Fig. 3B). some papillae but no thick fringe (Fig. 3E; Etymology.—The species is named for Monro 1939; fig. 4e; Uschakov 1962; pi. the late P. V. Uschakov, in recognition of 9F; Averincev 1972: pi. 13.5; Hartmann- his vast contributions to the study of the Schroder & Rosenfeldt 1990: pi. 1:1, 2; Polychaeta. VOLUME 110, NUMBER 4 545

Fig. 3. Eulagisca uschakovi, new species, A-D, paratype (BMNH 1941.3.3.17); E, paratype (USNM 58351): A, Dorsal view of anterior end, pharynx partially extended; missing: palps, styles of median and lateral antennae, dorsal and ventral tentacular cirri, except left one; segment 2 with nuchal fold partially folded back, right parapodium not shown: B, Right cirrigerous parapodium, posterior view; C, Tip of notoseta; D, Tips of two neurosetae; E, Right elytron, with detail of microtubercles, sharply pointed tubercles, spinous globular vesicles, and papillae. Scales =1.0 mm for A; 1.0 mm for B; 0.1 mm for C, D; 2.0 mm for E. 546 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Remarks.—The polynoids from the Da- to larger spiny tubercles posteriorly and vis Sea in 1 1-920 m, referred by Averincev medially, some rounded tubercles with me- (1972:131, pi. 13:1-7) to Eulagisca gigan- dial filamentous extensions (Fig. 4F). Dor- tea, were not available for examination. sal cirri with cylindrical cirrophores with The description and figures agree with E. glandular bulbous lobes on anterior and uschakovi. One of the specimens from El- posterior sides and distal brown bands; ephant Island (Antarctica) in 280-437 m, re- styles missing; dorsal tubercles nodular ferred to E. gigantea by Hartmann-Schrod- (Fig. AC). er and Rosenfeldt (1990:91, pi. 1:1, 2, not Bilobed prostomium with large cerato- 3-5), also agree with E. uschakovi. phore of median antenna in anterior notch, The elytra of E. uschakovi differ from the style missing; lateral antennae inserted on other species of Eulagisca in having sharp- anterior extensions of prostomium, with ly pointed tubercles and globular vesicles long papillate styles and filamentous tips; with few to numerous terminal spines. The two pairs of lateral eyes moderate in size; neurosetae of E. uschakovi agree with E. palps long, tapering, with six longitudinal puschkini Averincev by having neurosetae rows of papillae; tentaculophores lateral to tapering to sharp tips, rather than ending in prostomium and palps, each with projecting slightly hooked blunt tips, as in the other digitiform acicular lobe, missing dorsal and species of Eulagisca. This also applies to ventral tentacular cirri, and bundle of long the record of E. gigantea by Stiller (1996, setae (ca. 14); conical facial tubercle ventral pi. 10:figs. 3, 5) from the eastern Weddell to ceratophore of median antenna; dark Sea. brown bands on ceratophore of median an- Distribution.—Antarctic, off MacRobert- tenna, lateral extensions of prostomium for son Land, off Enderby Land, Davis Sea, lateral antennae, and tentaculophores (Fig. Bransfield Strait, off Palmer Archipelago, 4A). Segment II with semilunar nuchal pad Scotia Sea, Elephant Island, South Orkneys, and subcorneal nuchal fold, partially pulled Ross Sea, Weddell Sea, in 10-920 meters. back, first pair of large elytrophores, biramous parapodia, and long ventral buccal cirri, similar to lateral antennae (Fig. 4A). Eulagisca macnabi, new species Biramous parapodia with prominent pro- Fig. 4 jecting digitiform acicular lobes on both Material examined.—Antarctic: Off rami. Notopodium smaller than neuropo- South Orkneys, 61WS, 44°58'W, 283-329 dium, rounded, with projecting acicular m, Eastwind, Sta. 29, 11 Feb 1966, D. L. lobe on lower side; larger neuropodium

Pawson and D. F. Squires, coll., holotype with subcorneal presetal acicular lobe and (USNM 58352). shorter subcorneal postsetal lobe (Fig. 4B, Description.—Holotype 62+ mm long, C). Notosetae numerous, forming radiating 38 mm wide with setae, segments 24, plus bundle of three lengths, stouter than neu- small regenerating posterior end. Body flat- rosetae, acicular, with spinous rows and tened, with long parapodia (about as long rather long bare tips (Fig. 4B, D). Neuro- as body width); dorsum with transverse cil- setae numerous, forming fan-shaped bun- iated bands (ca 10 per segment) extending dle, with long spinous regions, upper ones on bases of elytrophores and dorsal tuber- more slender, all with rather long, slightly cles (Fig. 4A-C). hooked bare tips (Fig. 4B, E). Ventral cirri Elytra large, subreniform, covering dor- short, tapered, papillate (Fig. 4B, C). Ne- sum, with fringes of long border papillae phridial papillae beginning on segment VI. laterally, posteriorly and medially; surfaces Etymology.—The species is named for splotched with brownish pigmentation, con- the late Dr. James A. Macnab, a dedicated ical microtubercles on anterior part, small and inspirational teacher. VOLUME 110, NUMBER 4 547

Fig. 4. Eulagisca macnabi, new species, holotype (USNM 58352): A, Dorsal view of anterior end, missing: styles of median and right lateral antennae, right and left dorsal and ventral tentacular cirri, right ventral buccal cirrus, and elytra; nuchal fold partially pulled back; B, Right elytrigerous parapodium, anterior view, aciculae dotted; C, Right cirrigerous parapodium, posterior view, style of dorsal cirrus missing; D, Notoseta from same;

E, Middle and upper neurosetae from same; F, Right elytron, with detail of microtubercles, spiny vesicles, and papillae. Scales = 2.0 mm for A-C; 0.1 mm for D, E; 2.0 mm for F 548 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Distribution.—Antarctic, South Orkneys, Diagnosis.—Body subrectangular, taper- 300 meters. ing slightly anteriorly and posteriorly, with Remarks.—The elytra of E. macnabi dif- segments up to 37. Elytra and bulbous ely- fer from the other species of Eulagisca by trophores 16 pairs, on segments 2, 4, 5, al- the well developed border fringes of papil- ternate segments to 23, 26, 29, 32, 33. El- lae and the surface spiny tubercles. ytra large, covering dorsum, smooth, without tubercles or papillae. Dorsal cirri on Eulagisca puschkini Averincev, 1972 non-elytrigerous segments, with cylindrical cirrophores, bulbous basally, with papillate Eulagisca puschkini Averincev, 1972:131, styles extending to tips of setae; dorsal tu- pi. 13:8-12. bercles slightly bulbous. Prostomium bi- Remarks.—A single specimen was col- lobed, with two long finely papillate palps lected by aqualung in the Davis Sea, in 32 and three papillate antennae; ceratophore of meters, on rocky bottom, among sponges median antenna large, in anterior notch of and hydroids (Type deposited in ZIASL). prostomium, lateral antennae inserted on The type was not available for examination. anterior extensions of prostomium subter- The following was reported by Averincev. minal to ceratophore of median antenna; The large specimen is 175 mm long, 55 two pairs of lateral eyes, moderate in size. mm wide without setae, and 85 mm wide Tentaculophores of segment I lateral to pro- with setae, number of segments ?. The an- stomium, each with single seta, without terior end, including the prostomium, palps, projecting acicular lobe (as in Eulagisca), tentaculophores, facial tubercle, nuchal and papillate dorsal and ventral tentacular fold, and pharynx, agree with Eulagisca gi- cirri; facial tubercle weakly developed. Sec- gantea. The elytra differ from the other spe- ond or buccal segment with thick, fleshy cies of Eulagisca by lacking border and sur- nuchal fold, first pair of elytrophores, bi- face papillae, by having numerous micro- ramous parapodia, and long ventral buccal tubercles on the anterior part, with a red- cirri, lateral to ventral mouth. Parapodia bi- brown iridescent spot on the medial half ramous, noto- and neuropodia about equal (pi. 13:8). The biramous parapodia have in length, both subcorneal with projecting large digitiform processes with inner acic- acicular lobes. Numerous notosetae forming ulae. The notosetae are thicker than the radiating bundles, nearly as long as neuro- neurosetae, with transverse spinous rows setae, of two kinds: stout, acicular (about as and short bare tips (pi. 13:12). The neuro- stout as neurosetae), smooth or with faint setae have subdistal spinous rows and end spinous rows; slender, capillary, smooth or in tapered bare tips, and few with small sec- with few spines. Numerous neurosetae stout, regions long, ondary tooth (pi. 13:9-11). with long spinous and bare, slightly entire tips, The elytra of E. puschkini differ from hooked with or Ventral other members of Eulagisca by lacking bor- without minute secondary tooth. cirri short, tapered, papillate. Pygidium der papillae, surface tubercles and vesicles, with pair papillate anal cirri. Nephridial and with only numerous microtubercles on of papillae beginning on segment IV. the anterior half. The neurosetae of E. pus- Etymology.—Pareulagisca from Greek chkini, with tapered bare tips, agree with para = near + Eulagisca in allusion to the those of E. uschakovi new species. similarity of the two genera. Distribution.—Antarctic, Davis Sea, in 12 meters. Pareulagisca panamensis (Hartman 1939), new combination Pareulagisca, new genus Fig. 5 Type species. —Eulagisca panamensis Eulagisca panamensis Hartman, 1939:31, Hartman, 1939. Gender: feminine. pi. 3:figs. 38-42. VOLUME 110, NUMBER 4 549

Fig. 5. Pareulagisca panamensis, holotype of Eulagisca panamensis (LACM-AHF 10): A, Dorsal view of anterior end; B, Right elytrigerous parapodium, anterior view, aciculae dotted; C, Right cirrigerous parapodium, posterior view;

D, Three stout notosetae from same; E, Two slender, capillary notosetae from same; F, Lower and upper neurosetae from same; G, Right elytron. Scales = 2.0 mm for A; 1.0 mm for B, C; 0.1 mm for D-F; 2.0 mm for G.

Material examined.—Pacific: Panama, subrectangular, flattened, tapering slightly Pinas Bay, 07°34'N, 78°12'W, shore, rock, anteriorly and posteriorly. Elytra and bul- Velero Sta. 436, 28 Jan 1935, holotype bous elytrophores 16 pairs (not 15, as in- (LACM-AHF 10). dicated by Hartman) on segments 2, 4, 5, Description.—Holotype 42 nun long, 15 7, alternate segments to 23, 26, 29, 32, 33, mm wide with setae, 37 segments. Body dorsal cirri present on posterior four seg- 550 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON ments. Elytra large, covering dorsum, Table 1. —Comparison of characters for Eulagisca and Pareulagisca. fleshy, smooth, without tubercles or papillae, with rust-colored semilunar pigmenta- Eulagisca Pareulagisca tion on posterior half (Fig. 5G; Hartman Elytra 15 pairs 16 pairs 1939, pi. 3: fig. 42). Dorsal cirri with cylin- Palp 6 longitudinal rows without longitudi- cirrophores, basally, drical bulbous and of papillae nal rows of pa- long styles extending to about tips of setae, pillae with short papillae on distal half and brown Tentacu- with projecting acic- without projecting pigmentation on basal three-fourths; dorsal lophore ular lobe and acicular lobe group of long se- and single seta tubercles slightly bulbous (Fig. 5C; Hart- tae pi. fig. man 1939: 3, 39). Notosetae of one kind, long, of 2 kinds: 1) Bilobed prostomium suboval, without ce- spinous rows, ta- stout, acicular phalic peaks; ceratophore of median anten- pering to blunt 2) slender, cap- tips illary na in anterior notch of prostomium, very large, with long papillated style enlarged subdistally, with filamentous tip; lateral an- pi. 3, figs. 38, 41). Ventral cirri short, ta- tennae inserted on anterior extensions of pered, with short papillae (Fig. 5B, C). prostomium, subterminal to large cerato- Distribution. —Central Pacific Ocean, phore of median antenna, with shorter Panama, intertidal. styles; palps long, thick, tapered, finely pap- Remarks.—Pareulogisca panamensis is illated; two pairs of eyes moderate in size; unusual in having 16 pairs of elytra, instead tentaculophores lateral to prostomium, each of the more common 15 pairs, and in hav- with single seta, dorsal and ventral tentac- ing two distinct types of notosetae: stout ular cirri, similar to median antenna; facial and slender, capillary. The large size of the tubercle indistinct (Fig. 5A). Second or ceratophore of the median antenna is also buccal segment with triangular nuchal fold, unusual. first pair of bulbous elytrophores, biramous Pareulagisca differs from Eulagisca as parapodia, and long ventral buccal cirri, shown in Table 1. similar to tentacular cirri (Fig. 5A).

Notopodia of biramous parapodia almost Literature Cited as long as neuropodia, subcorneal, with pro- Augener, H. 1932. Antarktische und Antiboreale Po- jecting acicular lobes on lower sides; neu- lychaeten nebst einer Hirundinee.—Scientific ropodia subcorneal, with longer presetal results of the Norwegian Antarctic Expedition acicular lobes and shorter, rounded postsetal 1927-28. Det Norske Videnskaps-Akademii lobes (Fig. 5B, C; Hartman 1939: pi. 3, fig. Oslo, Pp. 1-88. 39). Notosetae numerous, forming radiating Averincev, V. G. 1972. [Benthic polychaete worms of the Errantia from the Antarctic and Subantarctic bundle of two kinds: acicular, about as stout in the material of the Soviet Antarctic Expedi- as neurosetae, mostly smooth and shorter, tions].—Akademiia Nauk SSSR Zoologisches- curved, with 6-12 widely-spaced spinous kii Institut 5:88-293. [In Russian] rows (Fig. 5D: Hartman 1939: pi. 3, fig. Benham, W. B. 1921. Polychaeta. Australian Antarc- tic Expedition 1911-14, Under the leadership of 40); and slender, tapering to pointed and Sir Douglas Mawson, D.Sc.B.E.—Scientific capillary tips, smooth or with few widely- Reports, Series C. Zoology & Botany 6(3): 1- spaced spines (Fig. 5E). Neurosetae numer- 128. ous, forming fan-shaped bundle, lower Desbruyeres, D. 1976. Bionomie benthique du plateau shorter to upper longer, with long spinous continental de l'archipel des lies Kerguelen- Macrofaune 3. Contribution a 1' etude des Pol- regions and rather long bare, slightly ynoidae (Annelides Polychetes) des Terres aus- hooked entire tips, upper with few minute trales et Antarctiques Franchises. —CMFRA secondary tooth (Fig. 5F; Hartman 1939: Prospections en Ocenographie Biologique et VOLUME 110, NUMBER 4 551

Bionomie Benthique aux abords des lies Ker- -. 1936. Polychaeteworms.il.—Discovery Re- guelen et Crozet No. 39:83-96. ports 12:59-198.

Hartman, O. 1939. Polychaetous annelids. Part I. -. 1939. Polychaeta.—Reports British Austra- Aphroditidae to Pisionidae.—Allan Hancock lian and New Zealand Antarctic Research Ex- Pacific Expeditions 7(1): 1-156. pedition, 1929-1931, series B 4(4):87-156.

. 1967. Polychaetous annelids collected by the Pettibone, M. H. 1989. A new species of Benhami- USNS Eltanin and Staten Island cruises, chiefly polynoe, (Polychaeta:Polynoidae:Lepidastheni- from Antarctic Seas.—Allan Hancock Mono- inae) from Australia, associated with the unat- graphs in Marine Biology 2:1—387. tached stylasterid coral Conopora adeta.—Pro- Hartmann-Schroder G. 1986. Die Polychaeten 56 Re- ceedings of the Biological Society of Washing- ise der "Meteor" zu den South Shetland-Inseln ton 102:300-304. (Antarktis).—Mitteilungen aus dem hambur- Stiller, M. 1996. [Distribution and biology of the gishchen zoologischen Museum und Institut 83: Aphroditides and Polynoids (Polychaeta) in the 71-100. eastern Weddell Sea and the Lazarev Sea (Ant-

, & P. Rosenfeldt. 1990. Dis Polychaeten der arctica).]. —Reports on Polar Research, No. 185 "Walther Herwig" Reise 68/1 nach Elephant Is- Heft Nr. 185/1996:1-200. [in German]

land (Antarktis) 1985 Teil 1: Aphroditidae bis Uschakov, P. V 1962. [Polychaetous Annelids of the Cirratulidae.—Mitteilungen aus dem Hambur- Families Phyllodocidae and Aphroditidae from gishchen zoologischen Museum und Institut 87: the Antarctic and Subantarctic Waters.—Bio- 89-122. logical Results of Soviet Antarctic Expedition

Kinberg, J. G. H. 1856. Nye slagter och arter af An- (1955-1958).] Issledovaniia fauny morei Zoo- nelider.—Ofversigt af Konglia Vetenkaps-Aka- logicheskii Institut Akameniia Nauk SSSR, demiens Forhandlingar, Stockholm 12:381-388. I(DC): 129-189. [in Russian]

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