Vol. 83, No. 47, pp. 527-544 9 February 1971 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON SOME POLYCHAETES OF THE SUPERFAMILY EUNICEA FROM THE NORTH PACIFIC NORTH ATLANTIC OCEANS^ FEB 9 1971 FIC AND^HS0/V7>s By Katharine D. Hobson Systematics-Ecology Program, Marine Biological LaZ?ora*e^^RARl^^ Woods Hole, Massachiisetts One new species of Onuphis is presented together with a discussion of and key to five other species of this genus and five new records of species of Lumbrineridae, Arabellidae, and Dorvilleidae. Most of the polychaetes from Washington were collected from the Strait of Juan de Fuca by the author while on Brown Bear cruise #365 (June 1965); from near Orcas Island by H. L. Sanders (July 1967), and obtained from the collections of K. Banse, K. D. Hobson, and F. H. Nichols from Puget Sound (February, April, and August 1963; October 1965). Some of the polychaetes from New England were collected from the slope off New England by H. L. Sanders ( May 1961, August 1962); and from Cape Cod Bay by the Biotic Census of the Systematics-Ecology Program (February 1966 through October 1969). The latter are now deposited in the Gray Museum of the Marine Biological Laboratory ( CM ) . Supple- mentary material was obtained from the following museums or institutions: the AUan Hancock Foundation (AHF), through O. Hartman and K. Fauchald; the British Museum of Nat- ural History (BMNH), through J. D. George; the Bmreau of Commercial Fisheries ( BCF ) , through R. L. Wigley ; the Mu- seum of Comparative Zoology (MCZ), through H. W. Levi; 1 Contribution No. 215 from the Systematics-Ecology Program, Marine Biological Laboratory, Woods Hole, Massachusetts 02543. This research was supported primar- ily by a grant from the Whitehall Foundation to the Systematics-Ecology Program. Collection of material was supported by NSF grant GB-747 and by Contract Nonr 3070(03) from the Office of Naval Research to the Systematics-Ecology Program. 47— Proc. Biol. Soc. Wash., Vol. 83, 1970 (527) 528 Proceedings of the Biological Society of Washington Fig. 1. Onuphis elegans (collection of Banse, Hobson, and Nichols, Puget Sound): a, Anterior view of 1st parapodium; b, anterior view of 25th parapodium; c, slender pseudocompound hook from 1st parapodium; d, stout pseudocompound hook from same. the Pacific Marine Station Museum (PMS), through J, A. Blake; and the Smithsonian Institution (USNM), through M. L, Jones. The manuscript benefitted from discussions and correspondence with K. Fauchald and K. Banse. Eunicean Polychaetes 529 SUPERFAMILY EUNICEA Onuphidae Onuphis Audouin and Milne Edwards All species of Onuphis with cirriform branchiae that begin on the first setiger were reviewed in order to better characterize O. elegans and O. iridescent. In some species important specific characters, such as first occurrence of branchiae and of subacicular hooks, were found to vary with the size of the worm. Although the species discussed below are very closely related, they can be distinguished by considering this variation together with other characters. Onuphis elegans (Johnson) Figure la-d Northria (sic) elegant Johnson 1901, p. 406, pi. 8, figs. 77-85. Nothria elegans. — Hartman 1968 (in part), p. 675, figs. 1, 2, 5 only. Not Hartman 1944, p. 88, pi. 5, figs. 113-117. Nothria sp. near N. elegans and JV. iridescens. — Banse, Hobson, and Nichols 1968 (in part, Form A only), p. 534. Onuphis elegans.— E. Berkeley 1927, p. 408.— C. Berkeley 1967, p. 1056. —Berkeley and Berkeley 1932, p. 312; 1948, p. 93, fig. 141. Material examined: WASHINGTON, syntypes of N. elegans (AHF; MCZ 1895) from Puget Sound. — Collections of Banse, Hobson, and Nichols from Puget Sound (10-23 m). CALIFORNIA, Tomales Bay (PMS 180241). — Velero station 3048 near Los Angeles Lighthouse (19 m) and five localities in central and southern California (intertidal to 18 m) (AHF). BRITISH COLUMBIA, Long Bay, E. Berkeley, col- lector (intertidal) (USNM 38275). Description: Pigmentation, when present, is usually as described by Johnson. The first 5 or 6 setigers are cylindrical, then the body becomes flattened. The inner paired antennae usually extend only to setiger 3-7 (range, 3-11). The branchia of the first parapodiimi (Fig. la) is longer than the dorsal cirrus and becomes relatively longer farther posteriorly. The branchiae are thick (Fig. lb) in setigers 10 through 60-80, rather than slender as in the other species treated here (c.f.. Figs. 2d, 3b, 5a). Ventral cirri are usually cirriform through setiger 5, (setiger 6 in the specimens from Long Bay, B.C. ) . Pseudocompound hooks occur in the first four setigers ( rarely 3 ) . The hooks are usually tridentate ( Fig. Ic, d ) but the third tooth may be very small (as in Fig. 2f). I have found bidentate hooks without a trace of a third tooth only in the specimens from Long Bay, B. C. and in a speci- men from Tomales Bay, California. These specimens also have a few quadridentate hooks. Supraacicular narrowly limbate setae are present from the first setiger, appearing faintly hairy. Pectinate setae are present INSTITUTiOW FEB 9 WH 530 Proceedings of the Biological Society of Washington Fig. 2. Onuphis farallonensis new species: a, Dorsal view of anterior end of holotype (AHF), setae omitted (stippling indicates location of pigment); b, anterior view of 1st parapodium; c, anterior view of 5th parapodium; d, anterior view of 26th parapodium; e, tip of slender pseu- docompound hook from 1st parapodium; f, stout pseudocompound hook Eunicean Polychaetes 531 from the first setiger, are distally slightly oblique, and have about 14 teeth. There are no compound spinigers. Subacicular hooks usually begin on setiger 9 ( occasionally on 10 or 11 ). The tube is parchmentlike with adherent sand grains. Remarks: There does not seem to be any variation of specific char- acters with size of this species. Onuphis elegam is characterized by its short antennae, thick branchiae, and first occurrence of subacicular hooks from setiger 9 ( occasionally 10 or 11 ) . Distribution: British Columbia to southern California, intertidal to 23 meters. Onuphis farallonensis new species Figure 2a-g Nothria elegam.— Hartm&n 1944, p. 88, pi. 5, figs. 113-117; 1968 (in part), p. 675, figs. 3-4 only. Not IV. elegans Johnson, 1901. Material examined: CALIFORNIA, Vehro station 887, east of Middle Farallon Island (68 m), Holotype and 6 Paratypes (AHF); Paratype (USNM 42026). Description: None of the specimens are complete. The holotype is 3.8 cm long for about 80 setigers, 1.2 mm wide at the 20th setiger. The body is subcylindrical. There are dorsal transverse reddish brown bands of pigment on the anterior margin of anterior segments (Fig. 2a) and traces of pigment on the bases of the prostomial appendages and on the dorsal surface of the parapodia. This pigmentation pattern occurs on about the first 30 setigers. The prostomivun is triangular, and the frontal antennae are elongate- oval. The ceratophores of the occipital antennae are distinctly annulated. The styli of the inner paired anteima are the longest and reach to setiger 10 to 13. The palpi are large and globular. The presetal lobes of the first few setigers are rounded and have a transverse fold (Fig. 2b, c). Postsetal lobes are elongate through about the first 10 setigers and gradually diminish in size posteriorly. Ventral cirri are cirriform through setiger 3 to 5, depending on size of the worm, and are padlike thereafter. The branchiae begin on the first setiger, ex- cept in the smallest worm (0.6 mm wide at the first segment) where they begin on setiger 4. When fully developed the branchiae are longer than the dorsal cirri ( Fig. 2c, d ) . The hooded pseudocompound hooks occur in the first 4 to 6 setigers. Their distribution in a paratype is as follows: the first setiger has 11 slender hooks, most of which are bidentate (Fig. 2e), the length of the blade is about 40 to 50 times the width of the blade at midlength, there from same; g, tip of subacicular hook from 26th parapodium (teeth broken). Figures b-g from dissected paratype (AHF). 532 Proceedings of the Biological Society of Washington Fig. 3. Onuphis iridescens ( F. H. Nichols' collection, Puget Sound ) : a, Anterior view of 1st parapodium; b, anterior view of 25th parapodium; c, pseudocompound hook from 1st parapodium; d, pseudocompound hook from 2nd parapodium of another specimen. are 2 stout barely tridentate hooks (Fig. 2f), the second setiger has at least 8 slender hooks (7 bidentate, 1 tridentate) and 1 stout bidentate hook, the third setiger has 8 slender and 1 stout bidentate hook, the fifth setiger has 3 fairly slender and 3 stouter bidentate (or barely tridentate) hooks and 2 supraacicular limbate setae. The long slender pseudo- compoimd hooks are most noticeable in the first 3 setigers. Hartman (1944) has given additional figures of the pseudocompound hooks. Eunicean Polychaetes 533 There are no compound spinigers. Pectinate setae are distally slightly oblique, have about 11 teeth, and are first evident on the fifth setiger. Subacicular hooks (Fig. 2g) begin on the 10th setiger. The mandibles of some specimens are calcified anteriorly. Maxilla I of a paratype is falcate. MaxiUa II has 9 teeth on the left and 10 teeth on the right. Left maxilla III has 10 teeth and left maxilla IV has 6 teeth. The fused right maxilla III and IV has 11 teeth. Both maxillae V have a single tooth. Sand grains of various sizes (0.1 to 2.0 mm diameter) adhere to the parchmenthke tube. Diagnosis: An Onuphis species with reddish brown dorsal bands, cirriform branchiae from the first setiger, ventral cirri cirriform through the 5th setiger, mostly bidentate pseudocompound hooded hooks with very long slender blades, and subacicular hooks from the 10th setiger. Remarks: This species most resembles O. shirikishinaiensis (Ima- jima) in having bidentate pseudocompound hooks Mdth long slender blades. Onuphis farallonensis differs from O. shirikishinaiensis in having mostly bidentate hooks in the first 4 to 6 setigers, and in the first occur- rence of subacicular hooks. Distribution: Southern California, in 68 meters. Onuphis irtdescens (Johnson) Figm-es 3a-d; 4a-c Northia (sic) iridescens Johnson 1901, p. 408, pi. 8, figs. 86-87, pi. 9, figs. 88-92. Nothria sp. near N. elegans and N. iridescens. — Banse, Hobson, and Nichols 1968 (in part. Form B only), p. 534. Nothria iridescens. — Fauchald 1968, p. 24, pi. 7, fig. a. — Hartman 1944 (in part), p. 87, pi. 5, figs. 99-104; 1968, p. 681, figs. 1-3.— Moore 1911, p. 255. Not Hartman 1965, p. 104. Onuphis iridescens. — ^E. Berkeley 1927, p. 408. — C. Berkeley 1967, p. 1056.— Berkeley and Berkeley 1948, p. 93, fig. 140. Material examined: BRITISH COLUMBIA, Holotype of N. iridescens from Victoria (MCZ 1887). — Off Ruxton Passage, E. Berkeley, collector (USNM 38283). WASHINGTON, two specimens from Orcas Island (40 m) H. L. Sanders, collector. — Many specimens collected by the author from the Strait of Juan de Fuca ( 140-170 m ) . — Banse, Hobson, and Nichols' collections from Puget Sound (13-200 m). SOUTHERN CALIFORNIA, Vehro stations 893, 981, 1130, 1133, 1135, 1137, 1157, 1348, 2175, 2354, 4785, 7028 (25^30 m) (AHF). Description: The inner paired antennae of this species usually reach to setiger 13 to 20 (range, 7 to 25). The branchiae begin on the first setiger except in smaller specimens (see Fig. 4a). The first few pairs of branchiae are shorter than the dorsal cirri ( as in Fig. 3a ) , but exceed the dorsal cirri in length thereafter. The branchiae are slender throughout 534 Proceedings of the Biological Society of Washington Q^^H^ 6 X - X X X yxK. X X ^^^B . 4 . xx>«<>« ^iSfl^df^S^ • o ocQ) a ▲ - . b - O (CD CD - hi 6 - » O^A4«4^0»« O ▲ - O _ CkOC^^ OODCCD O . }- III 4 . x>s< X >060O««« X X - 2 - X X X 1 1 1 1 1 1 1 1 1 - c 14 - ^Bottcsb CI o c3) d) - 12 Z# •i^ • • • A ; CO • o» 9 o avo • 1 o o 10 - X X X >x <• Qi - 8 - w )«^S«««>« 1 1 1 1 1 r - 2 0.6 1.0 1.4 1.8 mm A = British Columbia A= Strait "\washinqton (x= juveniles) •-Puget SoundJ orCalifornia Fig. 4. Variation in specific characters of O. iridescens versus width of 1st segment (ram) : a, First setiger with branchiae; b, last setiger with cirriform ventral cirrus; c, first setiger with subacicular hooks. Eunicean Polychaetes 535 (Fig. 3b). The ventral cirri are usually cirriform through setigers 5 to 7, but through only 2 to 4 on smaller specimens (see Fig. 4b). The first 3 to 5 setigers have tridentate pseudocompound hooks (Fig. 3c, d ) . The distal tootli of these setae usually extends further than in the setae of O. elegans; however, this character does not separate the species reliably. Narrowly limbate setae are present from the first setiger and appear faintly hairy. I have found pectinate setae no earlier than the 4th setiger. Subacicular hooks usually begin on setigers 11 to 14, but as early as 8 to 10 in smaller specimens (see Fig 4c). The tube is parchmentlike with adherent sand grains or a thick coating of mud. Remarks: The size of individuals of O. iridescent must be considered when defining this species because of the size-dependent variation of some important characters (summarized in Figm-e 4). The juveniles ( :^ 0.6 mm viddth at first segment) depicted in Fig. 4a-c all have slender branchiae, and were collected from stations where the only Onuphis species known to occur are O. iridescens or O. iridescens and O. elegans. Therefore these specimens probably do represent juveniles of O. irides- cens. Onuphis iridescens occasionally has been regarded as being close to O. elegans, if not synonymous (Hartman 1944, 1961; Monro, 1930). In Washington and British Columbia the adult specimens of the two species are easily distinguished by the shape of their branchiae, the first occur- rence of subacicular hooks, and the length of the inner paired antennae. In California, however, some specimens have been found that appear to be intermediate between the two species (see below). Onuphis iridescens occurs in deeper water than does O. elegans. Moore (1911) reports the former species from as deep as 1460 m in Monterey Bay; Fauchald (1968) reports it from as deep as 2400 m in western Mexico; Ushakov (1950) reports it from 252-1643 m in the Sea of Okhotsk; and Monro (1930) reports it from 650 m near the southern tip of South America. I have not verified the records from the western Pacific or southern Atlantic by examination of material. Distribution: British Columbia to western Mexico, possibly also in the Sea of Okhotsk and the South Atlantic Ocean. In 13 to 2400 meters. Onuphis sp. "intermediates" Onuphis iridescens. — Hartman 1944 (in part), p. 87. Material examined: CALIFORNIA, Tomales Bay (PMS 180240, 180242).— Southern California, Velero stations 887, 893, 1126, 2114, 2314, 3048, and 4842 (8-143 m) (AHF). Description: These specimens range from 0.6 to 1.4 mm in width at the first segment. They may have either no pigmentation pattern or dorsal transverse reddish brown bands of pigment on the posterior part of each segment or in the intersegmental grooves. The inner paired antennae reach to setigers 7 to 17; tlie branchiae are slender throughout; ventral cirri are 536 Proceedings of the Biological Society of Washington Fig. 5. Onuphis opalina (BCF): a. Anterior view of 24th para- podium; b, pseudocompound hook from 1st parapodium. cirriform through setigers 5 or 6; hooded pseudocompound hooks are tridentate and occasionally have long slender blades; pectinate setae occur from the 5th setiger; and subacicular hooks begin on the 9th setiger. Remarks: The "intermediates" resemble O. iridescens (see above) in the shape of the branchiae and O. elegans (see above) in the first occurrence of subacicular hooks. These specimens have been found in samples with O. iridescens, O. elegans, or O. farallonensis. How- ever, I have found no samples from California in which one of these three species co-occurs with another. "Intermediates" are not known for Washington and British Columbia. In Puget Sound, O. iridescens and O. elegans do co-occur in the same grab sample. The "intermediates" may result from interbreeding between O. elegans and O. iridescens ( and perhaps also O. farallonensis); they may represent a new species; or they may belong to a highly variable O. iridescens. Only through a laboratory Eunicean Polychaetes 537 14 ▲ AmA M M A . ▲ A AM A daxDO) O CIT) O) O o c& CD 12 CD CO O O o o UJ A ax> CD o CO CCD O O O ^ O o AO ; in 10 8 o o o -