JAMES A. BLA Revision of the Genus Polydora from f The East Coast of North America Polychaeta: Spionidae) SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY NUMBER 75 S E R I A L P U B L I C A T I O N S OF T H E S M I T H S O N I A N I N S T I T U T I O N The emphasis upon publications as a means of diffusing knowledge was expressed by the first Secretary of the Smithsonian Institution. In his formal plan for the Insti- tution, Joseph Henry articulated a program that included the following statement: "It is proposed to publish a series of reports, giving an account of the new discoveries in science, and of the changes made from year to year in all branches of knowledge." This keynote of basic research has been adhered to over the years in the issuance of thousands of titles in serial publications under the Smithsonian imprint, com- mencing with Smithsonian Contributions to Knowledge in 1848 and continuing with the following active series: Smithsonian Annals of Flight Smithsonian Contributions to Anthropology Smithsonian Contributions to Astrophysics Smithsonian Contributions to Botany Smithsonian Contributions to the Earth Sciences Smithsonian Contributions to Paleobiology Smithsonian Contributions to Zoology Smithsonian Studies in History and Technology In these series, the Institution publishes original articles and monographs dealing with the research and collections of its several museums and offices and of professional colleagues at other institutions of learning. These papers report newly acquired facts, synoptic interpretations of data, or original theory in specialized fields. These pub- lications are distributed by mailing lists to libraries, laboratories, and other interested institutions and specialists throughout the world. Individual copies may be obtained from the Smithsonian Institution Press as long as stocks are available. S. DILLON RIPLEY Secretary Smithsonian Institution SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY NUMBER 75 James A, Blake Revis ion of the Genus Polydora from The East Coast of North America (Polychaeta: Spionidae) SMITHSONIAN INSTITUTION PRESS CITY OF WASHINGTON 1971 ABSTRACT Blake, James A. Revision of the Genus Polydora from the East Coast of North American (Polychaeta: Spionidae). Smithsonian Contributions to Zoology, num- ber 75, 32 pages, 1971.?Twelve species of Polydora from the east coast of North America are described on the basis of recent collections as well as material de- posited in various museums. The study includes 11 previously described species, one new species, 10 synonyms, and one nomen nudum. Diagnosis of the genus Polydora, with a key to the species covered, are included. Official publication date is handstamped in a limited number of initial copies and is recorded in the Institution's annual report, Smithsonian Year. UNITED STATES GOVERNMENT PRINTING OFFICE WASHINGTON : 1971 For ale by the Superintendent of Document*, U.S. Government Printing Office Wathington, D.C. 20408 ? Price 45 cent* (paper cover) James A. Blake Revision of the Genus Polydora from The East Coast of North America (Polychaeta: Spionidae) The family Spionidae is among the largest, in num- ber of species, of the Polychaeta. They are typically marine bottom-deposit feeders and occur mostly in coastal waters. The family is rare in the deep sea. The spionids have a pair of long peristomial palps which are used in feeding. Each palp has a ciliated groove along which particles move toward the mouth; some of these particles are used to build tubes while others are utib'zed as food. The sorting mechanism, however, is not as well de- veloped as in some other polychaete families. The Spionidae has been divided into several subfamilies (Sdderstrom, 1920; Orrhage, 1964). The subfamily Spioninae contains the genera Pygospio, Spio, Paraspio, Polydora, Bpccardia, and Pseudo- polydora. The three latter genera comprise the polydorid complex in which the fifth setigerous segment is modified and includes specialized setae or spines. The structure of the spines form useful taxonomic characters. Major systematic treatments of the polydorids include works by Carazzi (1893), Mesnil (1896), Sdderstrom (1920), Fauvel (1927), Hartman (1941), Woodwick (1964), and Day (1967). Our knowledge of spedes of Polydora from the east coast of North America comes largely from the studies of Verrill (1880, 1881, 1885a, b), Webster (1879a, b), Andrews 1891a, b), Moore (1907), and Dr. James A. Blake is on the staff of the Pacific Marine Sta- tion, University of the Pacific, Dillon Beach, Marin County, California, 94929. Hartman (1942, 1943, 1944, 1945, 1965). These five authors described eleven species of Polydora. Four additional spedes, previously known from other geographic areas, were reported. Special men- tion should be made of the paper by Bosc (1802), in which the description of the type spedes of the genus, P. cornuta Bosc from Charleston, South Carolina, was given. Unfortunately, the description of that spedes was superfidal and spedfic deter- mination is today impossible. Thus, the type-spedes of Polydora is indeterminable. Considerable confusion exists with regard to the four spedes of Polydora erected by Verrill. Only for P. concharum Verrill (1880) are adequate de- scriptions and figures given (Verrill, 1885a; Hart- man, 1942, 1944) and type-material available. Polydora gracilis Verrill (1880) and P. tubifcx Verrill (1885b) were described but never figured, while P. littorea Verrill (1881) is a nomen nudum. Hartman (1944) belatedly published a number of Verrill's unpublished plates, which contained some figures of Polydora, including P. concharum and P. littorea. Webster (1879a, b) described P. hamaia,P. caeca, and P. ligni. Polydora caeca, however, was dedared a junior homonym by Hartman (1943) and re- named P. websteri. Polydora hamata was referred to Boccardia by Blake (1966). Polydora aggregata was named and the larvae described by Blake (1969b). The other spedes described from eastern North 2 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY America include Polydora commensalis Andrews (1891a), P. colonia Moore (1907), and P. tetra- branchia Hartman (1945). Four species from other regions have also been reported to occur in eastern North America, including P. ciliata (Johnston) and P. quadrilobata Jacobi, reported by Procter (1933); P. socialis (Schmarda), reported by Hart- man (1945); and P. caulleryi Mesnil, reported by Hartman (1965). Despite these numerous references and others, only Hartman (1945) provides useful keys and de- scriptions to a number of species. Unfortunately, that study is limited to North Carolina and ex- cludes several species found in adjacent areas. The present study is an attempt to reevaluate older rec- ords and to provide new information on species of Polydora for the entire eastern seaboard. New col- lections come from Newfoundland to South Caro- lina. Most were collected by the writer, while others were collected by various individuals, acknowledged in the text. The bulk of these collections have been deposited in the United States National Museum. Field collections of adult Polydora were made intertidaUy and subtidally from shells and sediment. As a result of the author's location in Maine, ex- tensive collections were made in that area. Sub- tidal samples were obtained with either a biological dredge or a 0.05 m* Ponar Grab. Sediment was washed with sea water through fine-mesh screens. The tubes were removed and placed in jars in run- ning seawater. In the laboratory, the tubes were carefully dis- sected and the worms removed. Polydora from shells were removed after cracking the shells with a ham- mer or pliers. Adults were relaxed in 0.15% propylene phe- noxytol prior to fixation in 10% formalin or Bouin's fluid. After several days in the fixative, the worms were removed and stored in 70% ethyl alcohol. Older collections of Polydora, deposited in the Allan Hancock Foundation, University of Southern California (AHF), Academy of Natural Sciences of Philadelphia (ANSP), United States National Museum (USNM), and the Yale Peabody Museum of Natural History (YPMNH) were also examined. The following people assisted by arranging for the loan of specimens from their respective Mu- seum: Marian H. Pettibone (USNM), Willard Hartman (YPMNH), and Olga Hartman (AHF). The following people were of assistance during visits to their Museums by helping locate specimens and providing working space: Olga Hartman and Kristian Fauchald, of the Allan Hancock Founda- tion, during my visit in September 1968; R. Tucker Abbott, of the Academy of Natural Sciences in Philadelphia, during my visit in October 1968. On numerous occasions, Keith H. Woodwick of Fresno State College, Fresno, California, provided working space, offered advice and allowed me to examine his extensive collections of Polydora from California. Additional collections were received from David Grant and Katherine D. Hobson of the Systematics- Ecology Program, Woods Hole, Massachusetts (SEP); Anders Eliason of the Naturhistoriska Museet, GSteborg, Sweden (NMGS); Francois Rullier of the University Catholique d'Angers, France; Luden Laubier, Laboratoire Arago, Banyuls-Sur-Mer, France; P. Korringa, Rijksinsti- tuut voor Visserij-Onder, Bergen-Op-Zoom, The Netherlands; John W. Evans, Memorial University of Newfoundland, St. John's, Newfoundland; John D. Davis, Smith College, Northampton, Massachu- setts; and Sarah A. Haigler, Virginia Institute of Marine Sdence, Gloucester Point, Virginia. Specimens of Polydora concharum collected by the writer in August 1968 aboard the R/V Hero off West Greenland are also included. Complete results of the Hero cruise will be published later. Assistance rendered by Mrs. Crystal M. Blake, Barbara Hilton, Thomas L. Richards, Michael A. Mazurkiewicz, David Dean and George S. Noyes is gratefully acknowledged. The contents of this paper represent portions of a dissertation submitted in partial fulfillment of the degree of Doctor of Philosophy, University of Maine, Orono, Maine. This study was supported in part by a University of Maine Graduate Fellowship, National Sdence Foundation Summer Fellowship, N.S.F. Graduate Traineeship, and N.S.F. Grant GB-4892. Genus Polydora Bosc, 1802 TYPE-SPECIES.?Pofydora cornuta Bosc, 1802. DIAGNOSIS.?Prostomium rounded or bifid an- teriorly, extending back as a caruncle. Eyes present or absent. First setigerous segment with or without notosetae. Setiger 5 greatly enlarged, with spedal- NUMBER 75 ized spines and companion setae arranged in a single curved row. Posterior notosetae may include spe- cialized spines or hooks as well as capillaries. Hooded hooks usually begin on setiger 7. Branchiae begin on segments posterior to setiger 5 and are not fused to the notopodial lobes. Pygidium enlarged, saucer-like or divided into lobes. The following species of Polydora are covered in this report (synonyms in parentheses; those marked with an asterisk are new synonyms): 1. P. ligni Webster, 1879b (?P. littorea Verrill, 1881. Nomen nudum) (P. amarincola Hartman, 1936) 2. P. webiteri Hartman, 1943 (P. caeca.?Webster. 1879a.?Not Oersted. 1843) S. P. aggregata Blake, 1969b 4. P. tetrabranchia Hartman, 1945 5. P. caulleryi Mesnil, 1897 (P. carazzi Mclntosh, 1909) (P. brachycephala Hartman, 1936) 6. P. quadrilobata Jaoobi, 188S (?P. tubifex, Verrill, 1885b) (P. littorea.?Hartman, 1944) 7. P. colonia Moore. 1907 (?P. ancistrata Jones, 1962) (P. hoplura inhaca Day, 1957) 8. P. commensalis Andrews, 1891a (P. ciliata brevipalpa Zaii. 1933) 9. P. concharum Verrill, 1880 10. P. socialis (Schmarda, 1861) (Leucodore socialis Schmarda, 1861) (P. socialis plena Berkeley and Berkeley, 1936) (?P. gracilis Verrill, 1880) 11. P. anoculata Moore, 1907 12. P. hartmanae new spedes (P. anoculata.?Hartman, 1945.?Not Moore, 1907) Key to the Species of Polydora from the East Coast of North America (Figure numbers in parentheses) 1. Hooded hooks begin on setiger 7 2 Hooded hooks begin on setigers 10-17. Palps very short (Ha). Modified spines of setiger 5 with long lateral sheaths (116, c) P. commenmUs (p. 17) 2. Hooded hooks with a constriction on the shaft (2d,e) 3 Hooded hooks without a constriction on the shaft (8/) 6 S. Nuchal tentacle present (la) P. Upd (p. 5) Nuchal tentacle absent 4 4. Modified spines of setiger 5 bifid, with subterminal collars (lOd-f). Posterior notopo- dial hooks present (lOh-k) P. coUmim (p. 15) Modified spines of setiger 5 falcate, with lateral accessory structures. Posterior notopodial hooks absent. 5 5. Modified spines of setiger 5 with lateral sheaths or flanges (36-g). Bores in calcareous stniaures. P. websteri (p. 6) Modified spines of setiger 5 with weakly developed lateral shelves or knobs (46-?). Forms mat of tubes on rocky shores. P. mggregtlm 6. Posterior notopodial spines present, awl-shaped (8g; 9;) Posterior notopodial spines absent 7. Modified spines of setiger 5 large, falcate, with pectinate or bushy tops (8c, d) P. caulleryi Modified spines of setiger 5 bifid, or with bushy tufts between the teeth (9e, g) P. qusdrOobatm 8. Modified spines of setiger 5 bifid, or with well developed lateral teeth or sheaths. Modified spines of setiger 5 simple, falcate or with a weakly-developed accessory thelt 11 9. Branchiae begin on setiger 7. Modified spines of setiger 5 bifid, with two large teeth (Jb, c) P. tetrmbrmnchU (p. 10) Branchiae begin on setigers 9-12. Modified spines on setiger 5 falcate, with lateral sheath or tooth 10 10. Modified spines of setiger 5 falcate, with lateral sheath or tooth and fine hairs in concavity (166-A). Branchiae begin on setigers 11-12 P. hartmmnme (p. 25) Modified spines of setiger 5 falcate, with lateral sheath or tooth not covered with fine hairs (15fr-i). Branchiae begin on setigers 9-10. P. anocuUtm (p. 25) 11. Modified spines of setiger 5 falcate, with weakly-developed lateral shelf (126-d). Pygidium with four unequal lobes (12;). Caruncle extending no further than setiger (p. (p- (p- 8) 7 8 11) 13) 9 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY Ficuu l.?Polydora Kgni: a, Anterior end, dorsal view; b, setiger 1, anterior view; c, posterior end, dorsal view. NUMBER 75 5. Larval pigment retained only in juveniles. Posterior capillary notosetae of three sizes P. concharum (p. 20) Modified spines of setiger 5 falcate, with subterminal boss (146). Pygidium with 1-3 lobes (13c, d). Caruncle extending to setigers 4-9. Larval pigment retained on most adults. Posterior capillary notosetae of one size P. sodattt (p. 20) A FIGURE 2.?Polydora ligni: a, Companion seta of setiger 5; b, c, heavy spines and companion setae of setiger 5; d, e, hooded hooks. Polydora ligni Webster FIGURES 1-2 Polydora ciliatum Agassiz, 1867, pp. 323-330, pis. 9-10.? Verrill, 1873, pp. 51, 309, 345, 364, 453. 603, pi. 14. fig. 78. Not Johnston, 1838. Polydora littorea Verrill, 1881, p. 301 (nomen nudum).? Hartman, 1944, pp. 336, 340, pi. 18: fig. 10, not pi. 18: fig. 9. Polydora ligni Webster, 1879b, p. 119. pi. 5: figs. 45-47.? Webster and Benedict, 1884, p. 729.?Sdderstrom, 1920, pp. 265-267. figs. 170-174.?Cowlcs, 1930, p. 341.?Berkeley and Berkeley, 1936. pp. 471-472; 1952. p. 19. figs. 31-33; 1954. p. 464.?Friedrich, 1937, pp. 345-347, figs. 5-6; 1938. p. 133. ?Rioja, 1943, pp. 232-244; 1947, p. 207.?Mortenson and Galtsoff. 1944, pp. 164-165.?Graham and Gay, 1945, p. 379. ?Hartman, 1941, pp. 309-310, figs. 47-49; 1945, p. 32; 1951. p. 82; 1954, p. 10; 1961, p. 29; 1969, pp. 137-138, figs. 1-6.? Hartman and Reish, 1950, p. 28.?Smidt, 1951, pp. 63-65 ? Hannerz, 1956, pp. 106-111, figs. 37-38.?Jones. 1961, p. 266.?Eliason, 1962, p. 52.?Reish, 1963a, pp. 25, 30; 1963b, pp. 265-270; 1964a. pp. 202-207; 1964b, pp. 86-92.?Wells and Gray, 1964, p. 73.?Galtsoff, 1964, p. 421-425.?Cory, 1967, pp. 71-89.?Berkeley, 1968, p. 560.?Blake, 1969b, pp. 4-10, figs. 1-4. Polydora amarincola Hartman, 1936, p. 49, figs. 6-10. MATERIAL EXAMINED.?Maine (Cobscook Bay; Lamoine Beach; Penobscot River; Damariscotta River; Sheepscot River), Massachusetts (Woods Hole; West Falmouth; Hadley Harbor), Connecti- cut (Noank), New Jersey (Raritan Bay, coll. D. Dean), North Carolina (Beaufort). DESCRIPTION.?Largest specimens measure 32 mm in length and have up to 80 segments. The pro- stomium is bifurcated and flares laterally. There are four eyes, arranged as a trapezoid. A nuchal ten- tacle is present on the caruncle at about the level of setiger 1. The caruncle continues posteriorly as a narrow ridge through setiger S. A ciliated groove lies on either side of the caruncle (Figure la ) . Setiger 1 has well-formed parapodial lobes but contains only capillary neurosetae (Figure 16). Setigers 2-4 and 6 contain fascicles of finely winged capillary setae. After setiger 6 the number of notosetae gradually diminishes, until in posterior segments only a tuft of long capillary setae remains. Bidentate hooded hooks begin on setiger 7 (Figure 2d, e). There are up to 15 hooks per neuropodium and no accompanying capillary setae. Hooks of the same neuropodium may have the main fang either at a right angle to the shaft or bent to about 70?. A constriction and curve are prominent on the shaft. A striated hood covers the teeth. Setiger 5 is modified to include a row of heavy spines with closely adhering companion setae (Fig- ure 2a-c). The former have a small accessory tooth which may be considerably worn. The latter have a delicate feathery end; when depressed, the tip is seen to be forked. Branchiae are long and thin, beginning on setiger 7 and continuing to near the end of the body. The pygidium is a large, flaring cup with a dorsal gap (Figure lc ) . Color in life is light tan with red blood vessels. Except for lateral black pigment spots often present in juveniles, the body is unpigmented. SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY REMARKS.?Polydora littorea Verrill is herein considered synonymous with P. Ugni. The species was named by Verrill (1881) but was never de- scribed. Verrill erected the name littorea to replace the P. ciliatum of Agassiz (1867), which he con- sidered to be distinct from the European P. ciliata. The figures of the adults and larvae of Agassiz's P. ciliatum appear to be P. Ugni. Plate 18, Figure 10, was cited by Verrill but not published until Hart- man (1944) issued Verrill's plates. A nuchal ten- tacle is not evident and Figure 10 is indeterminable m other respects. Figure 9, however, cited by Hart- man as P. littorea, is not the same species. It re- sembles P. quadrilobata Jacobi both in general shape and structure of the pygidium. The species in North America most closely re- lated to P. Ugni is P. nuchalis Woodwick from California (Woodwick, 1953), both species having a nuchal tentacle. Polydora nuchalis differs in that no accessory structure is present on the modified spines of setiger 5. Further, the companion setae are of different structure. ECOLOGY.?Polydora Ugni is a common inhabi- tant of estuaries in North America. It constructs fragile tubes of silt on tidal flats or attached to shells, rocks and wood. The species does not bore into calcareous structures. Galtsoff (1964) reported that on several occasions in Delaware Bay the re- production of P. Ugni was so rapid that nearly every live oyster of the affected area was killed by a de- posit of mud several inches thick. Mud particles sus- pended in the water are caught on the worm's palps and carried toward the head by ciliary action. The silt is then formed into a tube in the manner de- scribed for other species by Hempel (1957b) and Dorsett (1961). DISTRIBUTION.?East and west coasts of North America, Gulf of Mexico and northern Europe. Polydora websteri Hartman FlGUKE S Polydora ciliata.?Procter, 19SS, p. 142.?Kavanaugh, 1941, pp. 32-34.?Lunz, 1940. p. S10; 1941, pp. 273-283.?Abbott, 1946, pp. 15-18, pi. 3.?Medcof, 1946, pp. 498-515.?Plaine, 1952. pp. 121-123. fig. 1.?Turner and Hanks, 1959. pp. 109-111.?Landers, 1967, pp. 63-66, figs. 1-2. Not Johns- ton, 1838. Polydora caeca.?Webster, 1879a, pp. 252-253, pL 9: figs. 119-122.?Andrews. 1891a, p. 291. Not Oersted. 1843. Polydora websteri.?Hartman. 1943, pp. 70-72, fig. 1; 1945, p. 33; 1951, pp. 81-82; 1954. p. 415; 1961. pp. 99-100, pis. 16-17; 1966b, p. 223; 1969, pp. 151-152, 4 figs.?Loosanoff and Engle, 1943, pp. 69-78.?Grice, 1951. pp. 1-10.?Mackin and Cauthron, 1952. pp. 14-24.?Menzel, 1955, p. 112.? Hopkins, 1958, pp. 268-277, figs. 1-6.?Owen, 1957, pp. 35-46.?Rioja, 1960, p. 304.?Mackenzie and Shearer, 1961, pp. 105-111?Wells and Gray, 1964, p. 73.?Galtsoff. 1964, pp. 421-425.?Forbes, 1966, pp. 280-281.?Davis, 1967, pp. 67-72, figs. 2-3.?Evans, 1969, pp. 775-782.?Blake. 1969a, pp. 814-815. fig. 2; 1969b, pp. 10-16, figs. 5-11.?Haigler, 1969. pp. 821-828, figs. 1-3. MATERIAL EXAMINED.?Maine (Lamoine Beach; Damariscotta River; Boothbay Harbor region. Ocean Point), Newfoundland (St. Mary's Bay, 10- 20 meters, coll. J. W. Evans), Quebec (Gaspe Peninsula, coll. J. D. Davis), Connecticut (Milford, Type AHF 1569; Noank), Virginia (Gloucester Point, coll. J. L. Simon), North Carolina (Beau- fort), South Carolina (Charleston, coll. M. Cham- berlain), California (Morro Bay, coll. K. H. Woodwick). DESCRIPTION.?Polydora websteri is a small slen- der species, measuring up to 20 mm in length and having about 100 segments. In life it is light tan with red branchiae, palps and blood vessels. Body pigmentation is variable, absent in some specimens while others have dense anterior and posterior black pigment. A prostomial anterior incision may be distinct, weak or absent (Figure 3a). Specimens from New England and eastern Canada tend to have a weakly incised or rounded prostomium, while in specimens from the middle Atlantic and southern states, it tends to be more strongly incised. Postlarval forms from Maine have a rounded prostomium. The caruncle terminates bluntly on setiger 2. Eyes are present or absent; if present, there are four, dis- posed as in Figure 3a. Setiger 1 has only capillary neurosetae and ele- vated notopodial lobes. Setigers 2-4 have blunt notopodial lobes with fascicles of winged capillary setae. The neuropodial lobes are greatly reduced and contain winged capillary setae. Setiger 6, and those immediately following, contain dorsal fascicles of winged capillary setae. The number of setae per notopodium gradually diminishes in the middle segments. In far posterior segments, only three or four winged capillary setae are found in each noto- podium. On setiger 7, bidentate hooded hooks re- place the neuropodial winged capillary setae. The hooks number about six in a series and have the NUMBER 75 FicintE S.?Poiydora websteri: a, Anterior end, dorsal view; b-g, heavy modified spines of setiger 5 showing different views and degrees of wear; h, pennoned companion seta of setiger 5; i, superior dorsal winged seta of setiger 5; /, hooded hook; k, posterior end, dorsal view; I, pygidium, posterior view. 8 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY main fang at a right angle to the shaft. Each hook has a constriction on the shaft (Figure 3;*). Setiger 5 is twice as large as either 4 or 6; the parapodia contain a row of heavy modified spines (Figure 36-g) alternating with pennoned compan- ion setae (Figure 3/t) . One or two superior dorsal, winged setae (Figure 3t) and a ventral fascicle of short winged setae are present. The modified spines are falcate and have a lateral flange or sheath. Fig- ure Sb-g illustrates different views and degrees of wear of these spines. Branchiae begin on setiger 7. They are small at first, reaching full size on setigers 11 and 12 and are absent from posterior segments. The pygidium is cup-shaped with a distinct dor- sal notch (FigureS*,/). REMARKS.?During the course of these studies, no specimens of P. ciliata have been identified from eastern North America. Hartman (1945) has al- ready suggested that P. ciliata may not even occur in eastern North America. On this basis earlier records are herein referred to P. websteri. Both Blake (1969a) and Haigler (1969) reported that the hooded hooks of P. websteri begin on setiger 7 and not 8, as previously reported by Hartman (1943). ECOLOGY.?The species is best known as a pest of commercial oysters, but is abundant in numerous intertidal and shallow water calcareous substrates, especially in eastern North America. In Maine the species has been found boring into shells of the scallop Placopecten magellanicus and the gastro- pods Littorina littorea, Thais lapillus, Crepidula fornicata, and Lunatia heros. DISTRIBUTION.?East coast of North America from Quebec and Newfoundland to Florida, Gulf of Mexico, California, Oregon, and Hawaii. Polydora aggregate Blake FICUBES 4-6 Blake, 1969b, pp. 16-21, figs. 12-15. (Description of larvae.) MATERIAL EXAMINED.?Maine (Lamoine Beach State Park; Boothbay Harbor region, Ocean Point). The holotype (USNM 42246) and 10 paratypes (USNM 42247-8) have been deposited in the United States National Museum. Additional para- types have been deposited in the Allan Hancock Foundation. DESCRIPTION.?This is a small species measuring 6-8 mm in length and having 35-40 segments. The body has a characteristic reticulate, dark-brown pig- ment pattern of varying intensity on the peristo- mium and first six to eight segments (Figure 4a). The caruncle has two lateral clear areas which in life are seen to contain ciliary grooves. Posterior segments are generally not darkly pigmented, al- though the pygidium is often rimmed with black. The prostomium is bifid, although in some pre- served specimens the anterior end curves downward and appears rounded when seen from above. The caruncle ends abruptly on setiger 2. There are four eyes, the anterior pair cup-shaped and the posterior pair oval (Figure 4a). Setiger 1 has only neurosetae, although a well developed notopodial lobe is present. Setigers 2-4 have blunt notopodial lobes with long and short, winged capillary setae. The neuropodial lobes are small, with those of setigers 2-4 becoming progres- sively shorter. Beginning on setiger 6, the notopodia contain fascicles of long and short, winged capillary setae. The number of setae per fascicle gradually diminishes in the middle region, and in posterior segments only a few long, thin, capillary setae are present. The winged capillary neurosetae are re- placed by bidentate hooded hooks beginning on setiger 7 (Figure 4g). Each hook has a constriction on the shaft. Five or six hooks are present on most segments, although fewer are found in posterior seg- ments. Setiger 5 is about twice the size of preceding seg- ments. The parapodia contain a curved row of heavy spines (Figure 4b-e) alternating with pen- noned companion setae (Figure 4/) . Tufts of winged setae are located above and below the heavy spines. The heavy spines are falcate, with a small lateral accessory structure appearing as a small elevation, cleft on one side and rounded on the other. This structure can be classed neither as a tooth nor flange. Deeper unworn setae have a long tapered mucron on their tips. Branchiae begin on setiger 7 and are absent from posterior segments. The pygidium is cup-shaped with a wide dorsal gap (Figure 4h). REMARKS.?The name Polydora aggregata was prematurely introduced in a larval study (Blake, 1969b). The adult description is given here for the first time. NUMBER 75 Ficuu 4.?Pofydora aggregata: a, Anterior end, dorsal view; b-e, heavy modified spines of setiger 5 in different views; /, pennoned companion seta of setiger 5; g, hooded hook; h, posterior end, dorsal view. SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY ? F K U U 5.?Lamoine Beach, Maine: a, Foregrouod ihows a ?olid mat of P. aggregate tubes; b, clustered tubes of P. ag- gregat* between rocks and broken shells of Mytilus edulis. The snail is Uttorina tittorem. Polydora aggregate is closely related to P. limicola (Annenkova, 1954). The two spedes are similar in habitat and morphology. They differ, however, in that P. limicola has a small, triangular tooth on the side of the modified spines of setiger 5, whereas P. aggregate has the structure described above. ECOLOGY.?At Lamoine Beach, P. aggregata oc- curs in dense mats on and between rocks, exposed at low tide (Figure 5a). The mats are so thick in some areas they appear to stabilize the beach. At the lowest tide levels, the dense mats of tubes give way to tube dusters or dumps, which are attadied to rocks, shells, or algae (Figures 5b; 6a, b). Oc- casionally, the orbiniid Naineris quadricuspida (Fabricius), phyllododds, chironomid larvae and nematodes are associates in the mats of tubes. At Ocean Point the spedes was found among holdfasts of Chondrus crispus. Ficuu 6.?Lamoine Beach, Maine: a, Rock with tube duster) of P. aggregata; b, tube dusters of P. aggregata. DISTRIBUTION.?Maine, on semiexposed rocky shores. Polydora tetrabranchia Hartman FlCUKE 7 Hartman, 1945, p. 54, pi. 1: figs. 7-10. MATERIAL EXAMINED.?North Carolina (Beaufort, Types, AHF N1563). DESCRIPTION.?This is a small spedes, measuring about 6 mm in length and having about SO seg- ments. The prostomium has a weak anterior inci- sion. There are no eyes or nuchal tentacle. The caruncle extends to setiger 1. In the spedmens examined, notosetae were not seen on setiger 1, although Hartman (1945) indi- cated they were present Slender, fine, winged, NUMBER 75 11 FicmtE T.?Polydora tetrabranchia: a, Hooded hook; b, c, heavy modified spines of setiger 5. capillary notosetae occur in setigers 2-4, 6, and suc- ceeding segments. The neuropodia of setigers 1-4 and 6 contain winged capillary setae. Hooded hooks begin on setiger 7 with no more than three hooks per segment. They have the main fang bent at nearly a right angle to the recurved shaft (Figure 7a). The modified spines of setiger 5 are distally bifid, with two large teeth (Figure 7b-c). There are usually two or three spines in each parapodium, ac- companied by slender pennoned setae. The pygidium is a simple, collar-like ring. Branchiae begin on setiger 7 and usually number only four pair. DISTRIBUTION AND ECOLOGY.?Polydora tetra- branchia bores in shell fragments and is known only from Beaufort, North Carolina. Polydora cauUeryi Mesnil 8 Polydora cauUeryi Mesnil, 1897. pp. 88-89, pi. 3:figs. 12-14.? Southern, 1914, p. 104.?Mclntoah, 1915, pp. 210-212, pi. 100:fig. 8. pi. 106:fig. 5.?Fauvel, 1927, pp. 54-45, fig. 19/-/I.?Hartman, 1951, pp. 83-84; 1965, pp. 149-150. pi. 28:figs. b-d.?Pettibone, 1954, pp. 282-284, fig. 32*^.? Hannerz. 1956, pp. 123-126, figi. 43-44.?Eliason, 1962, p. 53.?Blake. 1969b. p. 52. fig. 38. Polydora carazxi Mclntoth, 1909, pp. 172-173, pi. 5: figs. 4-6. Polydora brachycephala Hartman, 1936, pp. 48-49, figs. 3-5; 1941, p. 307; 1944, p. 238; 1954, p. 10; 1961, p. 29; 1969, figs. 1-3.?Filfce. 1958, p. 190.?Reish, 1959. pp. 38, 61, 67. 70, 75; 1964b, pp. 86-92. MATERIAL EXAMINED.?Maine (Damariscotta River), Connecticut (Noank), Massachusetts (Cape Cod Bay, coll. SEP), California (Dillon Beach, coll. K. H. Woodwick), Sweden (Kattegatt, NMGS). DESCRIPTION.?Largest specimens measure 50 mm in length, are 2.2 mm wide, and have over 150 seg- ments. Rust colored pigmentation has been observed dorsally on the anterior segments of some sped* mens. The color in life is light tan with prominent red blood vessels. A female which had recently spawned was yellow. The anterior end is prominently wedge-shaped (Figure 8a). The prostomium is deeply incised anteriorly with two rounded lobes. The caruncle extends back through setigers 3 or 4. Eyes are pres- ent or absent; if present, a small oval pair lies pos- terior to a more widely spaced cup-shaped pair. Palps are slender. Setigers 1-4 have flattened, bladelike parapodial lobes, the notopodial lobes being somewhat auricu- late (Figure 8b, h, t) . The notosetae of setiger 1 are short, slender capillaries (Figure Sh) while those of setigers 2-4 are long, slender, winged capillaries (Figure 8t). Setiger 6 and succeeding segments contain notopodial fascicles of long, winged capillary setae. In far posterior segments, the capillary notosetae are replaced by a curved row of stout, sharply pointed spines (Figure 8g). Neuro- setae of setigers 1-4 and 6 are long, slender capil- laries. Bidentate hooded hooks begin on setiger 7, accompanied by a tuft of capillary setae. The hooks are recurved, with a main fang at nearly right angles to the shaft and have a smaller secondary tooth (Figure 8/). There are about 13 hooks on anterior setigers, the number gradually diminishing until in far posterior segments only about 5 can be found in a parapodium, accompanied by one or two capillary setae. Setiger 5 is large, with a heavy musculature, overlapping setiger 6. The characteristic large, bushy-topped modified spines are arranged in an 12 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY FICUBE S.?Pofydora caulleryi: a, Anterior end in dorsal view; b, anterior end, lateral view; c, heavy modified spine and companion capillary seta of setiger 5; d, heavy spine of setiger 5; e, dorsal seta of setiger 5; /, hooded hook; g, posterior notopodial spine; h, parapodium of setiger 1, anterior view; i, parapodium of setiger 3, anterior view; /, pygidium, posterior view. NUMBER 75 IS anteriorly elevated semicircle, accompanied by a few very fine, capillary companion setae (Figure 8c, d). A vertical row of dorsal, winged setae is located anterior to and above the heavy spines (Figure 8e). The neurosetae consist of a tuft of curved capillary setae. Branchiae begin on setiger 7, being small at first and reaching full size on setiger 11. They are absent from the last third of the body. The pygidium consists of four equal lobes (Fig- ure 8;). REMARKS.?The synonymy of Polydora caulleryi and P. brachycephala, proposed by Pettibone (1954), is herein supported. California specimens possess posterior spines and agree in other respects with New England representatives. Specimens from the Kattegat (Sweden) did not differ from Amer- ican material. Pelagic larvae obtained in New Eng- land waters agree well with those described by Hannerz (1956) from Sweden (Blake, 1969b). DISTRIBUTION AND ECOLOGY.?Polydora caulleryi occurs in Europe, New England, Alaska and Cali- fornia. It ranges from low intertidal (Pettibone, 1954; personal observations) to continental shelf depths of 200 meters (Hartman, 1965). Polydora quadrilobata Jacobi FlCUKE 9 Polydora quadrilobata Jacobi, 1883, pp. 1-37, 2 pis.?Mesnil, 1897, pp. 87-88, pi. 3: figs. 9-11.?Mclntosh, 1909, pp. 170-172. pi. 5:figs. 2-3; 1915. p. 236, pi. 98:figs. 13, 17. pj. 106: fig. 4.?Eliason, 1920, pp. 45-46; 1962, p. 52.?Fauvel, 1927, p. 52, figs. 18 /-n.?Annenkova. 1931, pp. 203-205; 1932, p. 139, figs. 6-9.?Procter, 1933. p. 142?Friedrich, 1938, p. 133.?Berkeley and Berkeley. 1943. p. 130; 1954. p. 464.?Uschakov, 1955. p. 278. figs. 94a-d; 1965, p. 250* fig. 9ia-d? Rasmussen, 1956, p. 64.?Hannerz, 1956, pp. 122-123.?Hempel, 1957a, pp. 276-278; 1957b, pp. 100-135, fig. 5.?Hartman, 1961, p. 29; 1969, pp. 145-146, 4 figs.? Khlebovitsch, 1961, p. 201.?Jones, 1961, p. 288.?Blake, 1969b, pp. 37-51, figs. 27-37. Polydora tubifex Verrill, 1885b, pp. 438-439.?Sumner et al., 1913, p. 624. Polydora littorea.?Hartman, 1944, pi. 18: fig. 9. Not Verrill. 1881. MATERIAL EXAMINED.?Maine (Cobscook Bay; Lamoine Beach; Damariscotta River), Connecticut (Noank), Sweden and Denmark (NMGS). DESCRIPTION.?Individuals measure up to 15 mm in length and have about 70 segments. The anterior end has brown, reticulated pigmentation dispersed on the sides of the peristomium and on the anterior and posterior margins of setigers 1-7. The intensity of the pigmentation varies between individuals, but has been found in some degree on all specimens examined from New England. The color in life is light tan,with red blood prominent in the bran- chiae, palps, and blood vessels. The prostomium is distinctly bifid on its anterior margin and continues posteriorly as an indistinct caruncle to about the middle of setiger 3 (Figure 9a). Four to six eyes are present in nearly a straight transverse line. The two outer pairs may be fused into a single mass. The palps are long and flexible, with a prominent ciliated groove. Setiger 1 has capillary notosetae and neurosetae (Figure 96). Setigers 2-4 and 6 have neuropodial fascicles of finely-winged, capillary setae. The notosetae of setigers 2-4 are winged capillaries (Figure 9c). Bidentate hooded hooks begin on setiger 7 (Figure 9i) and are accompanied by two or three fine capillary setae. There are usually no more than three or four hooks in any one para- podium. The notosetae of setiger 6 and succeeding setigers include fascicles of winged capillaries. The number of capillary notosetae diminishes in the middle segments, and they are completely replaced by heavier, sharply pointed spines in far posterior segments. These spines are disposed in a semicir- cular arrangement and give the posterior end a spinous appearance (Figure 9j-k). Setiger 5 lacks parapodial lobes and has three types of setae (Figure 9d) . The dorsal setae consist of a horizontally curved row of heavy, modified spines (Figure 9e, f) and a vertical curved row of shorter limbate setae (Figure 9g, h) located at the anterior end of the horizontal row, resulting in a J-shaped arrangement when the specimen is viewed from the sides. The heavy spines have a unique structure (Figure 9e, f). The distal end curves and is bifurcated at the tip, forming two nearly equal teeth between which is located a fine bushy tuft. When worn, the teeth and tufts may be reduced. A small fascicle of winged setae is located ventral to the modified spines. Branchiae begin on segment 7. They are short and stubby, not reaching full size until setiger 12 or 13. They are absent from the posterior third of the body. The pygidium has four subequal lobes (Figure 14 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY FicwtE 9?Potydora quadrilobata: ?, Anterior end, donal view; b, setigrr 1, anterior view; c, ?etiger 3, anterior view; d, anterior end, lateral view; e, f, heavy ipincs of tedger 5; g, h, limbate ?etae of wtiger 5; i, hooded hook; /, posterior notopodial spinet; k, posterior end, donal view. NUMBER 75 15 9k). When seen on living specimens under reflected light, they are white. REMARKS.?Soderstrom (1920) considered P. quadrilobata to be a juvenile of P. caulleryi. This synonymy has not been supported by subsequent workers. There is no question, however, that the two species are closely related. A comparison be- tween European and American specimens of P. quadrilobata revealed no morphological differences. Polydora tubifex Verrill (1885b) is here referred to P. quadrilobata. The original description was incomplete and no setal details were given. The description as given by Verrill, would apply to several species. The eyes were said to be nearly in a straight line; the "caudal appendage" was small and consisted of four short lobes; and the anterior segments had "greenish-black" marks between seg- ments. Polydora quadrilobata is the only spedes un- covered to date in New England to which this combination of characteristics agrees. J. Percy Moore (MS) described and figured P. tubifex. His description, however, differs markedly from that given' by Verrill. Moore's spedmen in the Academy of Natural Sdences of Philadelphia was examined. Consisting of a single specimen in two parts, it agreed with Moore's description but dearly be- longed to Polydora commensalis. ECOLOGY.?Polydora quadrilobata is a common spedes in New England. It is easily collected inter- tidally at Lamoine Beach and Cobscook Bay State Parks (Maine) where it occurs in sandy mud. At Noank, Connecticut, the spedes is abundant off the pier at the Marine Research Laboratory of the University of Connecticut, where it occurs in silty mud. The tubes stand erect in the substrate, are rusty colored, and are formed of fine silt. The spedes is usually associated with other spionids, maldanids, and sabellids. It was dredged in 10-12 meters in Clark Cove on the Damariscotta River, Maine, where it occurs in muddy substrate and is associated with Polydora socialis and a sabellid, Laonome sp. DISTRIBUTION.?Europe, eastern Canada, New England, southern California, western Pacific. Polydora colonia Moore FIGURE 10 Polydora colonia Moore, 1907. pp. 199-201, pi. 15: figs. 18-23. ?Sumner, et al., 1915, p. 625.?Annenkova, 1938, p. 178.? Hartman, 1945. pp. 32-33.?Uschakov, 1955. p. 273, fig. 94g-;; 1965. p. 252. fig. 94g-;. Polydora ancistrata Jones, 1962, pp. 185-187, figs. 55-65. Polydora hoplura inhaca Day, 1957, p. 99, fig. 6ft-/; 1967. p. 468, fig. 18.2n. MATERIAL EXAMINED.?Massachusetts (Holotype, ANSP 2S24, Woods Hole; Hadley Harbor), North Carolina (Beaufort, AHF 1546), Jamaica (para- types of P. ancistrata, in the collections of Keith H. Woodwick). DESCRIPTION.?Polydora colonia is a small spedes, measuring up to 5 mm in length and having about 35 segments. The prostomium appears anteriorly rounded in preserved specimens (Figure 10a). In life it is seen to have a slight anterior bifurcation, which is turned under when preserved. The car- uncle extends to setiger 2. The palps are short and reach posteriorly to setigers 8 or 9. Eyes and other body pigmentation are lacking (Figure 10a). Setiger 1 contains only capillary neurosetae. The neurosetae of setigers 2-4 and 6 are short, winged capillaries. Hooded hooks begin on setiger 7. They are bidentate with a large fang and small secondary terminal tooth; the shaft has a constriction (Figure lOg). The notosetal fasddes of setigers 2-4, 6 and succeeding segments contain long capillary setae. The notosetae diminish in number in posterior segments and, in the last six or seven segments, are accompanied by one or two specialized hooks (Figure 10/). The more anterior hooks are slender (Figure \Qh, t) and not as curved as those which follow (Figure 10;, k). The latter are recurved and falcate. Setiger 5 is larger than the preceding and suc- ceeding setigers. The modified spines are bifid, with two unequal teeth and a subterminal collar extending halfway around the spine (Figure lOd-f). Alternating with the three or four large spines are pennoned companion setae (Figure 10c). There is a small bundle of slender, winged, dorsal setae lo- cated above the modified spines (Figure 106). A ventral tuft of three or four winged capillary setae is also present. Branchiae begin on setiger 7, continuing through the middle segments; they are absent from the pos- terior one-half of the body. They are small and do not overlap dorsally (Figure 10a). The pygidium consists of a flattened disk, the sides of which gradually merge into the last seg- ment, leaving a wide dorsal gap (Figure 10/). 16 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY FIGUK 10.?Potydora coIonia: a, Anterior end, dorsal view; b, superior dorsal seta of setiger 5; e, pennoned companion seta of setiger 5; d-f, heavy spines of setiger 5 in different views, g, hooded hook; h-k, posterior notopodial hooks representing a sequential change in structure on progressively more posterior segmenu; /, posterior end, dorsal view. NUMBER 75 17 REMARKS.?Polydora ancistrata Jones is herein referred to P. colonia Moore. The differences be- tween the two species, as discussed by Jones (1962), have not been substantiated by recent, well- preserved material collected from the type locality of P. colonia (Woods Hole, Massachusetts). Poly- dora hoplwra inhaca Day (1957) also belongs in the synonymy, as suggested in a footnote by Day (1967). The species most closely related to P. colonia is P. spongicola Berkeley (emend. Woodwick, 1963). The specimens of the latter from California and MazatlAn, Mexico (courtesy of Dr. Keith H. Wood- wick) were examined by the writer. In P. spongicola the specialized spines of setiger 5 have a large ter- minal tooth surrounded by a flange, while in P. colonia there are two teeth surrounded by a collar. ECOLOGY.?Polydora colonia inhabits tubes within sponges, forming compact, soft masses at- tached to pilings and rocks in areas of water move- ment. DISTRIBUTION.?Massachusetts to North Carolina, Jamaica, South Africa, northwest Pacific. Polydora commensalis Andrews FlCUftE 11 Polydora commensalis Andrews, 1891a, pp. 25-35, 2 pis.? 1891b, pp. 291-292, pi. 15: fig. 27.?Cowles, 1930, p. 344.? Berkeley and Berkeley, 1936, pp. 469-471; 1952, pp. 18-19, figs. 29-30.?Annenkova, 1938, p. 178, fig. 14?Hartman, 1941, p. 308; 1945, p. 32; 1961, p. 29; 1969, pp. 133-134, 4 figs.?Rioja, 1943, p. 229.?Hartman and Reish, 1950, p. 28.?Uschakov, 1955. p. 272, fig. 93; 1965, p. 250, fig. 93 ? Wells and Gray, 1964, p. 73.?Hatfield, 1965, pp. 356-368, figs. 1-5.?C. Berkeley, 1968, p. 560.?Blake. 1969a, pp. 815-816, fig. 3; 1969b, pp. 21-24. figs. 16-18. Polydora sp. Berkeley, 1927, p." 420. Polydora ciliata brevipalpa Zaks, 1933, p. 129. MATERIAL EXAMINED.?Maine (Damariscotta River), Massachusetts (Cape Cod Bay, coll. SEP; North Falmouth, as P. tubifex coll. J. P. Moore, ANSP 1516), Connecticut (Noank), North Caro- lina (Beaufort). DESCRIPTION.?Specimens measure up to 30 mm in length, are 2.5 mm in width, and have over 100 segments. The prostomium is rounded or weakly emarginate on its anterior margin; the entire head is usually contracted into the first setiger (Figure l l a ) . There is no caruncle. Four small eyes are arranged in a nearly straight line. The palps are unusually short and covered with numerous, fine papillae, each with a sensory hair. The color in life is dark tan with a greenish cast on numerous seg- ments. Red blood vessels are prominent. Setigers 1-4 have well developed noto- and neuro- podial lobes with fascicles of long capillary setae (Figure 1 la) . Se tigers 3 and 4 have some additional short capillary notosetae. Bidentate hooded hooks begin on setigers 12-17 (rarely 10-17). The hooks are accompanied by a few capillary neurosetae. The hooks have a main fang surmounted by a shorter secondary tooth (Figure lid). The secondary tooth is reduced in hooks from far posterior setigers and, when viewed from certain angles, is difficult to see (Figure l i e ) . There are 5 or 6 hooks in anterior setigers, increasing to about 13 in middle setigers, and reduced to 4 or 5 posteriorly. The setae of setiger 5 are modified to include a row of large curved spines and a small tuft of capil- lary neurosetae. The heavy spines have a character- istic long lateral flange or sheath (Figure lib, c). Branchiae are long and straplike, with mem- branous margins. They begin on setiger 6 and con- tinue to the end of the body. The posterior end of the animal is dorsoventrally flattened (Figure I If). The anus is surrounded by a ring of small papillae, the number of which is variable, depending upon the age of the animal- In young juveniles 4 papillae may be present, increas- ing to 14 in the largest adults. An intermediate stage with 8 papillae is shown in Figure 11/. REMARKS.?Polydora commensalis deviates widely from other spedes of the genus and should be re- moved from Polydora. However, the writer has knowledge of two undescribed species in California which appear to be closely related to P. commen- salis. Until these new collections are studied in de- tail, the generic placement of P. commensalis had best be delayed. The specimens, identified as P. tubifex Verrill and described by Moore (MS), were examined and found to be P. commensalis (ANSP 1516). Polydora tubifex Verrill (1885b) is here (p. 13) referred to P. quadrilobata. ECOLOGY.?Polydora commensalis is known only from shells containing hermit crabs. In Maine, the mollusc shells include Littorina littorea and Thais lapillus, occupied by the hermit crab Pagurus longicarpus. Shells covered with the hydroid Hydractinia echinata appear to have a higher in- festation of P. commensalis. 18 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY Ficuu 11.?Poly dora commetualis: a. Anterior end, donal view; b, c, heavy spines of se tiger 5; d, e, hooded hooks; /, posterior end, donal view. NUMBER 75 19 Fsscu 12.?Polydora concherum: a, Anterior end, dorsal view; b-d, heavy ipinet of ledger 5 In different view*; e, pennoned companion seta of ledger 5; /, superior dorsal geniculate sett of ?etiger 5; g, setiger 1, anterior view; h, posterior setiger, dorsal view (not to scale); i, posterior end, dorsal view; /, pygidium, posterior view; *, /, hooded books; m, n, winged capillary notosetae. 20 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY DISTRIBUTION.?East and west coasts of North America; North Japan Sea. Polydora conchamm Verrill Ficinti 12 Polydora concharum Verrill, 1880, pp. 174-176.?Webster and Benedict. 1884, p. 729; 1887, p. 737.?Sumner, et al., 1913, p. 624.?Procter, 1933, p. 142?Hartman, 1942, pp. 64-66, figs. 121-123; 1944, pp. 336-340, pi. 19: fig. 10 (not fig. 8).? Evans, 1969, pp. 775-782.?Blake. 1969a, p. 816, fig. 4; 1969b, pp. 32-36, figs. 23-26. Dipolydora concharum.?Verrill, 1881, p. 320; 1885a, p. 688, pi. 43: fig. 186. MATERIAL EXAMINED.?West Greenland (Hero Station 29F) , Newfoundland (St. Mary's Bay, 10-20 meters, coll. J. W. Evans), Maine (Damariscotta River; Gulf of Maine, Types YPMNH 2687), Mas- sachusetts (Cape Cod Bay, coll. SEP). DESCRIPTION.?Polydora concharum is a long, slender spedes reaching lengths up to 140 mm and having over 300 segments. In life it is light tan with red branchiae and blood vessels. Juveniles have been found to retain larval pigment for a time after metamorphosis. In adults the body is largely color- less, although some specimens have been found with a line of dark pigment on the anterior borders of some segments, first appearing in setiger 6. The prostomium is distinctly bifid on the an- terior margin. The caruncle continues posteriorly as a low ridge to the anterior border of setiger 5 (Figure 12a). Eyes are either present or absent; when present, there are four in a trapezoidal ar- rangement. The palps are relatively short in com- parison to the length of the animal and extend posteriorly for only 15 to 20 segments. Setiger 1 has both noto- and neurosetae (Figure 12g). The former are fine capillaries, while the latter consists of winged capillaries. Setigers 2-4 have well developed postsetal lobes and large spreading fascicles of winged capillary noto- and neurosetae. Notosetae of setiger 2 are arranged in two successive rows, while those of setigers 3, 4, 6, and those immediately following have notosetae in three rows, those of each successive row being longer (Figure 12m, n ) . Notosetae from more posterior setigers lack wings; they are represented by three sizes of straight capillary setae (Figure 12h). The winged neurosetae of setigers 2-4 and 6 are ar- ranged in two rows. Bidentate hooded hooks (Fig- ure 12A, /) begin on setiger 7 and are accompanied by two or three capillary setae. Setiger 5 is larger than proceeding and succeeding setigers. The setae are arranged in three bundles: a group of geniculate setae (Figure 12/) located anterior and dorsal to a semicircular row of large heavy spines (Figure \2b-d), alternating with smaller pennoned companion setae (Figure 12e); and a small tuft of ventral winged neurosetae. The heavy spines have a weakly-developed accessory shelf, horizontal to the main shaft. The structure was not observed by Verrill (1880) or Hartman (1942). Branchiae begin on setigers 7-9. They are small anteriorly, reaching full size on setiger 11 or 12 and are absent from the posterior half of the body. The pygidium is small and has four lobes, the dorsal pair being smaller than the ventral pair (Figure 12i,;) . ECOLOGY AND DISTRIBUTION.?Cape Cod, Massa- chusetts, to Newfoundland, in dredged shells and decayed wood; West Greenland. Polydora sociaUs (Schmarda) FIGURES 13, 14 Leucodore socialis Schmarda, 1861, p. 64, figs, a-c, pi. 26: fig. 209. Polydora socialis.?Mesnil, 1896, pp. 193-194, pi. 12: figs. 30-32?Ehlers, 1901, p. 165.?Hartman, 1941, pp. 310-311, pi. 48: figs. 41-42; 1945, pp. 33-35; 1948, p. 37; 1951. p. 83; 1953. pp. 43-44; 1961, p. 29; 1966a, p. 19. pi. 19:figs. 1-2; 1969, pp. 147-148. 2 figs.?Hartman and Reish, 1950, p. 28. ?Rioja, 1943, p. 230.?Hartmann-Schrdder, 1962, pp. 137-138, figs. 167-168; 1965. pp. 209-211, figs. 200-203 ? Wells and Gray, 1964, p. 73.?Long, 1968. p. 348.?Blake, 1969a. pp. 816-817. fig. 5; 1969b. pp. 24-31. figs. 19-22. Polydora socialis plena Berkeley and Berkeley, 1936, pp. 468-469; 1952, p. 22?Reish, 1968, p. 82. ?Polydora gracilis Verrill, 1880, p. 174. MATERIAL EXAMINED.?Maine (Damariscotta River), Massachusetts (Cape Cod Bay, coll. SEP), Connecticut (Mystic River), North Carolina (Beau- fort) , South Carolina (Charleston, coll. M. Cham- berlain), California (Mugu Lagoon, AHF N1417; Cayucos), British Columbia (paratypes of P. socialis plena, USNM 32704), Chile (Lund University Chile Expedition 1947-48, deposited AHF). DESCRIPTION.?Specimens of Polydora socialis from New England measure up to 55 mm in length and have over 400 segments. Individuals examined NUMBER 75 21 Ficuu IS.?Polydora sodalis: a, Anterior end, dorsal view; b, diagram of ledgers 4-7 in lateral view (not to scale); c, pygidium, posterior view; d, posterior end, dorsal view. from North and South Carolina were not as large. Sexually mature females are orange. Larval pig- mentation is retained on most adults (Figure 13a), including both dorsal and ventral bands. The prostomium is deeply notched on its anterior margin. The caruncle extends to setigers 4-9. Eyes are present or absent; when present, they number four to six: a circular pair high up on the pro- stomial ridge, and more anterior second and third pairs, which may be fused laterally on the pro- stomial ridge (Figure 13a). The palps are long and prehensile in life; upon preservation, however, they contract and extend posteriorly only to setigers 12- 15. Setiger 1 has capillary setae in both noto- and neuropodia (Figure 14a). Setigers 2-4, 6 and suc- ceeding setigers have two types of posteriorly di- rected notosetae, an anterior row of short curved capillaries and a posterior row of long, stout capil- laries. In far posterior setigers only a few laterally- directed capillary notosetae are present The neuropodia of setigers 2-4 and 6 have posteriorly directed capillary setae. Bidentate hooded hooks begin on setiger 7. Two or three capillary setae ac- 22 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY Fwtnut 14.?Poly dor a socialis: a, Se tiger 1, anterior view; b, fascicle of modified spines and pennooed setae from aetiger 5; e, hooded hook from anterior setiger; d, hooded hook from median setiger; e, f, hooded hooks from posterior sedgers; g, gisard-like structure from an adult (not to scale). NUMBER 75 23 company the hooks for several setigers, then disap- pear, and appear again in far posterior setigers. In anterior setigers, the hooks have a 45? angle between the main fang and the secondary tooth (Figure 14c). In posterior setigers, the main fang is longer and narrower, while the upper tooth is smaller and closely adhering to the shaft and diminishing the angle (Figure 14e). Setiger 5 is large and well developed (Figure 15a, b). The setae include a small bundle of dorsal geniculate setae lying at the anterior end of a semicircular row of alternating heavy modified spines and smaller pennoned companion setae (Fig- ure 14b). The heavy spines are falcate and have a subterminal protuberance. Ventral and posterior to the above is a small tuft of winged setae. Branchiae begin on setiger 8 (rarely 7-9) and continue to near the posterior end. The pygidium typically has a large ventral lobe and two small dorsal ones, with a dorsal gap (Fig- ure 15c, d). In some specimens, however, the three lobes are continuous. In P. socialis an unusual grinding apparatus has been observed between the esophagus and intestine. In young specimens, this "gizzard-like" structure is thin and transparent, consisting of four teeth, each with a central, inwardly directed cusp. In older specimens, the teeth are fragmented and lie as in- clusions in four longitudinal muscles (Figure 14g). This structure is similar to one figured by Carazzi (1893) for P. flaw. REMARKS.?The type material of P. socialis (Schmarda) from Chile (Vina del Mar) was re- described by Mesnil (1896). Hartman (1941, 1948, 1961), Rioja (1943, 1947), and Hartman and Reish (1950) have extended the species range from Mexico to Oregon and Alaska. Hartman (1953, 1966a) has reported the species from the Falkland Islands. Records from eastern North America in- clude Hartman (1945) and Wells and Gray (1964) from the Carolinas and Hartman (1951) from Flor- ida. In each of these reports, the specimens lacked notosetae on setiger 1. A new subspecies, P. socialis plena, was described by Berkeley and Berkeley (1936, 1952) from Brit- ish Columbia. Reish (1968) reported P. socialis plena from Baja California. This subspecies differed in having notosetae on setiger 1. With the record of Reish, P. socialis plena overlapped the range of P. socialis (sensu stricto), a fact which seemed un- usual to this investigator. Early in the present study it became apparent that all specimens of this species from the east coast of North America, from Maine to South Carolina, possessed notosetae on setiger 1, thus agreeing with the subspecies plena. Since the notosetae are small, it seemed possible that they had been overlooked in earlier east and west coast reports. During a visit to California, it was possible to examine specimens of P. socialis collected by Dr. Keith H. Woodwick from various California localities. All possessed notosetae on setiger 1. Collections in the Allan Hancock Foundation were also examined. The specimens which Hartman (1941) had reported from Mugu Lagoon, southern California, were found to have notosetae on setiger 1, as were speci- mens identified as P. socialis from the Lund University Chile Expedition (unpublished), now de- posited in the Allan Hancock Foundation. Further, specimens of P. socialis from localities in Chile de- scribed by Hartmann-Schrdder (1962, 1965) also possessed notosetae on setiger 1. The presence of notosetae on setiger 1 on all specimens from Chile to British Columbia, and from South Carolina to Maine eliminates the need for the subspecies plena. The "gizzard-like" structure described above was found on all specimens examined and the structure of the modified spines of setiger 5 was found to be the same. Specimens, described as Polydora gracilis Verrill by J. P. Moore (MS), were examined in the Phila- delphia Academy of Natural Sciences. They agreed with P. socialis. Blake (1969a) placed P. gracilis in synonymy with P. socialis but this is at least ques- tionable, since no type material of P. gracilis exists and the description is not adequate for specific de- termination. ECOLOGY.?Polydora socialis constructs tubes in sediment and may form extensive beds (Hartman, 1941). In the Damariscotta River, Maine, P. socialis is found subtidally in sediment and to a lesser ex- tent in shell. In the former, it forms tubes of silt with a firm mucoid lining, and is associated with P. quadrilobata and Laonome sp. In the latter it bores into shells of dead Mercenaria and living Placopecten magellanicus associated with P. con- charum and P. websteri. DISTRIBUTION.?East and west coasts of North America; Gulf of Mexico; Chile; Falkland Islands. 24 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY FKUBE 15.?Potydora anoculata: a. Anterior end, dorsal view; b-i, heavy modified spine* ef tetiger 5 seen in different views and degrees of wear; /, hooded hook. NUMBER 75 25 Polydora anoculata Moore FICUBE 15 Polydora anoculata Moore 1907, pp. 197-199, pi. 15:fig?. 7-17, 25.?Sumner et al., 191S, p. 625. MATERIAL EXAMINED.?Massachusetts (Cape Cod Bay, coll. SEP; Holotype, ANSP 2338). DESCRIPTION.?Polydora anoculata is a thin, slen- der species measuring up to 20 mm in length and having about 100 segments. The prostomium is anteriorly bifurcated. Eyes are absent. The caruncle extends posteriorly to setiger 5 (Figure 15a). Setiger 1 has both noto- and neurosetae. The former are thin, slender capillaries, while the latter have narrow wings. Setigers 2-4, 6 and succeeding setigers contain winged capillary notosetae. The slender capillary neurosetae of setigers 2-4, and 6 are almost completely replaced by hooded hooks on setiger 7. The hooks have a recurved shaft and no constriction (Figure 15/). Setiger 5 contains three groups of setae: a bundle of pennoned dorsal setae anterior to a row of heavy spines alternating with pennoned companion setae, and a ventral tuft of delicate pennoned setae. The heavy spines have a lateral flange, which resembles a tooth when worn (Figure 15b-i). An additional small spur is occasionally seen on some spines. Branchiae begin on setigers 9 or 10 (Figure 15a). They are small at first, attaining full size on setiger 11; they are absent from posterior setigers. The pygidium is disklike, with a deep dorsal notch. REMARKS.?Polydora anoculata resembles -P. giardi Mesnil and P. trideniiculata Woodwick in branchial arrangement. It differs from the latter in the structure of the modified spines of setiger 5 (Woodwick, 1964). The differences between P. giardi and P. anoculata, however, have not been entirely resolved in the present study. Specimens labeled P. giardi were obtained from Europe, but they proved to be another species. It may be that the two species are conspecific. Additional material is needed for comparison. ECOLOGY.?Moore (1907) reported P. anoculata associated with colonies of Amaroecium pelluci- dum, on piles among Cynthia, and in sponge to- gether with P. colonia. In Cape Cod Bay it was found on a shell-gravel bottom. DISTRIBUTION.?Known only from the Woods Hole region, Massachusetts. Polydora hartmanae, new species FIGURE 16 Polydora anoculata?Hartman 1945, p. SS. Not Moore 1907. MATERIAL EXAMINED.?The specimens from Bogue Sound, near Beaufort, North Carolina, de- scribed by Hartman (1945) as P. anoculata, were reexamined. They proved to be an undescribed species. The holotype and one paratype are de- posited in the Allan Hancock Foundation. DESCRIPTION.?The types consist of two anterior ends and two posterior fragments, with the py- gidium missing. One anterior end with 78 segments measured 10 mm in length. The prostomium is distinctly bifid in its anterior margin, extending posteriorly as a low caruncle to setigers 4 or 5. Eyespots and nuchal tentacle are lacking. The peristomium is greatly enlarged on the lateral and anterior margins (Figure 16a). Setiger 1 has fascicles of capillary setae in both the noto- and neuropodia. The neurosetae are finely unilimbate. Setigers 2-4 contain spreading fascicles of winged, capillary noto- and neurosetae. The notopodia of setiger 6 and succeeding setigers con- tain double fascicles of long and short, winged capil- lary setae, the number gradually diminishing posteriorly. No specialized posterior notosetae were observed. Bidentate hooded hooks begin on setiger 7, with 5 or 6 per neuropodium; they have a re- curved shaft. The teeth are delicate and have a re- duced angle between them. (Figure 161). Setiger 5 contains three groups of setae: a cluster of 4 or 5 large, geniculate setae (Figure 16A), lo- cated anterior and dorsal to a semicircular row of heavy spines (Figure 16b-h) alternating with pen- noned companion setae (Figure 16*,;), and a small tuft of winged neurosetae below the large spines. The heavy spines are distally falcate, with a lateral flange, the latter being broken or worn on all ex- cept newly-formed spines. Tufts of fine hairs arise from the concavity separating the flange from the main fang and, to a lesser degree, on the exterior of the flange (Figure 166-g). Branchiae begin on setigers 11 and 12, being short and stubby anteriorly and reaching full size on about segment 16. They are absent from pos- terior segments. The pygidial structure is unknown. REMARKS.?Polydora hartmanae differs from 26 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY TABLE 1.?Some taxonomic characteristics of Species of POLYDORA aggregate anoculata caulleryi colonia commensalis concharum hartmanae ligni quadrilobaU sodalit tetrabranchia websteri branchiae: a. begin on setiger b. on posterior segments a. 7 b. absent on posterior half a. 9-10 b. abtent on posterior half a.7 b. absent on posterior third a.7 b. absent on posterior half a. 6 b. present a. 7-9 b. absent on posterior half a. 11-12 b. absent a.7 b. present a.7 b. absent on posterior third a. 8 (rarely 7-9) b. absent on last segments a.7 b. absent from setiger 11 a.7 b. absent Notosetae on first setiger absent present present absent present present present absent present present absent absent Modified posterior notopodial spines or hooks absent absent awl-shaped spines falcate recurved hooka absent absent absent absent awl-shaped spines absent absent absent the species Modified spines of 5th setiger falcate. with accessory knob falcate. with accessory flange falcate. with bushy top bifid tip. with sub- terminal collar falcate. with long accessory sheath falcate. with lateral sheath falcate, with lateral flange and fine hairs falcate, with small accessory tooth bifid tip, with fine hairs be- tween teeth falcate. with sub- terminal boss bifid tip, with 2 large teeth falcate, with lateral flange or sheath of Polydora from eastern North America Eyes 4 0 0-4 0 4 0-4 0 4 4-6 0-6 4 4 Hooded hooks: a. begin on setiger b. have constric- tion on shaft a.7 b. present a.7 b. absent a.7 b. absent a.7 b. present a. 10-17 b. absent a.7 b. absent a.7 b. absent a.7 b. present a.7 b. absent a.7 b. absent a.7 b. absent a.7 b. present Pygidium disk-like disk-like 4 equal lobes disk-like 4-14 papillae 4 lobes, dorsal pair smaller rii*n ventral pair ? disk-like 4subequal lobes 3 lobes or disk-like collar-like disk-like Other features pigmented pattern in anterior region - - - commensal in gastropod shells occupied by hermit crabs; palps short comparatively long species; S sizes of posterior capillary notosetae _ nuchal tentacle present pigmented pattern in anterior body region gizzard-like struc- ture present; larval pigment re- tained on adults very small species _ NUMBER 75 27 Ficuiut 16.?Potydora hartmanae new species: a, Anterior end, donal view; b-h, heavy modified spines of setiger 5 seen in different views and degrees of wear; i-j, companion setae of setiger 5; k, donal geniculate seta of setiger 5; I, hooded hook. 28 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY other species in the structure of the modified spines of setiger 5. DISTRIBUTION AND ECOLOGY.?The species is known only from North Carolina. It was reported by Hartman (1945, p. 33) as being ". . . dredged from a bottom of broken shell fragments." It is not known whether the species actually bored into the shells or inhabited sediment. Summary Twelve species of Polydora have been identified from collections obtained from various localities along the east coast of North America. A key to the species, descriptions and figures are included. The adults of P. aggregata are described for the first time. Polydora hartmanae is new to science. New synonymies include P. littorea Verrill, referred to P. ligni Webster, P. tubifex Verrill, referred to P. quadrilobata Jacobi, and P. ancistrata Jones, re- ferred to P. colonia Moore. Four species to date have been reported only from eastern North America: Polydora aggregata from Maine, P. hartmanae and P. tetrabranchia from North Carolina and P. anoculata from Massa- chusetts. Polydora concharum ranges from New England to Newfoundland and West Greenland. Each of these species may be expected to have a wider range when additional collections become available. The other seven species of Polydora have a wider distribution. Five species were found boring into shells or other calcareous structures: P. commensalis, P. con* charum, P. socialis, P. tetrabranchia and P. websteri. Five species were found in sediment: P. aggregata, P. caulleryi, P. ligni, P. quadrilobata and P. socialis. The latter species is found in both sediment and shell. Polydora colonia is found in sponges, while P. anoculata occurs in association with tunicates and sponges. Some taxonomic characteristics of the twelve spe- cies are summarized in Table 1. Literature Cited Abbott, D. P. 1946. Some Polychaetous Annelids from a Hawaiian Fish Pond. University of Hawaii Research Publication, 23:1-24, 5 plates. Agassiz, A. 1867. On the Young Stages of a few Annelid*. 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Further Studies on the Benthic Fauna in a Recently Constructed Boat Harbor in Southern California. Bulletin of the Southern California Academy of Sciences, 62:23-32, 3 figures. 1964a. Studies on the Mytilus edulis Community in Ala- mitos Bay, California:?II. Population Variations and Discussion of the Associated Organisms. Veliger, 6:202-207,3 figures. 1964b. A Quantitative Study of the Benthic Polychaetous Annelids of Catalina Harbor, Santa Catalina Island, California. Bulletin of the Southern California Academy of Sciences, 63:86-92,1 figure. 1968. A Biological Survey of Bahia de Lot Angeles, Gulf of California, Mexico n. Benthic Polychaetous An- nelids. Transactions of the San Diego Society of Natural History, 15:67-106,29 figures. Rioja, E. 1943. Estudios anelidologicos Vm. Datos acerca de las especies del genero Polydora Bosc de las costas Mexicanas del Padfico. Anales del Instituto de Biologic, 14:229-241,25 figures. Mexico. 1947. Estudios anelidologicos XVII. 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A Biological Survey of the Waters of Woods Hole and Vicinity. Section 3. A Catalogue of the Marine Fauna of Woods Hole and Vicinity. Bulletin of the United States Bureau of Fisheries, 31:549-794. 32 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY Turner, H. J., and J. E. Hanks 1959. Infestation of Pecten irradians by Polydora. Nau- tilus, 72:109-111. Uschakov, P. V. 1955. [Polychaeta of the Far Eastern Seas of the USSR.] Oprcdeliteli Po Faune, SSSR, 56:1-445, 164 figures. (In Russian.) 1965. Polychaeta of the Far Eastern Seas of the USSR. English edition of the 1955 publication. Trans- lated by the Israel Program for Scientific Transla- tions, United States Department of Commerce, Washington, D. C, 419 pages, 164 figures. Verrill, A. E. 1873. Report Upon the Invertebrate Animals of Vineyard Sound and the Adjacent Waters, with an Account of the Physical Characters of the Region. Report of the United States Commissioner of Fisheries, 1871-1872, pages 295-778. 1880. Notice on Recent Additions to the Marine Inverte- brata, of the Northeastern Coast of America, with Descriptions of New Genera and Species and Critical Remarks on Others. Part n. Annelida, Gephyrea, Nemertina, Nematoda, Polyzoa, Tunicata, Echino- dermata, Porifera. Proceedings of the United States National Museum, 2:165-205. 1881. New England Annelida Part I. Historical Sketch, with Annotated List of Species Hitherto Unrecorded. Transactions of the Connecticut Academy of Arts and Sciences, 4:285-324, 12 plates. 1885a. Results of the Explorations made by the Steamer Albatross off the Northern Coast of the United States in 188S. Report of the Commissioner of Fish and Fisheries for 1883, pages 503-699, plates 1-44. 1885b. Notice of Recent Additions to the Marine Inverte- brata of the Northeastern Coast of America, with Descriptions New Genera and Species and Critical Remarks on Others. Proceedings of the United States National Museum, 8:424-448. Webster, H. E. 1879a. Annelida Chaetopoda of the Virginian Coast. Transactions of the Albany Institute, 9:202-269, 11 plates. 1879b. Annelida Chaetopoda of New Jersey. Annual Re- port of the New York State Museum, pages 101-128 (the 7 plates were not published until reprinted in 1886). Webster, H. E., and J. E. Benedict 1884. The Annelida Chaetopoda from Province town and Wellfleet, Massachusetts. Report of the United States Commissioner of Fisheries, 1881, pages 699- 747, 7 plates. 1887. The Annelida Chaetopoda from Eastport, Maine. Report of the United States Commissioner of Fish- eries, 1885, pages 707-755, 8 plates. Wells, H. W., and I. E. Gray 1964. Polychaetous Annelids of the Cape Hatteras Area. Journal of the Elisha Mitchell Scientific Society, 80:72-78. Woodwick, K. H. 1953. Polydora nuchalis, a New Spedes of Polychaetous Annelid from California. Journal of the Washing- ton Academy of Sciences, 43:281-283, 1 figure. 1963. Taxonomic Revision of Two Polydorid Spedes. (Annelida, Polychaeta, Spionidae). Proceedings of the Biological Society of Washington, 76:209-216, 2 figures. 1964. Polydora and Related Genera (Annelida, Poly- chaeta) from Eniwetok, Majuro and Bikini Atolls, Marshall Islands. Pacific Science, 18:146-159. 4 fig- ures. Zaks, I. 1933. [Polychaeta of the North-Japanese Sea.] Issledo- vaniya Morei SSSR, 19:125-137. (In Russian with German Summary.) U.S. GOVERNMENT PHINTINO OPPICBt 1*7f Publication in Smithsonian Contributions to Zoology Manuscripts for serial publications are accepted by the Smithsonian Institution Press subject to substantive review, only through departments of the various Smithsonian museums. Non-Smithsonian authors should address inquiries to the appropriate department. If submission is invited, the following format requirements of the Press should govern the preparation of copy. 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