THE ATLANTIC COAST SURF · with a partial bibliography

UNITED STATES DEPARTMENT OF THE INTERIOR AND WILDLIFE SERVICE BUREAU OF COMMERCIAL

Circular 288 DEPARTMENT OF THE INTERIOR

U.S. FISH AND WILDLIFE SERVICE BUREAU OF COMMERCIAL FISHERIES

THE ATLANTIC COAST SURF CLAM - with a partial bibliography

By

ROBERT M . YA 'CEY and WALTER R. WELCH

Circular 288

Washington, D . C. June 1968 CONTENTS Page

Introduction ......

Hi -tory of the 1 Ea rly period - - 1 70 's to 1 Q42 2 Developm'ntal period -- 1943-4q 2 Recent p nod - - 1950- 5 ..... 2

HandHng. process1f~g. and marketing 3

Biology ....•... 4 Cia" s lilc tion a net range 5 Morphol0 ical varia lon. 5 Habltat .•... 5 . 6 Growt}, .•... 7 R lative abundanc.e . 7 Mortali y and pathology. 7

Pa ~ lal biblIography ...... 9

iii f OL }( - U ~ . THE ATLANTIC COAST SURF CLAM with a partial bibliography

By ROBER T M . YANCEY, Fishery Biologist Bureau of Commercial Fisheries Biological Laboratory Oxford, Md. 21654 and WALTER R. WELCH, Fishery Biologist Bureau of Commercial Fisheries Biological Laboratory West Boothbay Harbor, 04575

AB STRACT

The surf clam, solidissima solidissima (Dillwyn), supports an important commercial fishery along the coasts of the Middle Atlantic States. The major portion of the catch has been taken off New Jersey since 1949. Surf clam vessels, equipped with hydraulic jet dredges, take the from depths of 100 feet or less and land them the same day. Most of the catch is processed for use as ; a minor portion is used as bait for fish. The total catch in 1965 was 44.1 million pounds of . Surf clams are found in waters of oceanic salinity and or gravel bottoms from the low-tide line to depths of 480 feet. They range from the Gulf of Saint Lawrence to Cape Hatteras, N.C. The sexes are separate. Spawning usually occurs twice a year. , under experimental conditions, failed to develop in salinities below 23 parts per thousand. Young clams may reach a length of 1 3/ 4 inches in 1 year. Mature clams are commonly 6 to 7 inches long. Surf clams are very abundant in some areas. Populations of 1 to 2 million juvenile clams per mile of beach have been found at Wallops Island, Va., and Sandy Hook, N.J. As many as 5 million mature clams per mile of beach have been washed up on Long Island, N. Y., by a storm.

INTRODUCTION has been taken by highly efficient gear. Ques­ tions of sustained yield, growth, and recruit­ The surf clam, Spisula solidis sirna solidis­ ment to the fishery must be answered to s irna (Dillwyn), is an important provide information for the management of res ource in the United States. In recent years the fishery. (1 949 - 65) the commercial fishery for surf This report summarizes existing informa­ clams has grown rapidly. The adoption of tion about the surf clam fishery and its history advanced technology in fishing and processing as well as information about the distribution, has enabled the fishery to harvest increasingly abundance, and biology ofthe . A partial greater amounts each year--reaching 44.1 bibliography is included. million pounds of clam in 1965. A basic life history study of the surf clams was made in New England by Belding (1910). HISTORY OF THE FISHERY Many subsequent reports have been concerned with faunal surveys and aspects of physiology The history of the fishery for surf clams is and where the surf clam was of divided into three periods: the early period, s econdary importance. when the fishery was small; the develop­ Truly intensive studies of biology and popu­ mental period, when it was growing rapidly; lation dynamics were first established in and the recent period, when it is well­ New Jersey where most of the catch since 1949 developed. Early Period -- 1870's to 1942 used mainly for bait, and the landings there dropped sharply owing to wartime restrict ions Large quantities of surf clams, thrown up on party boat (sport) fishing. In the Long on the beaches by storms, were gathered and Island, N.Y., area, the increased demand for eaten by the Indians in precolonial days. Early more clams of bette r quality made the indus­ settlers on our Atlantic coast from Cape Cod, try realize that fishing gear and methods Mass., to New Jersey occasionally used the needed improving. Late in 1945, when the clams for food in times of hardship, but more forerunner of the modern hydraulic jet dredge often used them for fertilizer or food for was developed by the industry, catch per unit swine and poultry. of effort and quality of clams increased The first organized fishery began on Cape markedly. Landings increased fourfold in 1945 Cod during the 1870's. Clams were raked or because (1) the fleet increased to 50 boats by tonged up by hand, shucked, and salted down in the end of the year, (2) the new hydraulic jet barrels to be used as bait in the handline dredge doubled the efficiency of gear, and fishery for cod. In a good year (1877), 3,000 (3) boats could successfully dredge areas barrels, worth about $18,000, were produced formerly considered to have too few clams. (Ingersoll, 1887). Although early catch statis­ landings had reached nearly 5.0 tics are scanty, it appears that the fishery million pounds of clam meats by 1949, but continued at a level below that of 1877 until those of Massachusetts, Rhode Island, and 1929, w,hen power with scrape-type New Jersey were far less. dredges began at Long Island, N.Y. Dredging began at Sheepshead Bay and spread to areas of Rockaway Inlet and Jones Inlet and southern Recent Period -- 1950-65 New Jersey. The development and use of Industry found that the yield of meats per scrape-type dredges facilitated the taking of bushel of clams was 12 pounds in New York larger quantities of clams and the use of beds and 17 pounds in New Jersey. Because this in de epe r wa te r s, although tong s continue d to greater yield and the increased demand for be used. surf clam meats for food, the industry began The fisheries in New York and New Jersey exploring for clams in 1949 off the coasts of developed equally from 1929 to 1942, and ir­ New Jersey, Delaware, and in 90 to regular landings were made in Massachusetts 100 feet of water. Beds of clams were found. and Rhode Island. The peak year for this early These beds were much more extensive than the period was 1938, when 1.6 million pounds of Long Island beds, and the clams yielded more meats were landed. meat per bushe1. In this middle Atlantic area l a fishery subsequently developed for surf Developmental Period -- 1943-49 clams to be used as food. New Jerseylandings Before 1943 many attempts were made by inc reas ed from 0.4 million pounds of clam private industry to promote surf clams for meats in 1949 to 6.9 million pounds in 1953, food. All attempts failed, mainly because of and Maryland landings reached 2.5 million inability to rid the clam meats of sand. In 1943, pounds by 1953 (June and Reintjes, 1957; wartime demands for brought Reintjes and Roithmayr, 1960). The fishery in about a renewed interest in the surf clam re­ .0,------, source. The F. H. Snow Canning Company of Pine Point, Maine, using clams landed at Free­ port, Long Island, N. Y., solved the problem of sand removal with a drum washer and soon 40 increased use of surf clams from small ex­ perimental quantities to 1,500 bushels per day. By the end of 1943 other processors had be­ come interested and were preparing for 1944 30 production (fig. 1). The resulting increase in landings for food occurred mainly in New York. Bylate summer of 1945, the area between Ejast Rockaway Inlet and Fire Island Inlet was being dredged for surf clams and continued to be the most im­ portant area through 1949. Freeport and West Sayville, Long Island, N. Y., were ports where 10 clams were landed. New Jersey clams were

l Westman, James R., and Milton H. Bidwell, 1946.

The surf clam. Economics and biology of a New York 1943 45 47 49 51 marine resource. Unpublished manuscript. In files of Bureau of Commercial Fisheries Biological Laboratory, Figure 1.--Surf clam landings 1943- 65. The important Oxford, Md. 21654. gr owth of the fisher y came after 1943.

2 Maryland was short-lived, however, as the About 100 boats were dredging in 1957, and New Jersey grounds proved to be more pro­ most of them were privately owned and oper­ ductive. By 1957, most of the Maryland clam ated; now the fleet has about 60 boats, of which dredging fleet had moved to New Jersey. nearly all are company-owned and operated. Since 1958, nearly all the Atlantic coast clam The present boats are generally larger than landings used for food have been taken off those in use 10 years ago. New Jersey. The total landings rose from 16.0 million pounds of clam meats in 1956 to 44.1 HANDLING, PROCESSING, million pounds in 1965. New Jersey contributed 96 percent of the total landings in 1965. The AND MARKETING major part of the New Jersey landings has The clams are handled and processed by corne from the area between Point Pleasant various methods. As the clams are dumped on and Beach Haven; some clams are taken off deck from the dredge (fig. 2), they are sepa­ Cape May and landed there. The New York rated from sand and dead shell byhand picking fishery has been unable to meet the competi­ or are shoveled directly into a I-bushel tion of the New Jersey fleet which dredges measure if the haul is clean. When full, the densely populated beds; New York landings measure is dumped into a burlap bag, and the dropped from 2.4 million pounds of clam meats filled bags are stacked on deck. One company in 1956 to 1.5 million pounds in 1965. fills 30-bushel wire cages (fig. 3) rather than The hydraulic jet dredge and accessory I-bushel bags and stores them on deck. Clams equipment were continuously evaluated and im­ are landed the same day they are taken. A proved during this period of increase. The dock hoist unloads six bags or one cage at a gear now is so efficient that the boats of the time. Most clams are then trucked to nearby fleet generally work a 5-day week and are shucking plants, but some are hauled as far often on daily quotas that are met within a 12- as Delaware or Maine. Clams are usually un­ to 14-hour workday (including sailing time). loaded in the afternoon, wetted down, and stored Daily quotas frequently have been imposed overnight in refrigerated rooms, or trucked because the capacity of the processing plants overnight to Maine in refrigerated trucks. is limited. Next morning the clams, still in bags or cages.

Figure 2.--A dredge load of clams about to be dumped on deck.

3 Figure 3.--Thirty-bushel cages of surf clams awaiting delivery to the shucking plant. are dipped in hot (150 0 F.) for 30 Surf clam meats compete with other clam or 40 seconds to open the shells slightly so meats because they are cheaper and make an the meats will be easier to shuck, and then acceptable product when processed. None are cooled in fresh water to prepare them for the sold as food on the fresh market. The landed shuckers. Meats are shucked by hand (fig. 4) value of surf clams used for food is low and and placed in a washer to remove ~and and stable at about 7 cents per pound of meat. The pieces of shell. wholesale value of the processed meats is Washed clam meats are packed in 130-gallon about five times the landed value. Industry stainless steel tanks, lightly iced, and taken to must operate at peak efficiency and depends the processing plant. The gonad and the vis­ on the continued availability of clams in ceral mass are squeezed out by hand (fig. 5), heavily populated, easily accessible beds. leaving the strap, siphons, foot, and The surf clam fishery for bait has a limited adductor muscles. The clam meats are washed market among sport and commercial fisher­ again, and pieces of shell and the black edge men. The landed value per pound of bait clams of the mantle are stripped off. At this point is higher than that of clams for food because the meat may be minced (l/4-inch pieces), of the demand for a special product. The land­ chopped (1 / 2-inch pieces), or sliced into strips, ings in New York (practically all bait clams) depending upon what product is desired. The in 1961-64 had an average value of $0.09 per juices and wash water from this operation are pound of meats. sometimes s a v ed, concentrated, clarified, salted, and packed as clam juice. Meats may BIOLOGY be canned, made into or clam cakes, refrigerated, or frozen and shipped to other Knowledge of the biology of the surf clam processors of specialty products. has previously been based mostly on the work

4 of Belding (1910) and Verrill (IS73) in Ma~ a­ is db' 0 1 on chusetts. The questions of growth, di tribu ion, found b w Cap Cod and reproductive patterns in the ar a of gr at­ Canyon (Chamberlin and S est abundance in the Middle Atlantic Bight mon nam ha hav b e need to be answered by further research. Per­ clam ar sonnel at the B . C . F. (Bureau of Commercial m r, dipp r,b achclam, Fisheries) Biological Laboratory, Oxford, Md., comm rcially lmpor an S. are now studying surf clam biology and popu­ the only p Cle r por d 0 lation dynamics, with the ultimate goal of providing information to industry. The B.C.F. Morphological Vanallon Exploratory Fishing and Gear Research Base, Gloucester, Mass., is surveying the distribu­ tion of the species.

Classification and Range

The surf clam bel 0 n g s to the family , which includes over 200 species found in most of the world IS shallow . It chusetts clams. alve is separated into two subspecies: Spisula waters, less than 40 fe~t d p. of solidissima solidissima (Dillwyn, lS17), the prominent check marks hleh ar large commercial form which is found along the result of environm ntal dl our Atlantic coast from the Gulf of Saint Clams from one sampl Lawrence to Cape Hatteras, N.C., and Spisula shape and length. solidissima raveneli (Conrad, IS31), a diminu­ tive form found from Cape Hatteras, N.C., south to northern Mexico (Merrill and Webster, Habitat 1964). North of Cape Cod, Spisula polynyma The surf clam inhabi (Stimpson, 1860), may be confused with S. from the lower edge of solidissima, but no problem should arise alo;:;g a depth of about 4 0 fee . the Middle Atlantic Bight where S. polynyma Ba y clams a re not comme

Fi r 4.--Yoor· rs sh surf clams,

5 Figure 5.--Women evis<..~rating shucked clams. waters; but in more northern waters of the commonly associated with surf Gulf of Saint Lawrence, the Bay of Fundy, clams in dredge samples taken between Long Maine, and Massachusetts Bay, they live im­ Island, N.Y., and Cape Hatteras, N.C., at mediately below the low-tide line. Clams have depths to 180 feet are: (Cancer, been found at depths of 480 feet along the . Ocellatus, and Libinia); moonsnails Middle Atlantic Bight, and at a depth of 250 (Polinices heros and Lunatia duplicata); razor feet off Digby Gut, Nova Scotia. clams ( directus); mahogany quahogs Surf clams require waters of near oceanic (); southern quahog salinity. They are usually found on open beaches (Mercenaria campechiensis); and va rio us and open bottom, and are generally of the classes Holuthuroidea, absent from enclosed bays and estuaries. We Asteroidea, Echinoidea, and Ophiuroidea. have found surf clams at Chincoteague Inlet, Va. (1.5 miles from the open ocean at Reproduction Chincoteague Point), where records show a minimum salinity of 29 p.p.t. (parts per Reproduction of surf clams is poorly known thousand) (McGary and Sieling, 1953). Pre­ (studies are in progress). The sexe s of surf liminary experiments with adults held at 500 F. clams are separate- -only one hermaphrodite indicate that salinities as low as 15 p.p.t. are has been found in 2,000 clams (John W. Rope s , lethal. personal communication 2 ). Some Massachu­ Surf clams live in sand or gravel bottoms. setts clams have been reported to be sexually Shells, shell fragments, and fine sediments mature when 1.9 inches long and to spawn may be mixed with the basic bottom types. Clams are most common in a bottom of coarse 2 John W. Ropes, B.C.F. Biologica l Laboratory, sand and gravel. Oxford, Md. 21654.

6 at 2.0 inches (Belding, 1910). The gametes showed that clams in a natural environment are extruded into the water, where fertiliza­ can reach a length of 1 inch by the end of tion takes place. Clams spawn during June their first 6 months and 1 3/4 inches by the and July in Massachusetts; essentially the end of their first year. same period was reported for Long Island, N . Y. (see footnote 1). Our preliminary studies of the reproductive cycle show that New Jersey Relative Abundance clams usually spawn from mid-July through Surf clams are extremely abundant in some mid-August; the n they rapidly redevelop areas. From Maine to Maryland, tremendous gametes and spawn again from mid-October quantities of clams have been reported washed through early December (John W. Ropes, per­ ashore b y wave action of winter storms sonal communication (see footnote 2)). The (fig. 7). Jacot (1920) gave an estimate of 5 c lams again begin gametogene sis in February, million clams washed ashore per mile of followed by rapid development of mature beach at Rockaway Beach, Long Island, N.Y., gametes in March and April. Westman and in the winter of 1919-20. Sampling for young Bidwell (see footnote 1) reported that Long clams at Wallops Island, Chincoteague, Va . , Island, N.Y., populations of clams begin to in 1965 by the B.C.F. Biological Laboratory, spawn at 590 F. Fecundity of the clams is Oxford, Md., indicated a density of 1 to 2 unknown. million live clams per mile between low- and Experiments on the rearing of larvae were high-tide marks; densities were similar in that performed at the B.C.F. Biological Labora­ year at Sandy Hook, N .J. (fig. 7). tory, Boothbay Harbor, Maine. Eggs failed to The relative abundance of surf clams from develop normally at salinities below 23 p.p.t. Montauk Point, Long Island, N . Y., to Cape at any temperature used (Alden P. Stickney, Hatteras, N.C., in depths from 40 to 480 feet personal communications 3 ). At 32 p.p.t. the has been determined from recent surveys with optimum tempe ra ture range for development a jet dredge and a grab. The area of greatest was 570 to 68 0 F.--mortality increased at abundance extends from Barnegat, N.J., to higher and lower temperatures. Some larvae, Cape May, N .J. Abundance is moderate in a reared on the unicellular alga Dicrateria narrow strip along the south shore of Long inornata, survived and grew at salinities as Island at depths to 100 feet and an offshore low as 16 p.p.t., and could survive brief ex­ strip 70 to 90 feet deep off Chincoteague, Va. posure (24 hours) to salinities as low as 8 The present middle Atlantic fishery is de­ p.p.t., if temperatures were sufficiently low pendent upon the continued availability of ex­ (46 0 F.). At 32 p.p.t. growth was optimum at tensive and densely populated beds off New 680 F. but was somewhat inhibited at higher York, New Jersey, Delaware, and Maryland. and lower temperatures. The larvae meta­ morphosed 18 to 30 days after fertilization 0 when held at 68 F. Mortality and Pathology Growth Small surf clams are subject to many types of . Bottom-feeding fish, diving Studies of growth of surf clams are essen­ ducks, and gulls feed on the clams. Moon tial for practical resource management. The of the genera Lunatia and Polinices time required for clams to reach commercial bore holes through the shells and eat the length (5 inches) must be known to estimate meats. About one -half of the empty paired rate of replacement of clams taken by the shells taken from a beach windrow at Wallops fishery. Figure 6 compares the informa­ Island, Va., had been bored. As previously tion available from two published sources mentioned, storms often wash enormous num­ (Kerswill, 1944; Belding, 1910), one unpub­ bers of surf clams ashore, where most die. lished report (Westman and Bidwell, see The impact of predation and storms on surf footnote 1), and data obtained at the B.C.F. clam populations is unknown. Biological Laboratory, Boothbay H arb 0 r, No diseases of the surf clam are known, Maine. Clams propagated during the summer and records list only one parasite, Pro­ at Boothbay Harbor averaged 0.04 inch (maxi­ boscidosaccus enigmaticus (of unc e rtain mum 0.4 inch) long by late fall. Studies at the taxonomic position), in the related Spisula B.C.F. Biological Laboratory at Oxford, Md., solida (L)--Gallien (1949). G onads of some New Jersey surf clams taken in 1962-65 have been infested with an unidentified trematode. S Alden P. Stickney, B.C.F. B i 0 log i ca l Laboratory, The effect of this parasite on the clam is West Boothbay Harbor, Maine 04575. not known.

7 u z. T 1: •

C) Z ..J ..J ..J WJ :r VI

Figure 7.--Surf clams washed upon rhe beach by a wlnrer srorm. LI htcr objects arc urf cl ms.

8 PARTIAL BIBLIOGRAPHY OF THE SURF CLAM (As terisks indicate citations in text)

ADAMS , HENR Y, and AR THUR ADAMS. CARPENTER, HORACE F. 1 85 8. The genera of recent . 1887. The shell-bearing Mollusca of Rhode 2 : 374- 388. J ohn Van Voorst, London. Island. Conchologists' Exchange 2: A L LEN, ROBERT DAY. 78-79. 1951a. Antimitotic substances secreted CARSON, RACHEL L. f rom eggs . [Abst ract.] BioI. Bull. 1945. Fish and shellfish of the Middle (Woods Hole) 101: 214. Atlantic coast. [U.S.] Fish Wildl. Serv., 1951b. The role of the nucleolus in spindle Cons. Bull. 38, iv + 32 pp. formation. [Abstract.] BioI. Bull. CHADWICK, GEORGE H. (Woods Hole) 101: 214. 1906. Shells of Prince Edward Island. 1951c. The use of Spisula solidissima eggs 19: 103-104. in cell res earch. J. Cell. Compar. CHAMBERLIN, JOSEPH LOCKWOOD. Physiol. 37: 504-505 . 1954. Studies in the Mactrinae. Unpub­ 1952a. A visible response to stimulation lished Ph.D. Thesis, Harvard Univ., in Spisula eggs. [Abstract.] BioI. Bull. 132 pp. (Woods Hole ) 103: 283. CHAMBERLIN, JOSEPH LOCKWOOD, 1952b . The behavior of Spisula eggs with JAMES B. ENGLE, CARL. N. SHUSTER, respect to potassium ions. [Abstract.] JR., and WALTER R. WELCH. BioI. Bull. (Woods Hole) 103: 290-29l. 1962. Report on the surf clam. Minutes 1953a. Fertilization and artificial activa­ 20th Annu. Meeting, Atl. States Mar. tion in the of the surf-clam, Fish. Comm. (1961). Appendix MA 2, Spisula s olidis sirna. Biol. Bull. (Woods 3 pp. [Mimeo.] Hole) 105: 213-239. *CHAMBERLIN, JOSEPH LOCKWOOD, and 1953b. Fertilization and artificial activa­ FRANKLIN STEARNS. tion in the egg of the surf- clam, 1963. A geographic study of the clam, Spisula solidissima. Unpublished Ph.D. Spisula polynyma (Stimpson). Amer. Thesis, Univ. Pennsylvania, 43 pp. Geograph. Soc. N.Y., Ser. Atlas Mar. ANONYMOUS. Environ. 3, 12 pp. 1955. Bar clam resources. Trade News CONRAD, T. A. 8(3): 8-10. 1868. Catalogue of the family Mactridae. 1959. Canada's shellfish resources. Trade Ame r. J. Conchol. 3: 30-47. News 11(9): 5-7, 9 . DALL, WIL LIAM HEALEY. ASHTON, FRANCIS T. 1894. Synopsis of the Mactridae of North 1960. Germinal vesicle breakdown in the America. Nautilus 8:25- 28. eggs of Spisula and Hydroides. Un­ FISCHER, PAUL. published Ph.D. Thesis, Univ. Pennsyl­ 1887. Manuel de conchyliologie. Pp.1114- vania, 38 pp. 1120. Librarie F. Savy, Paris. BARNES, E. W. FORD, E. 1910. The plague of clams at Easton's 1925. On the growth of some lamelli­ Beach, Newport. Fortieth Annu. Rep. branchs in relation to the food- supply Comm. Inland Fish. Providence, R.T. of . J. Mar. Biol. Ass. U.K. (1909): 183-185. 13: 531-559. *BELDING, DAVID L. FORD, JOHN. 1910. The growth and habits of the sea 1889. List of shells of the New Jersey clam ( solidissima). Rep. coast sou t h of Brigantine Island. Comm. Fish. Game Mass. 1909. Publ. Nautilus 3: 27-28. Doc. 25: 26-4l. FOX, WILLIAM J. BOND, R. M., and J. C. MEDCOF. 1891. List of Mollusca of Glo.ucester Co., 1958. Epidemic in New N.J. Nautilus 4: 113-115. Brunswick, 1957. Can. Med. Ass. J. FROMMHAGEN, LAURENCE H., MARVIN J. 79: 19 - 24. FAHRENBACH, JOHN A. BROCKMAN, BOUSFIELD, E. L., and A . H. LEIM. JR. , and E. L. R. STOKSTAD. 1958. The fauna of Minas Basin and Minas 1953. Heparin-like anticoagulants fro m Channel. Nat. Mus. Can., Bull. 166, Mollusca. Proc. Soc. Exp. BioI. Med. 30 pp. 82: 280-283. BURCH, HELEN B., and PHILIPP STRITT­ *GALLIEN, L. MATTER. 1949. Pro b 0 sci do sac c u s enigmaticus 1963. Enzyme patterns in eggs, developing nov.g., nov.sp., parasite de Mactra embryos and adult tissues of L. (Note preliminaire~) solidissima. [Abstract .] BioI. Bull. Bull. Soc. Zool. France 74: 280- (Woods Hole) 125: 373. 283.

9 GLAl 'CY, JOSEPH B. Ir\GERSOLL, ERNEST, and JOHN A. 1945. Utilization of the surf clam, Mactra RYDER--Con. solidissima. Trans. 10th r\. Amer . and fishery industries of the United Wildl . Coni.: 294-297. States, sect. 1: 703-710. U.S. Govern­ G01'SE, PIERRE H. ment Printing Office, Washington, D.C. 1959. Respiration and motility in Spisula ITO, SUSUMU, and CECILIE LEUCHTEN­ spermatozoa. [Abstract.] BioI. Bull. BERGER. (Woods Hole) 117:433-434. 1955. The possible role of the DNA content GOULD, AUGUSTUS A. of spermatozoa for the activation 1870. Report on the Invertebrata of Massa­ process of the egg of the clam Spisula chusetts. 2d edT Wright and Potter, solidis sirna. Chromosoma 7: 328- 339. Boston, 524 pp. + 22 plates. JACOBSON, MORRIS K. GRA 'D, C. G. 1944. Variations of Spisula solidissima 1938. Intracellular pH studies on the ova Dillwyn. Nautilus 57: 100-104. of Mactra solidissima. [Abstract.] JACOT, ARTHUR. BioI. Bull. (Woods Hole) 75: 369. 1919. Some marine Mollusca about New HAGERMAr\, DWAIN D ... FREDERICA M. York City. Nautilus 32: 90-94. WELLINGTO T and CLAUDE A. *1920. Notes on some Mollusca about New VILLEE. York City. Nautilus 34: 59- 60. 1957. Estrogens in marine . JOHNSON, CHARLES w. BioI. Bull. (Woods Hole) 112: 180-183. 1890. Annotated list of the shells of St. HARDING, CLIFFORD V. Augustine, Fla. Nautilus 4: 4- 6. 1958. The 0 s mot i c behavior of marine 1934. List of marine Mollusca of the oocyte nuclei. [Abstract] BioI. Bull. Atlantic coast from Labrador to Texas. (Woods Hole) 115: 371-372. Proc. Boston Soc. Nat. Hist. 40: 1-204. HAYASHI, TERU, RAJA ROSENBLUTH, and *JUNE, FRED C., and JOHN W . REINTJES . HAYES C. LAMONT. 1957. Survey of the off 1959. Studies of fibers of acto- and para­ Delaware Bay. U.S. Fish Wild!. Serv., myosin fro m lamellibranch muscle. Spec. Sci. Rep. Fish. 222, iv + 55 pp. [Abstract.] BioI. Bull. (Woods Hole) KEITH, M. L., G. M. ANDERSON, and 117: 396. R. EICHLER. HAYES, F. R., and D. PELLUET. 1964. Carbon and oxygen isotopic composi­ 1940. Inorganic constituents in molluscan tion of mollusk shells from marine and blood and muscle. Rep. Woods Hole fresh-water environments. Geochim. Inst. Oceanogr. 1939: 1920. (ColI. Re­ Cosmochim. Acta 28: 1757-1786. prints WHOI 1940.) KELLY, JOHN W. .HEILBRUNN, L. V., and W. L. WILSON. 1954. Metachromasy in the eggs of fifteen 1958. A physical study of the ground sub­ lower animals. Protoplasma 43: 329- stance of the Spisula egg. [Abstract.] 346. BioI. Bull. (Woods Hole) 115: 328. KENNEDY, DONALD. HENDERSON, JEAN T. 1958. Neural photosensitivity in Mactra. 1929. Lethal temperatures of Lamelli­ [Abstract.] BioI. Bull. (Woods Hole) branchiata. Contrib. Can. BioI. Fish., 115: 338. n.s. 4: 399-411. 1960. Neural photoreception in a lamelli­ HOPKI S, HOYT S. branch mollusc. J. Gen. Physiol. 44: 1934. Catalase and oxidative processes in 277-299. tissues as possible factors in *KERSWILL, C . J. adaptation. BioI. Bull. (Woods Hole) 1944. The growth rate of bar clams. Fish. 67: 115-125. Res. Bd. Can., Prog. Rep. Atl. Coast Ht: 'TER, 'v'''. RUSSELL, and DAVID C . StaT 35: 18-20. GRAr\T. KIVY -ROSENBERG, EVELYN, and FRANCES 1962. Mechanics of the ligament in the RAY. bivalve Spisula solidissima in relation 1961. Krebs and pentose cycle dehydro­ to mode of life. BioI. Bull. (Woods genase systems in the eggs of Spisula Ho:e) 122: 369-379. as measured with a tetrazolium . *1 GERSOLL, ERNEST. [Abstract.] BioI. Bull. (Woods Hole) 188,. Distribution, methods of gathering, 121: 394. and uses of sea-clams. In George KIVY -ROSENBERG, EVE L Y N, J OS E P H Bro\\n Goode, The fisheries and fishery CASCARANO, and G. MERSON. industries of the United States, sect. 5, 1959. Examination of some D.P. N .- and 2: 08 - 61 I. U.S. Gove rnrnent Printing T.P.N. - dependent dehydrogenase activ­ Office, Washington, D .C. ity before and after relatively high GEkSOLL, ER EST, and JOH1' A. RYDER. doses of x-irradiation of Spisula eggs. 1 • Th!' blvalves--Lamellibranchiata. [Abstract.] BioI. Bull. (Woods Hole) !!: George Brown Goode, The fisheries 117: 415-416.

10 KIVY -ROSENBERG, EVEL YN, KAREN STEEL MEDCOF, J. C., and J. S. MacPHAIL. KAGEY, and JOSEPH CASCARANO. 1955. Survey of bar clam resources of the 1958. Dehydrogenase activity in develop­ Maritime Provinces. Fish. Res. Bd. mental stages of Spisula as measured Can., Bull. 102, vi .,. 6 pp. with a tetrazolium salt. [Abstract.] *MERRILL, ARTHUR S., and JOH R . Biol. Bull. (Woods Hole) 115: 354-355. WEBSTER. KRANE, STEPHEN M., and ROBERT K. 1964. Progress in surf clam biological CRANE. research. In C. J . Sindermann (edi­ 1960. Effect of nicotinamide on pyridine tor), The Bureau of Commercial Fish­ nucleotide levels and cleavage rate eries Biological Laboratory, Oxford, of eggs of Spisula sol i dis s i m a. Maryland: Programs and perspectives, [Abstract.] Biol. Bull. (Woods Hole) pp. 38-47. U .S. F1Sh Wildl. Serv., 119: 324. Circ. 200. KRANE, STEPHEN M., and LEONARD METZ, CHARLES B., and JOANNE E. LASTER. DONOVAN. 1962. The incorporation of nicotinamide- 1949. Fertilizin from the eggs of the clam, 7-C l4 into pyridine nucleotides of in­ Mactra solidissima. [Abstract.1 Biol. tact eggs and embryos of Spisula Bull. (Woods Hole) 97: 257. solidissima. [Abstract.] Biol. Bull. MITCHILL, SAMUEL. (Woods Hole) 123: 501-502. 1826. Facts and observations intended to LASTER, LEONARD, and ROBER T K. CRANE. illustrate the natural and economical 1962. Triphosphopyridine nucleotide for­ history of the eatable clam of ew mation and disappearance in the pres­ York and its vicinity. Amer. J . Sci. ence of extracts of eggs, embryos and Arts. 10: 287-293. adult of Spisula solidissima. MORSE, EDWARD S. [Abstract.] BioI. Bull. (Woods Hole) 1919. 0 b s e r vat ion s on living lamelli­ 123: 502. branchs of New England. Proc. Boston LEIDY, J. Soc. Nat. Hist. 35: 193-196. 1878. Remarks on Mactra. Proc. Acad. NAGABHUSHANAM, R. Nat. Sci. Philadelphia 30: 332-333. 1962. The histology of the neurosecretory LOOSANOFF, VICTOR L. system of the clam, Spisula solidissima. 1943. Soft and hard clams of the Atlantic [Abstract] Amer. Zool. 2: 543. coast of the United States. U .S. Fish 1963. Observations on the neurosecretory Wildl. Serv., Fish. Leafl. 13, 11 pp. cells in the lamellibranch, ~sula 1946. Commercial clams of the Atlantic solidissima (Dillwyn). Nucleus 6: 99- coast of the United States. U.S . Fish 102. Wildl. Serv., Fish. LeaH. 13(Rev.), NAT ION A L combined wit h 14 pp. MAINE COAST FISHERMAN (news­ 1954. New advances in the study of bivalve paper). larvae. Amer. Sci. 42: 607-624. 1961. Big stern dredge harvesting sea clams LOOSANOFF, VICTOR L., and H. C. DAVIS. for Borden firm. National Fisherman 1963. Rearing of bivalve mollusks. Adv. 42(7): 14. Mar. Biol. 1: 1-136. NOLAND, JERRE L. LOVE, ROBERT, and LAURENCE H. 1949. Determination of amino acids in in­ FROMMHAGEN. . [Abstract.1 Biol. Bull. 1953. Histochemical studies on the clam (Woods Hole) 97: 263-264. Mactra solidissima. Proc. Soc. Exp. PACKARD,ALPHEUSS. Biol. Med. 83: 838-844. 1865. Observations on the glacial phenom­ *McGARY, JAMES W., and FRED W. SIELING. ena of Labrador and Maine, 'with a 1953. Chemical and physical data, view of the recent fauna Chincoteague Bay area, July 1943 to of Labrador. Mem. Boston Soc .• 'at. June 1953. Institute, Hist. 1: 210-303. Johns Hopkins Univ., Data Rep. 15, RAPPOR T, MAURICE M. Ref. 53-10, 49 pp. [Mimeo.] 1961. The Cl, P -unsaturated e the r (plas­ MEDCOF, J. C. malogen) content of the tissues of 1958. Stock-taking of molluscan shellfish several mollusks . [Abstract.] BlOl. resources and prospects for improve­ Bull. (Woods Hole) 121: 376-377. ment. Fish. Res. Bd. Can., Prog. Rep. REHBUN, LIONEL 1. Atl. Coast Sta. 71: 21-25. 1956a. Electron microscopy of basophi~lc MEDCOF, J. C., A. H. LEIM, ALFREDA B. structures of some in v e r t e bra t e NEEDLER, A. W . NEEDLER, J. GIB­ oocytes. 1. Periodic lamellae and the BARD, and J. NAUBERT. nuclear envelope. J. Biophys. Biochem. 1947. Paralytic shellfish pois oning on the Cytol. 2: 93-103. Canadian Atlantic coast. Fish. Res. Bd. 1956b. Electron microscopy of basophilic Can., Bull. 75, 32 pp. structures of some invertebra e

11 REHBUN, LIONEL I._-Con. SCLUFER, EVELYN. oocytes. II . Fine structure of the yolk 1951. Respiration of Spisula egg s. nuclei. J. Biophys. Biochem. Cytol. [Abstract.] BioI. Bull. (W oods Hole) 2: 159-171. 101: 228-229. 1957. Vital staining of eggs of Spisu1a 1954. Respiration of the Spisula egg. sol i dis s i m a by methylene blue. Unpublished Ph.D. Thesis, Univ. [Abstract.] Biol. Bull. (Woods Hole) Pennsylvania, 6 7 pp. 113: 353. SCOTT, GEORGE T., andHUGHR.HAYWARD. 1958. Behaviour of metachromatic granules 1950. The influence of the glycolytic in­ during cleavage in Spisu1a. [Abstract.] hibitor iodoacetic acid on aging and Biol. Bull. (Woods Hole) 115: 325. on the potas sium and sodium content 1959. Studies of early cleavage in the surf of t he egg cells of Mactra solidissima. clam, Spisula sol i dis s i m a, using [Abstract. ] BioI. Bull. (Woods Hole) methylene blue and toluidine blue as 99: 363 - 364. vital stains. Biol. Bull. (Woods Hole) SIELING, FRED W . 117: 518-545. 1955. Surf clam industry at Ocean City 1961a. Application of freeze- substitution to expands. Maryland Tidewater News electron microscope studies of inverte­ 11(10): I, 2, 4. brate oocytes. J. Biophys. Biochem. 1960. Mass mortality of the , Cytol. 9: 785-798. Asterias forbesi, on the Atlantic coast 1961b. Electron microscope studies of of Maryland. Chesapeake Sci . 1: 73-74. frozen- sub s tit ute din v e r t e bra t e STEPHENS, GROVER C . , and ROBERTA. oocytes. [Abstract.] Anat. Rec. 139: SCHINSKE. 266. 1961. Uptake of amino acids by marine 1961c. Endoplasmic reticulum in the asters invertebrates. Limnoi. Oceanogr.6: of invertebrate eggs. [Abstract.] BioI. 175-181. Bull. (Woods Hole) 121: 367-368. STICKNEY, ALDEN P . 1963. Induced amoeboid movement in eggs 1963. Histology of the reproductive system of the surf clam Spisu1a solidissima. of the soft-shell clam (Mya arenaria) . Exp. Cell Res. 29: 593-602. BioI. Bull. (Woods Hole) 125: 344-351. *REINTJES, JOHN W., and CHARLES M. STRITTMATTER, CORNELIUS F., and ROITHMAYR. PHILIPP STRITTMATTER. 1960. Survey of the ocean fisheries off 1960. Separation and isolation of Spisu1a Delaware Bay. Supplement rep 0 r t, embryo cells . [Abstract.] BioI. Bull. 1954-57. U.S. Fish Wildi. Serv., Spec. (Woods Hole) 119: 340-341. Sci. Rep. Fish. 347, iv + 18 pp. STRITTMATTER, CORNELIUS F., PHILIPP RUGGIERO, MICHAEL. STRITTMATTER, and CAROLYN 1961. Equipment note no. 9 .... The surf­ BURDICK. clam fishery of New Jersey. Commer. 1960. Electron transport in eggs, develop­ Fish. Rev. 23(8): 11-13. ing embryos and adult tissues ofSpis ula SCHECHTER, VICTOR. solidissima. [Abstract.] BioI. Bull. 1937. Calcium and magnesium in relation (Woods Hole) 119: 341. to longevity of Mactra, Nereis, and S T RI T T MA T T E R, PHILIPP, and COR­ Hydroides egg cells . [Abstract.] BioI. NELIUS F. STRITTMATTER. Bull. (Woods Hole) 73: 392. 1961. Electron transport in eggs, develop­ 1941a. Experimental studies upon the egg ing embryos and adult tissues ofSpisu1a cells of the clam, Mactra solidissima, solidissima. J. Cell. Compar. Physiol. with special reference to longevity. 57: 8 7- 93. J. Exp. Zool. 86: 461-478. SULLIVAN, CHARLOTTE M. 1941b. Further studies on Mactra egg cells . 1948. Bivalve larvae of Malpeque Ba y, [Abstract.] Biol. Bull. (Woods Hole) P. E. 1. Fish. Res. Bd. Can., Bull. 81: 303. 77, 36 pp. 1956. The effect of water upon gametes, SWIFT, HEWSON. upon maturation and upon fertilization 1956. The fine structure of annulate lamel- and cleavage. Exp. Cell Res . 10: 619- 1ae. J. Biophy s. Biochem. Cytol. 2: 630. 415-419. 1958. Water relations of the Spisula' egg. THIELE, JOHANNES. [Abstract.] BioI. Bull. (Woods Hole) 1935. Handbuch der systematischen 115: 362-363 . Weichtierkunde, vol. 2. Gustav SCHUEL, HERBERT, and CHARLES B. METZ. Fischer, Stuttgart. Reprinted by A. 1960. Inhibition of fertilization of Asterias, Asher & Co., Amsterdam, 1963. 375 pp. Spisula and Chaetopterus eggs by THOMAS, LYELL JAY, JR. Arbacia dermal secretion. [Abstract.] 1951. A blood anticoagulant from surf Biol. Bull. (Woods Hole) 119: 297- clams. [Abstract.] BioI. Bull. (Woods 298. Hole) 101: 230-231.

12 THOMAS, LYELL JAY. JR.- - Con. .S. F ISH 1953. A heparin-lik blood an lcoa ulan 19'5 c . from surf clam . Unp blished Ph. D. Th sis, Univ. P nn ylvama, 42 pp. 1954. The localization of h parin-lik blood 195 anticoagulant ubs anc'" in th i ue of Spisula solidissima. BioI. Bull. (Woods Hoi) lOb: 129-138. TOmAS, JU LIA' 1. 1952. Som observations on th ruc ure of nerve in the clam. Mac ra sohdis­ sima. J. Cell. Compar;-phYsioI. 39": 161-164. TURNER, HAHR Y J., JR. 1949. Report on investigations of m thod of improving the shellfish resourc s of Massachusetts. Mass. Dep. Cons .. Div. Mar. }ish., 22 pp. (Contnb. SID, Woods Hole Oceanographic Ins itute ColI. Reprints, J 950.) ~' E 1953. A review of the biology of some 1 commercial molluscs of the ast coast of ' orth America. Sixth Rep. Invest. Shellfisheries Mass. 1ass. Dep. Nat. Res ., Div. Mar. Fish., pp. 39-74. TURNER, HARRY J., JR., and DAVID L. BELDING. It C .. 1957. The tidal migrations of Donax B 01. variabilis Say. Limnol. Oceanog;:--r: 120-124. [U.S.] FISH AND WILDLIFE SERVICE. 1949. Surf-clam fishery. Commer. Fish. Rev. 11(4): 33. 1955. Surf-clam industry at Ocean City expands. Commer. Fish Rev. 17(S): 31- 32. U .S . FISH AND WILDLIFE SERVICE. 1958a. Commercial fishery landings at Ocean City, 1957. Commer. Fish. Rev. 20(9): '14-45. 1958b. Hard-shell and surf clam explora- tion by 1/ V "Sunapee". Commer. Fish. Rev. 20(9): 45-4 .

3 Table 1.--Surf clam catch for the Atlantic coast of the United States, in pounds of meats, and value in dollars (Absence of catch data may mean lack of either catch or data)!!

Average pr~ce Massachusetts Rhode Island Nel< York Nel< Jersey Delal

----

1/ Data for 1901-61 from personal communication, Walter R. Welch, BCF Biological Laboratory, Boothbay Harbor, Maine. '1/ No survey in New Jersey, Delaware, or Maryland in 1946. 3/ Ocean quahogs and surf clams combined and listed as surf clams in Rhode Island in 1954. 4/ Ocean quahogs and surf clams combined and listed as ocean quahogs in Rhode Island in 1455. 5/ Less than 500 pounds or 500 dollars. !.J ~ 6/ 0 C/l Data from U. S. Fish and Wildlife Service Statistical Digest Nos . 57, 58, and 59. =11= '!/ Data from Current Fishery Statistics publications for the respective states ...... C1' lJl 00